THE A MER ICA N J O U R N A L

ED ITO R IAL B OAR D

CH A RL E S R . B A R D EE N G . C A R L H U BE R J P L A Y F A I R MCM U R R I C H
U n i versi t y of Wi sc on sm U n i v ersi t y of Mi c hi g a n U ni v ersx t y of T oro nt o

H E NRY H D O N A L D S O N
. G EO RG E S . H U NTINGTO N GEO RG E A PI E RS O L
T ha W‘etar I nst i t u t e Co l um i a b U iv
n ersi t y U iv n er s i t y of P e n n sy l v a n i a

SIM ON H . GA GE H EN R Y MCE . K N O W ER , S ECRET A RY

Co rn ell U ni vers i t y U iv
n e rs i t y of C i nc i nn at i

MAR CH —
S EP TEM BER , 1 9 2 0

THE W IS TA R IN S T IT UT E O F A N A T O M Y A N D B IO L O G Y
,

P HI L A D E LP H I A PA , .
 CO NTEN TS

NO . 1 . M AR CH

H . E . J OR D A N . S t di u es o n st r ip d e m u sc l e s t ru c t u r e . VI . The co m p a r at iv e h i s t o lo gy
of t he l eg an d w n i g mu sc le o f t h e w as p , w thi s p i ec a l r e e r en c e f t o the ph en o m en o n
o f st r e r e e rs a l ip v du i g r n c o n t r ac t o n i an d t he g en e t c i r e l at o ni b et w een c on t ra e

t i on b an d s an d i n t e r c a l at e d di F t y igh t sc s . or -
e fi g u r es
C H RI S TI A NN A S M I T H . A s t u dy f t h l i p id
o t e o c on en t of t he k id y ne tu b u le . F o u r t ee n

fi g u re s ( tw o pl at e s )

No . 2 . MAY
GE 0 . S . H U N T1 N G T 0 N . A it i q cr u e of t h e t h eo r i es o f p ulm o n ary e v o l u t on i i n t h e m am
m al i a F ift ee n fi g u r es ”
W1
.

E . A B A U M G A RTNE R M ,
. T . N E L S ON A ND 11 . D O CK . D v l pm e e o en t of t h e u t er in e
g l an d s i n m an Sv. e en fi g u r es
E L I OT R . CL A RK AND E L E A N OR L INTON C L A R K . R i
e ac t on s o f c e ll s i n t he t a l i of Am
p h i b i an l ar v ae to in j ec t e d c ro t o n o i l ( asep t i c i n fl am m at i o n ) . Fift ee n fi g u re s

No . 3 . JU L Y

J A ME S W . P A PE Z . H e ar t , m u s c u l at u r e of t he at r a i . E i gh t fi g u r es (f ou r pl at es )
H . E . J OR D A N . F u rt h e r st u di es on r ed b on e -
m a rr ow . I . Exp i m er en t a l . II Cyto
lo gi c, i
w th p i l f
s ec a r e e r en c e t o t he d a t a su gg i g int
es t n r ac e ll u l ar h em o c y t o g e n i c ac

t iv i ty on th p t f th gi
e ar o e an t c e l l s , an d to the s i g n i fi c an c e o f t he s o - c a ll e d m it t i o c

fi g u r es i n t h ese c el l s . O ne pl at e ( t w en t y ~
se v en fi g u res )
CH I K A N O S U KE O G AW A . T h e fi n er r a m i fi c at i o n s o f t h e h u m an lu ng . E i gh t fi g u r es
C H I KA N O S U KE O G AW A . C on t r i b u t i o n s t o t he h i st o l o gy of t he res pi ra t o r y p s a c es o f t he
v e rt e b r at e lu g s T h i t y i gh t
n . r -
e fi g u res .

NO . 4 . S EPT EMBER

A L E X A N D ER S BE G G . . Ab sen c e o f t he v en a c a v i f i
a n er o r in a 1 2 -mm . pig em b ry o ,
as s o

c i at e d i
w t h the d in g ra a e of t he p o rt al s y st e m int o t he c ar di n al s y st e m . Th ree

fi gu r es
H A Y A TO AR AI . O n t he p o st n at a l d v
e el o p m t f t h ov en o e ary ( al b i n o ra t ) ,
w th i es p i
ec a l

f
r e e r en c e t o the n um b er o f ov a . F ou h t r c ar s .

L E ON A U GUSTUS H A USM A N . A mi c ro l o gi l i v tig t i

ca n es a on of t he h a i r s t ru c t u r e o f t he
M o n o t r em a t a . Th r ee te xt fi g u r es an d fou p l t ( i r a es n n et y i ght
-
e fi g u re s )

H o l d
ra m A M E RICA N J O UR N AL o A N AT M Y
r O , vo n. 27 , N 0 . 1
R es umen por a utor Harvey Er n est J ordan
el , ,

E sc u ela Médica de la U n iversidad de V irgi n ia .

E st u dios sobre la estr u ct u ra del m fi sc ul o estriado V I His . .

t ol o g i a c om p arada del m fi scul o de la pata y ala de la av i spa ,

con especial me n ei em del f en é m en o de r ev er si é n de las estr i as

d uran te 1a c ont rac ci én y la rel aci c m gen ética e n tre las b andas
’

de c on t racci én y los discos i n tercalares .

El au tor a n aliza las pr u ebas e x pu estas previame n te en pro y

e n co n tra de la i n t er p ret ac i é n de un a r ev ersi é n de las estrias

d u ran te 1a c on t racci én presen tan do n u evas pr uebas derivadas

,

prin cipalme n te de un est udio del sarc o st il o del m fi sc ulo alar de

la avi spa q ue demu estra 1a existen cia de un a rev er si én gen uin a
,
,

de las es tri as em rel aci én c on la presen cm de un co n stit uyen te

f uertemen te ti n gible del sar c o p l asma E ste co n stit uye n te c ro .

m ét i c o segregado por el disco os c uro (Q ) de la fi b ra eu reposo

,
,
,

se m u eve con tra el t el ofragm a d u ran te l a c on t racci én y c on ,

t ri b uye a la fo rm aci én de la b anda de con t r acci é n E sta fil t i m a .

p u ede fi n alm en t e m o di fi c arse en fin disco i n tercalar L a sub .

s t an ci a f u erteme n t e ti n gible del disco os c uro y la b anda de c o n

t racci é n n o so n idén ticas a un co n stit uye n te ani sot ré p i c o

esp e ci fi c o del sar c o p l asma E1 caracter an i sé t ro p o clarame n te
.

est rat i fi c ad o del sar c o p l asma d u ra n te el repos e es un a fun ci é n

de la ori en t aci én semej an t e de las part i c ul e s sarCO pl é smi cas a lo

l arg o de l i n eas paralelas de resisten cia c uya con di ci é n se satis ,

face a 1i n mej or en los discos os c uros a c ausa de su co n sisten cia

’

, ,

fi uid a relativame n te mas co n siderable

, .

T r a nsl at i o n by J osé F N o m d ez
C
.

a rn e gi e In s t 1 t u t 1 0 n of W as h m g to n
A UT HOR ’
S u s mnw r
'
OF T B IB p u ma 15 5 0 1111)

BY T H E B BL I IOGR P C A HI s mnv mm , J A NU ARY 19

ST U D I E S ON ST R IP E D M U SC LE ST R U CT U R E
VI . COM P ARAT I V E HIST O L O GY O F TH E L E G AN D W I NG MU S CL E
TH E

O F TH E WA SP W ITH SP E C I AL R E F E R E NC E T O T H E P HE NOM E NON

,

OF STRIPE R EVE R S AL D UR I NG C O NTRA C TI O N AN D T O TH E

G E NETI C R E LA T I ON BE TW EEN C O NTRA C TI O N B AN D S AN D INTER
CA L A TE D D I S C S

H . E . J O R DA N
Dep a r tmen t of H i s tol ogy an d Embr yol ogy, Un i versi ty f Vi rgi ni a M edi c al
o

S c hool

FO R T Y E I G H T F I G U RE S
-

C O NTENTS

N m lt
o en c a u re .

R vi w f
e e o

a . L eg m u s c l e
b . Wi n g mu sc l e

Di s c u ss o i n
01 . T l p h gm
e o ra a an d

b . S m
ar c oso es

c . T he a c cesso r y di sc

d . T he i
an so t r o p i ub
c s st a n ce

6 . C o n t ra c t i o n an d t he c o n t ra c t o n i
f . I n t e r c al a t e d di scs

I . INTR O DU C TI O N
This i n vestigatio n ce n ters o n the wi n g m u scle of the wasp .

The reaso n for the selectio n of this partic ular material is the
fact that i t forms the basis of Schaefer s descriptio n of t he ’ 28

morphologic chan ges exhibited by striated mu scl e d ur in g

co n tractio n U p on this descriptio n has bee n b uilt a widely
.

prevale n t hypothesis of mu scle con tractio n the imbibition —

hypothesis .
2 H . E . JOR D AN

This hypothesis takes two disti n ct forms In o n e of these .

forms some relatively less fl ui d co n stit u en t is s upposed to i m

bibe a relatively more fl ui d co n stit ue n t of the i n t rasar co st yli c
sarcoplasm cau sin g i n co n seq u en ce a swelli n g an d shorten i n g
,

of the sarcomeres In the other f orm the swelli n g an d shorten

.
,

i n g of the sarcomeres are s upposed to res u lt from an imbibitio n

of i nt ersar co st yli c fl ui d m u ch i n the mann er i n which a hempe n
,

rope swells an d shorte n s when s u spe n ded i n water .

In an i n termediate f orm this hyp othesis occ u rs i n En g l em ann s

’

co n ceptio n of mu scle co n tractio n En gleman n explai n s co n .

4

tractio n as the res u lt of th e absorptio n of the isotropic material

of the sarcoplasm by the an isotropic co n stit u en t the latter bei n g ,

segregated i n the so called dim disc But the s u ccessful

-
‘ ’
.

operatio n of s u ch a co n tractio n mechani sm req uires s u spen sio n

i n a fl uid whose temperat u r e can be raised s u dde n ly A parallel .

of this m uscle co n tractio n m echan ism is believed to be give n i n

-

a stri n g of catg ut s uspen ded i n water the temperat ure of W t h ,

ca n be raised s u dde n ly by the passage of an electric c u rre n t .

This compromise theory of mu scle co n tractio n accordi n gly

makes an i n t rasar c ost yli e imbibitio n proce ss on the part of the ,

isotropic and an isotropic co n stit u e nts depen den t u po n an ,

imbibition of fl ui d by the sarcostyle as a whole .

S c h aefer fl
discards this thermodyn amic hypothesis of m uscle
' 28

co n tractio n an d explai n s co n tractio n as the res ult of the passage

,

of the relatively more fl uid hyalin e s u bstan ce of the clear disc

‘ ’

i nto pores W ithi n the less fl ui d sarco u s s ubstan ce of the dark

‘ ’ ‘ ’

disc th u s e ffecti n g a horizon tal wide n in g an d a lo n git udin al

,

shorte n i n g of the sarcomere But as will be shown belo w .

,

refere n ce to Schaefer s ill ustratio n of a so called co n tracted

’
-
‘

sarcostyle of the wasp s wi n g m uscle upo n W hich he b ases this

’
,

explan atio n makes it clear that this s upposed i n t rasarc omeri c

,

imbibitio n of a fl uid by a semisolid s ubstan ce deman ds that the

sarcostyle be bathed by a hypoto n ic sol utio n ( not an isoto n ic
medi um as n ormally ) an d this fact removes Schaefer s hypoth
, ,
’

esis to the same category as that of En glemann .

The later chief expo n e n ts of a fra n k imbibitio n hypothesis of

co n tractio n are M c D o u g all an d Meigs The latter claims to
14
.
15
 STRIPE D MU S CL E OF WA SP

be able to add s upport to M c D o ug all s theory by his data from ’

a detailed microscopic st u dy of th e sarcostyle of the wi n g mu scle

of the fl y These i n vestigators advocate a close parallel betwee n
.

the phe n ome n o n of mu scle co n tractio n an d the swelli n g an d c on

seq u e n t shorte n i n g res ultin g from e n dosmosis ; an d between
relaxatio n an d the red u ctio n of swelli n g an d the co n seq u e nt
le n gthe n i n g res ulti n g from exosmosis .

En glema n n claims to be able to demo n strate by mea n s of the

4

micropolariscope that the anisotropic s u bstan ce which he re ,

gards as co n stit uti n g the dim ban d does n o t chan ge its locatio n
‘ ’
,

d urin g co n tractio n Schaefer by mean s of R ollet s gold chlorid

.
,
’
—

tech n ic claims to have proved the same fact Schaefer ao

, .
,

co rdi n g l y i d en t i fi e s the s u bsta n ce which stai n s W ith gold chlorid

,

with the anisotropic material of the sarcoplasm The earlier .

in vestigatio n s of M erkel R ollet an d To urneux on the c on

,
17
,
22
,
30

t rary s eemed to demo n strate a reversal of striae d uri n g c o n

,

tractio n These i n vestigators claim that the an isotropic sub

.

stan ce of the dim ban d of un co n tracted m u scle divides alo n g

‘ ’

the mesophragma an d moves i n opposite directio n s toward the

termi n al telophr agmata of the sarcomere to form the an isotropic
co n tractio n ban ds of co n tracted mu scle Co n tractio n is by them .

co n ceived to be the res ult or at least an accompan imen t of the

, ,

moveme n t of an isotropic materials from mesophragma to telo

p hr a g rn a prod u ci n g th u s a reversal of striatio n s
,
N o attempt .

is made to explai n the f un dame n tal relatio n betwee n this move

me n t of the an isotropic co n stit u e n t of the sarcoplasm an d the
coin cide n t co n tractio n of the sarcostyle R utherford also c o n .
26

el u des for a reversal of striatio n d u ri n g co n tractio n a n d i d en t i fi es ,

the chromatic eleme n t ( dark disc ) of the sarcoplasm with the

‘ ’

an isotropic s ubstan ce .

The phe n ome n o n of co n tractio n as expressed primarily i n the ,

form atio n of a co n tractio n ban d has become i n timately related

,

to the q u estio n of the si g ni fic an ce of the i n tercalated discs of

cardiac mu scle It is n o w clear that n o n e of the earli er hyp o th
'

eses which i n terpreted these discs as i n tercell u lar cem en t li n es ,

ten do n s or developin g sarcomeres are an y lon ger te n able In a

,
.

series of papers I have developed the hypothesis that the i n ter

4 H . E . JOR D AN

c al at ed discs which I am n o w able to demo n strate also i n h uman

,

leg m u scle are esse n tially m o di fi ed irreversible co n tractio n

,
9

ban ds But Schaefer claims that a co n tractio n ban d is an

.

optical ill u sio n This claim is based chi efly u po n his descriptio n
.

of an alleged co ntracted sarcostyle of wasp s wi n g mu scle treated ’

with R ollet s gold chlorid tech n ic It became n ecessary there

’
-
.
,

fore i n the i nterests of my i nterpretatio n of i n tercalated discs

, ,

to rei n vestigate w asp s wi n g m u scle A comparative st u dy of

x
’
.

the sarcostyle of this m u scle after treatme n t with R ollet s an d ’

other tech n ics has co n vi n ced me that what Schaefer describes

,

as a f un ctio n ally co n tracted sarcostyle is i n fact o ne that has .

become swolle n an d i n co n seq u e n ce shorte n ed ; that is o n e ,

art i fi c i all y co n tracted thr o u gh the actio n chi efl y of the aci du

,

lated hypoto n ic ( formic acid water ) sol utio n employed i n R ol

- -

i et s tech n ic In a f un ctio n ally co n tracted sarcostyle of this

’
.

wi n g mu scle as i n i n sect leg m u scle and other str i ped m u scle

,

gen erally there occ urs a ge nui n e reversal of striatio n s d uri n g

,

co n tractio n as regards a darker colored an d more deeply stai ni n g

co n stit ue n t of the dim disc ‘ ’
.

It may be stated at o n ce that there is n o absol utely defini t e

correspo n de n ce betwee n the distrib utio n of the an isotropic con
st i t u en t s of the sarcoplasm an d the limits of the deeply stai n i n g

s ubstan ce o f the dark discs at the di fi eren t fun ctio n al phases .

A n u n stai n ed fi b er appears striped n o t beca u se the a n isotropic ,

material is segregated i n to defin i t e n arrow strata ( discs ) b ut ,

becau se of a chemically differe n t as demo n strated chi efly by a ,

di ffere n ce of stai ni n g reactio n an d a darker sarcoplasmic c on ,

st i t uen t segregated i n the darker ba n ds This same s ubstan ce ‘ ’

.
,

n ot a sp e ci fi c an isotropic s u bsta n ce stai n s deeply i n basic dyes ,

an d gives to microscopic preparatio n s as compared with livi n g ,

material a similarly tra n sversely ban ded appearan ce In the

,
.

followi n g pages atten tio n will be directed primarily to the b e

havior of this s ubstan ce d urin g co n tractio n an d to its relatio n ,

to the formatio n of in tercalated discs .

 STR IPE D MU S CL E OF WA SP 5

II . M ATERIAL AND M ETH O DS

In additio n to the wi n g an d leg mu scles of the wasp I u sed ,

for comparative st udy similar material from fl y bee an d eyed , ,

elater (A lan s o cul at us) These materials are practically i den t i

.

cal except that the sarcostyle of the wasps wi n g mu scle has a

,
’

slightly smaller diameter than that of the elater an d a slightly ,

greater diameter than that of the fl y The several tec hn ics of .

i n vestigatio n i n cl u ded examin atio n of fresh material i n R in ger s ’

sol ution i n distilled water an d i n a 2 per ce n t sodiu m chl orid

, ,
-

sol ution ; examin atio n i n Toiso n s sol utio n which proved ex ’

,

c ep t i on al l y satisfactory ; exami n atio n of alcohol fixed material -

i n water i n glycerin an d i n hypoto n ic salt sol utio n s ; R ollet s

’
, ,

gold chlorid tech n ic ; fix at i on i n 90 per ce n t alcohol i n stro n g

—
,

Flemmi n g s fl ui d an d 1 0 per ce n t formalin followed by iro n

’
, ,

hematoxylin stain in g with an d w itho ut eosin an d V an G ieson s ’

co u n terstai n .

II I . N OM E N C LATURE
The prese n t lack of un iformity i n the termi n ology employed
for the descriptio n of striped mu scle str u ct ure is co n f usi n g an d
adds an extra eleme n t of di ffi cult y to an i nhere n tly complex
s u bj ect The time seems ripe for i n siste n ce u po n a uniform
.
,

mean i n gful an d precise termi n ology In an e ffort to select

,
.

from the prof usio n of proposed terms the most satisfactory

design atio n s I shall be gu ided by two prin ciples : 1 ) The omis
,

sio n of proper n ames ; 2 ) prefere n ce for the term which expresses

most regardin g str u ct ure an d esse n tial relatio n s To these I .

shall add i n brackets the more widely u sed syn o n yms adv o ,

cati n g their early di scardur e .

S ar cos tyle wo u ld seem to be an acceptable desig n a tio n for the

u n it of m u scle str u ct u re an d from the viewpoi n t of comparative
, ,

histoge n esis preferable to its syn o n ym myofi bml The pr ot op

’

.
,

plasmic s ubstan ce of the sarcostyle may be design ated s arco

p la sm . Withi n the sarcostyle may un der certain con dition s be
discern ed con stit u en t still smaller fi b ri l s R egardless of the q u es .

tio n whether these s ubdivisio n are art i fi c i al separatio n prod u cts

6 H . E . JOR D AN

or ge n u i n e eleme n tary fi b ri l s they are co n ve n ie n tly design ated

,

metafi bri ls However fi t t i n g the term sarcolemma wo uld be for

.

desig n ati n g the membra n o u s e n velope of the sarcostyle this ,

word sho uld probably be retai n ed i n the se n se of referri n g to the

cell membran e e n velopin g the smooth m u scle cell or the myo ,

blast or the defin i t i ve striped m u scle fi b er take n as a whole

,
.

For the e n velope of the sarcostyle I wo uld s u ggest the term ,

sarcos tyli c mem br a n e In macerati n g fl ui d s a n d un der mecha n

.

ical stresses t h e sarcostyle becomes broke n up at defi n i t e levels

i n to reg ular segme n ts the sarcomeres (i n ok omm at a) These are
,
.

bo un ded termi n ally by defin ite membran es the telop hragmata ,

( Kra u se s gro un d membran es D obie s lin es i ntermediate mem

’ ’
, ,

bran es ) The telophragmata are co n tinu ed i n to the i n t ersar

.

c o st yli c fl u id spaces an d serve to bi n d together the adj ace n t

fi b ri l s of a m u scle fi b er E ach sarcomere co n tai n s a media n

.

darkly colored more deeply stai n i n g dark di sc ( dim disc tran s

, , ,

verse disc of Br ii ck e sarco u s eleme n t of B owman ) This dark

,
.

disc is un der certain co n ditio n s bisected by a light medi an di sc

( He n se n s disc
’
) which ,
is i n t u r n bisected by a delicate mem

bran e the mesop hm gma (median membran e of Heide n hai n )

,
.

Betwee n the term i n al telophragmata an d the median dark

disc lie the li ght di scs ( clear discs i n termediate discs of Krau se , ,

hyali n e s ubstan ce of Schaefer ) In i n sect leg m u scle the light .

discs are frequ e n tly bisected or crossed close to the t el op hrag ,

mata by a delicate dark granular disc the a ccessory di sc ( disc

, , , ,

of Merkel an d R ollet ) U n der these circ u mstan ces the portio n s

.
,

of the light discs betwee n the termi n al telophragmata of the

sarcomere an d the accessory discs are design ated the en d di scs
( discs of E n glemann ) .

This co n ditio n of stratificatio n of chemically an d physically

differe n t s u bstan ces withi n the sarcostyles gives to the fi b er ,

whe n Viewed u n der low m agnifi cat i on a striped or ban ded ap ,

p e a r a n c e a light
,
ba n d ‘
alter n ati n g reg u
’
larly with a dark ba n d ‘
.
’

It is obvio us however that we are act u ally deali n g with discs

, ,

( a n d membra n es ) n o t with,
ge n u i n e ba n ds or stripes B u t
‘ ’ ‘
.
’

the desig n atio n striped or striated may co n ti nu e to be em

‘ ’ ‘ ’

ployed rather than the more precise term st rat i fi ed as applyi n g ‘ ’

 STR IPE D M US C LE OF WA SP 7

to the appearan ce rather than the i n timate str u ct u re of the

, ,

mu scle fi b er an d the sarcostyle In referri n g to a disc the term .

,

len gth ( thick n ess ) may be u sed to i n dicate the lo n git u di n al

exte n t the ter m wi dth (breadth ) whe n speaki n g of the exte n t
,

tran sverse to the lo n g axis of the fi b er .

A system of i n dicati n g the several discs a n d membra n es by

letters has n ow bee n i n ge n eral u se si n ce the time of R ollet s ’

earlier papers o n striped mu scle str u ct u re These letters

are the first of the respective G erman n ames design atin g the
discs an d membran es If s u ch a system is u sed at all it see m s
.
,

better to retai n that employed by R ollet The o n ly chan ge .

that might possibly make any claim to e ffecti n g a gai n wo uld be

the s u bstit utio n of the letter T to design ate the telophragma ‘ ’
,

for the letter Z ( for Zwischen scheibe )

‘ ’
M co uld the n co n ti nu e .
‘ ’

to stan d for mesophragma as well as for M i t t elsch ei b e The ‘

.
’

terms telophragma an d mesophragma ( collectively i n O p h rag

mata) proposed by Heide n hai n are so precise an d co n ven ie n t
, ,

that their ge n eral adoptio n seems ass ured The so called con .
-
‘

tractio n ban d fun dame ntally a composite str u ct ure is act u ally
,
’
,

a con tra cti on di sc completed axially i n cardiac mu scle by the

telophragma ; an d the so called i n tercalated disc is ge n erally-
‘ ’

more n early a ban d ; that is a peripheral portio n of an earlier ,

co n tractio n disc The i n tercalated disc is a more or less deeply

.

exte n ded ba n d ; or it may e n circle the e n tire fi b er i n the fo rm o f a

rin g or perforated disc It seems stran ge that the act u al discs
.

are commo n ly referred to as ban ds an d that the o n ly str u ct u re ,

that is n ot really a disc n evertheless co n tin u es to be desig n ated

‘
i n tercalated disc But lo n g co n ti nu ed un iform u sage has too
.
’
-
,

fi r m l y establ i shed these l atter terms to warra n t cha n ge at th i s

time especially i n V i ew of a prevaili n g se n se of u n c ertai n ty
,

regardi n g the complete history an d i n timate compositio n of

these stru ct u res .

S ummarizi n g the foregoi n g s u ggestio n s the parallel design a ,

tio n s of the pri n cipal discs an d membran es i n terms of char ,

act ors an d abbreviatio n s that best seem to co m me n d themselves ,

are as follows ( fi g s 1 2 an d .
, ,
8 H . E . JOR D AN

Telophragma T (or Z)
L ight disc
D ark disc
Median disc
Mesophragma
A ccessory disc
En d disc
Co n tractio n disc ( ban d )
I n tercalated ban d ( disc )
IV . REVIE W OF LITERATURE
For the p urpose of p utti n g i n to sharper relief the ce n tral
poi nts at iss u e it seems desirable here to review the literat ure
,

an d to appraise the ill u stratio n s beari n g directly o n the sig

n i fi can c e of the co n tractio n ban d of f un ctio n ally co n tracted

mu scle an d o n the q u estio n of the similarity of this ban d to the

,

simplest typ e of i ntercalated disc .

The most sa tisfactory fi g ure ill u strati n g the chan ges i n striped
mu scle d uri n g co n tractio n is that by R ollet of a fi x ed an d 23

stai n ed lateral co n tractio n wave i n the leg m u scle of the chry

so m el id beetle Cassida e q u est r i s ( fi g
,
A t the right the fi b er .

is said to be i n exte n sio n ; at the left un der D oyere s hillock i n ,

’
,

the f ully co n tracted co n ditio n Co n tractio n i n volves a move

.

men t of the separatin g halves of the dark discs i n opposite di rec

tio n s an d a f usio n of the approachin g halves of adj ace n t dark
discs abo ut the telophragma This fusio n i n volves also the i n
.

terv en i n g accessory discs The res ult is a deeply stain i n g con

.

tractio n ban d comprisin g o n e telophragma two accessory discs

, , ,

an d two opposite halves of adj ace n t dark discs There has .

occ urred accordin gly a reversal of striatio n s d urin g co n tractio n

, ,

as regards a deeply stain in g s ubstan ce located i n Q and N i n

the exte n ded co n ditio n an d tran sferred to the co n tractio n ban d ,

origi n atin g abo u t the telophragma i n the co ntracted co n ditio n ,

.

R o l l etfl
agrees with the earlier co n cl usio n of Merkel
24
'

that the movi n g s ubstan ce i n co n tractio n is the an isotropic

co n stit ue n t of the sarcostyle R ollet s fi g ures of similar .
’ 23

fi b er s as Viewed un der crossed n icols show all of the deeply

, ,
 STR IPE D MU S CL E OF WA SP 9

S a r colemma

FI G . 1 . I ll u st r a t o n i of fix ed an d h em a t o xyl i n -s t a n ei d sp i m ec en of m u sc l e
i g v
‘

fib e r of C a ss id a e que st ri s , s h o w n at t he le t f a l a t e ra l ‘
co n t r a c t o n i wa e
’
in
c ludin g i gh t n t t n
e
‘
cco d in g t R ll t ) A t t h i gh t t A
ra c 1 o b d (
an s
’
a co r o o e . e r ,
a ,

t h w it
e r h dd d t w
er me
as a t i ll u t
e t hi n cep t i
o sar co f th di t i n
r es o s ra e s co on o e c on o

in t h l x d m u l fib Th i gh t h d b d f R l l t fi g u i d ’
-
e re a e sc e er . e r an or er o o e s re s , a c co r

in g t t h w i t int p t t i n t m i dp h f ti n ti n
’
o e r er s nt
er re a Th e nt o ,
a ase o co ra c o . co ra c o

b n d i n lu d pp i t h l v f t w u
a c es o os i v Q di t h i n t v n in g N di
e a es o o s c cess e -
s cs , e er e - sc s ,

an d t h bi t in g t l ph gm T h
e se c u du in g n t t i n t u ev l
e o ra a . er e o c c rs r co ra c o a r e r e r sa

o f st i at i n
r g d d p l y t in i g u b t n c f t h Q di c n d t h n
o a s re ar s a ee s a n s s a e o e - s s a e co

t ti n b nd
ra c o I l lu t t i n f i m i l fi b V i w d w i t h t h e m i p l i p
a s . s ra o s o s ar e rs e e c ro o ar s c o e,

s h w Q N
o nd Z
,
ni t pi in n tu
,
a ; h n
a w m y in f
s ro t h t R ll t
c g dd a re e ce e a er a o e re ar e

l
a so th c nt e ti n b ndo ni t
ra c p i n d in t p t d t h v l f t i t i n
o a as a so r o c a er re e e re e r sa o s r a o

as th ul t
e r es f ch n g f p it i n f t h
o a ni at pi ub t
e o f t h Q di
os o o e a so r O c s s an c e o e - s cs

o f th l x d fib
e re a e er .
10 H . E . JOR D AN

stai n i n g portio n s alo n g the right border of the fi g ure ( N Z , ,

a n d the separated halves of Q ) as a n isotropic i n physical co n sti

t ut i o n A ccordi n gly he i n terprets the co n tractio n ba n d as

.
,

composed of the an isotropic s ubsta n ce which was segregated

withi n the Q disc i n the relaxed fi b er The q u estio n of the .

distrib utio n of the an isotropic s ubstan ce at di ffere n t f un ctio n al

phases will be f ully disc u ssed below It will su ffi ce here to .

poi n t o ut that R ollet ide n tifies the an isotropic with the deeply
stai n i n g s ubsta n ces of the sarcoplasm It may be stated n o w .
,

i n a n ticipatio n of the s u bseq u e n t f u ller disc u ssio n that the ,

an isotropic co n stit u e n t of the sarcoplasm mai n tai n s n o precise

correspo n de n ce with the deeply stai n i n g portio n s of the
sarcostyle .

Two other poi n ts pertai n i n g to this ill u stratio n m u st be

to u ched : the festoo n i n g of the sarcolemma an d the abse n ce of
sarcomeres i n which the Q disc is un divided A festoo n ed .

sarcolemma is n o t an i n variable coi n cide n ce of co n tractio n It ‘

may be claimed that i n ge n eral the festoo n ed co n ditio n of the

sarcolemma is a fi x at i o n artifact A s to the seco n d poi n t the .
,

right ha n d border of the fi b er here ill u strated is probably n o t

-

i n exte n sio n ( n o r i n relaxatio n ) as is ass u med by R ollet,

At .

A I have added a diagram of two sarcomeres to ill u strate the

co n ditio n of a fib er i n repose The dark disc is n o t divided.

by the prese n ce of an H disc R ollet s fi b er is at a m i dp h ase

.
’

of co n tractio n at the right The occ urre n ce of an H disc is a

.

coi n cide n ce a n d an i n dex of co n tractio n The type of fi b er

, ,
.

show n at A is a very commo n o n e i n microscopic preparatio n s .

If the fi b er of R ollet s fi g ure were act u ally exte n ded at the

’

right the n somewhere betwee n the right an d left ( co n tracted )

,

borders of the ill ustratio n we sho uld expect to fi n d a co n ditio n

like that show n at A A s will be made clear below the ill u s
.
,

t r at i o n A represe n ts a sarcomere i n repose

, ,
S u ch a fi b er .
,

i n the exte n ded co n ditio n wo u ld s i mply have a lo n ger ( thicker )

,

Q disc I i
. n terpret R ollet s fi g u r e as ill ’
u strati n g a fi b e r at
m i dp h ase of co n tractio n at the right i n f ull co n tractio n at the,

left .

In my st u dy with Ba n ks of i n tercalated discs i n heart m u scle ,

11

I based my i n terpretatio n of the si g n i fi can c e o f these discs o n

12 H . E . JOR D AN

processes of a lo n git u di n al splitti n g of sarcostyles an d the

irreg u lar te n sio n s prevaili n g i n ab n ormal hearts Still an other .

b ut rare type of disc i n appearan ce little more tha n a

,

thicke n ed telophragma may be explai n ed as the telophragma

,

co n stit ue n t of a co n tractio n ban d from which a remn a n t of the

deeply stai n i n g s ubsta n ce failed of removal at relaxatio n the ,

whole s ubseq u e n tly becomi n g m o di fi ed by the acc u m ulatio n of

tiss u e fl uid This last type will be f urther disc u ssed below
. .

In order more fi rm ly to establish my hypothesis that i n ter

c al at ed discs represe n t esse n tially m o d i fi e d irreversible co n

tractio n ban ds as above b ri efly o utli n ed I un dertook a thoro u gh

, ,

histologic st u dy of the co n tractio n phe n ome n a i n the leg mu scle -

of the sea spider

—
I co n vi n ced myself i n the st udy of this
.
7
,

material of the esse n tial acc uracy of R ollet s descriptio n of

,
’

the formatio n of the co n tractio n ban d Figure 2 take n from .

,

an earlier paper is here reprod u ced to f urther emphasize my

,

agreeme n t W ith R ollet i n regard to the formatio n of the c on

tractio n ban d an d to ill u strate the details i n o ur disagreeme nt
,

regardi n g the sp e cifi c morphologic characteristics of the relaxed

co n ditio n of the sarcomere Portio n s of fo u r fi b ers ( A to D )
.

are shown arran ged i n horizo n tal series from left to right
,

accordin g to the degree of co n tractio n Fiber A is i n a c o n .

ditio n of repose A rrows Z mark the limits of a si n gle sar

.
, ,

c o m ere
. The dark disc ( Q ) stai n s very deeply thro u gho ut
an d exists as a compact u n divided disc The accessory disc .

( N ) is fai n tly visible o n either side of Z i n the light disc ( J ) .

Fiber B is j u st passi n g i n to an early phase of co n tractio n The .

dark disc has become lo n ger while the sarcomere as a whole

has become shorter Moreover the dark disc is becomin g
.
,

lighter medially the co n stit ue n t m yo fi b rils havi n g become

,

disti n ctly k n obbed an d if possible still more deeply stai n i n g

, , ,

termi n ally This co n ditio n clearly demo n strates a movemen t

.

of the deeply stain i n g s ubstan ce toward Z In fi b er C co ntractio n .

is approximately at m i dp hase The dark disc is no w bisected .

by the seco n darily formed median disc ( H ) The res ulti n g .

halves of Q have f used i n part with the accessory discs en co un t

ered i n their passage toward Z The dark disc i n cl udi n g H .
, ,
 ZN

F iv T
fl

FI G 2 .
(A B C , , ,
and D) o u r s u c c ess e f un c t i o n a l s t a g es o f a l eg -m u sc l c
p id F ib A xd
l

fib e r of the sea - s er, An O p l o da ct yl u s l en t u s . er i s i n t he rela e co n

di t i on , fi b er s B, C , an d Di n su c cess iv ly l t phe a er a ses o f c o n t ra c t 1 o n . Id ti en cal

st a g es , i n t he i i
sa m e se r a t o n , a re m t w it h wh n
e e t he four fib e rs a r e co n s id e re d
in a i z n t l di t i ( b t w n t h t l ph gm t Z ) n d w h fi b D i
hor o a rec on e ee e e o ra a a, ,
a en er s

co n id s d i n v t i l di t i
e re aThi er p n d n b tw n t h d f
ca re c on . s c o r r es o e ce e ee e or er o

s u i n f t h di ff n t l v l f fib D d t h d f i t i n f t h f u
c c e ss o o e e re e e s o er an e or er o se r a o o e o r

fib ers w uld o m t p v thi h iz t l

see i ti n
o rot eDu i g t t i n
s or on a se r a o co r r e c . r n co n ra c o ,

th f
er e o r e , th d p l y t i n i g ub t n f t h Q di
e ee s af th l x d fib
n s(A ) s a ce o e - scs o e re a e er

di v id n d fl w i pp i t di t i f m t h m id l in ( m p h gm a) t w d
es a o s n o os e re c on s ro e e e so ra o ar

th dj n t t l p h gm t ( B) m n wh i l in v l v i g t h N di ( C ) n d u i t i n g
e a ace e o ra a a ,
ea e o n e — scs a n

a b ut t h t l ph g m t (D ) t f m t h
o e e o rat ti n b
a a d ( Cb ) f t h n o or e c on ra c o an s o e co

t t d fib e
ra c e Th ru . h a t u v l f t i ti n
e re o c c rs g d t h d p ly
er e r e re e r sa o s r a o as re ar s e ee

t i n i n g ub t n
s a s f Q du i g t h f m t i
s a ce f th
o nt t i n b nd
r n Th e or a on o e co ra c o a s . e

H di - isc in id n f n t t i
s a co c F i b D i t t h d i n i t m idd l p t i n
e ce o co rac on . er s s re c e s e or o

( di g t J d : A n t R
a c co r n o l 1 0 p 4 93
or an a . ec .
,
vo .
,
.
,

13
14 H . E . JORD AN

is still lo n ger than i n fiber B b ut the sarcomere as a whole is

,

shorter In fi b er D appear s u ccessively later phases i n the for

.

matio n of the co n traction ban d (levels 3 4 5 an d The , , ,

sarcomere has become red uced to less than half the le n gth of
that of the relaxed fi b er The co n tractio n ban d co n sists of the
.

f u sed opposite halves of s u ccessive Q discs an d the two i n volved ,

accessory discs the W hole bisected by a telophragma That the

, .

horizon tal seriatio n as here given is correct is proved by the

fact that t he same phases follow i n the same order i n passin g
from top to bottom of fi b er D which is still relaxed above b ut ,

f ully co n tracted below .

Three additio n al importan t data are co n trib uted by this

ill ustratio n The sarcolemma of the co n tracted fi b er ( D ) is
.

n ot n ormally festoo n ed The seco n d dat um relates to the modi

.

fi c a t i o n s prod uced i n the str u ct u re of a fi b er by stretchi n g A .

fi b er may be stretched at a n y phase of co n tractio n This simple .

fact is of prime importan ce ; it may n ot safely be ign ored i n the

i n terpretatio n of any partic ular fi b er whe n j u dgin g of its fun c
t i o n al co n ditio n o n morphologic gro un ds In fi b er D by reaso n .
,

of the co n ditio n of full co ntractio n at the lower en d the middle ,

portio n which is o n ly at m i dphase of co n tractio n has become

, ,

greatly stretched The e ffect o f this stretchi n g i n fl uences

.

c hi efl y the le n gth of Q Compariso n of the le n gth o f the sarco

.

meres of this portio n of the fi b er with the len gth of the sarcomeres
i n an un stretched fi b er i n the relaxed co n ditio n i n disregard of ,

the special co n dition obtain i n g i n the case of the former fi b er ,

wo uld lead to “the erro n eo u s co n cl u sio n widely held at the ,

prese n t time that the prese n ce of an H disc is an i n dicatio n of

,

stretchi n g s uperimposed u po n a sarcomere i n repose This .

matter will also be f urther disc ussed below The portio n of .

fi b er D here un der co n sideratio n discloses a third importan t

, ,

fact n amely that the appare n t abse n ce of a mesophragma may

, ,

be an ill usio n for the reaso n that its str u ct ure is too delicate t o
,

come W ithi n the limits of microscopic Visio n The lo n git u din al .

te n sio n imposed u po n the middle portio n of this fi b er by the

stretchi ng relaxes the horizo ntal te n sio n un der which the meso
phragma exists i n the resti n g an d especially i n the co n tracted
 STR IPE D MU S CL E OF WA SP 15

co n dition of the fi b er an d so by reaso n of an i n here n t elasticity

, ,

permits the mesophragma to co n tract an d coarse n to the poi n t

of possible observatio n S u ch observatio n is f u rther aided
.

by the special f un ctio n al co n ditio n s obtain in g i n thi s portio n of

the fi b er ; the prese n ce of the H disc at the m i dp hase of c o n
tractio n allows the darker mesophragma to sta n d forth mor e
co n spic u o u sly withi n the clear (H ) portio n of the dark (Q ) disc .

Figure 3 of a fi x ed an d stai n ed mu scle fiber from the lobster s

,
’

an te n n a i n terpreted as relaxed above an d co n tracted below

, ,

accordi n g to D ahlgre n an d Kep n er shows with exceptio n al ,

3

clearn ess exactly the same co n ditio n s with respect of the un di

v i de d character o f the dark disc of the sarcomere i n repose an d ,

the steps i n the formatio n of the con tractio n ban d However .

,

this ill u stratio n prese n ts o n e serio u s di ffi cul t y : while the sarco

mere of the relaxed portio n meas ures i n the fig ure 1 1 mm i n .

le n gth that of the portion i n terpreted as i n co ntractio n meas ures

,

o n ly 3 mm less But the len gth of a co n tracted sarcomere as

. .

compared with its relaxed associate is approximately two thirds -

less Si n ce the ch aracter of the striatio n s of this fi b er i n dicates

.
‘ ’

co n tractio n at the lower en d the n early equ al len gth of the ,

sarcomeres at the two en ds mu st be explai n ed o n the basis ,

s upplied by the data relati n g to the sea spider leg mu scle of a -

,

s uperimposed te n sio n at the lower en d .

Fig ure 4 showi n g a wave of co n traction passin g over a livin g

,

(un fix ed an d un stain ed) leg mu scle fi b er of the beetle ( D ytisc u s

-

margi n alis ) accordin g to Schaefer

,
demo n strates the same ,
28

esse n tial poin ts regardin g the formatio n of the co ntractio n ban d ,

n amely a n origi n by f u sio n alo n g the telophr agma of the opposite

,

halves of s u ccessive dark discs It ill u strates likewise a genuin e .

reversal of striatio n co n trary to the opin ion of Schaefer w ho

, ,

in terprets the specime n as demo n stratin g that the apparen t

reversal is n o t real b ut simply an optical ill usio n Schaefer s
,
.
’

explan atio n of the basis of this optical ill usio n however har ‘
,
’
,

mo nizes better with o ur descriptio n of what occ ur s d urin g

co n tractio n than with his descriptio n of the esse ntial phe n omen o n
of co n tractio n i n wasp s wi n g m uscle with W hich the D ytisc u s
’

-
“
leg mu scle is said to agree H e says The dark ban ds of t he .
,

TH E A M ERIC A N J OU R N A L op A N AT OM Y vo n 27 , NO 1
h
. .
,

Ma rc , 1 920
16 H . E . JOR D AN

“m
H aar an 1) . v

FI G 3 I ll u st ra t o n i of a fix ed and st a 1ne d p im s ec en o f a co n t ra c t n i g m us c l e
f
fib e r ro b
m t h e l o s t e r s a n t en n a , sh o w n a r e
’
i g v e r sa l of st r i a t 1 o n as re g d ar s a

d p ly
ee i
st a n i n g
su b s t a n c e o f t h e Q
‘

sc o f t he -
di r el a xdp e o rt on i ( ab o v e ) of the
fib e r (a c c o r d i g t D hlg
n o a re n a n d K p e ner ) . J, l ig h t d1sc ; d k di ; Z t l
Q ,
ar sc ,
e o

p h gm
ra a ; M m ph gm
,
e so ra a x i
; m , e t e n s o n s o f t e l o h ra p g m t i nt p m u l
a a o er c ea r

pl
sa r c o as m; c on n 1 t en d on fi b r1 l s w h 1 c h a t t a c h t h e m u s c l t t h hyp d m i
e o e o er s .

FI G . 4 I ll i
u s t r a t o n o f c o n t ra c t l o n wa v e on 11 V 1 n g l e g m u sc l e fi b e r o f D y t l s c u
-

m a rg m a h s ( a c co r d m g t o S f
c hae e r ) . Th e
‘
a pp a re n t
’
v
re e r s a l o f s t r ae i n t h e i
fo rm a t 1 o u o f t h e c o n t ra c t i on b a n d s o f t h e m idd le p o rt i o n 18 e xp l i d b y S
a ne f
c h ae e r

as d ue t o a c c u m u la t 1 o n s o f sa r c o pl a sm ,
a pp ea r n i g as d k ar l 1 1i es , wh i ch o b s cu r e
t he c o n t i nu l t y o f t ho fi b r i l s an d by c o n t r a s t c a u se t h e w ho le of the sar c o m e res
b e t w ee n t h em t o a pp ea r l1ght . T he w r l t e r i n t e r p re t s th i s 1 l l u st r a t 1 o n a s b e in g
in i
s tr ct a cco r d Wi t h t h i llu e i
s t ra t o n s o f fix ed an d s t a n e i d fib e rs , fi g u res an d 3;
t h er ef o re , as d om o n s t ra t m g a t ru e re v e r sa l of st n ae , i n t he f m ti
or a on of t he
n t ra c t i o n b and ,
as 1 e g d ar s a d p
ee e r c o lo r e d su b st a n c e of t he d k l
ar ( lSC .
 STRIPE D MU S CL E OF WA SP 17

co n tractio n wave are seen to be really d ue to acc u mulatio n s of

sarcoplasm These acc umu lation s appear as dark lin es which
.

obsc ure the co ntinuity of the fi b ril s an d by co n trast cau se the

,

W hole of the sarcomeres betwee n them to appear light (p .

The acc u mulatio n s of sarcoplasm that Schaefer here post u lates

‘ ’

are i n ter sar co st yl i c acc um ulatio n s at the levels of the telo

—

p h r a g m at a B u t this
. expla n atio n ass u mes a beaded co n ditio n

of the sarcostyle with co n strictio n s at the telophragma levels .

Beadin g however is n ot a n ormal accompan ime n t of co nt r ac

, ,

tio n b ut as will be demo n strated below a fi x at i on or osmotic

, , ,

artifact Moreover it mu st be emphasized that the si n gle

.
,

sarcostyle of wi n g m u scle likewise shows this phe n ome n o n of

stripe reversal d urin g co n tractio n In fact this phe n ome n o n as
.
,

ex hibited by a complete fi b er is simply the sum e ffect of the

stripe reversals i n the co n stit u e n t m yo fi b ri l s The explan atio n .

of the phen ome n o n o n the basis of an acc u m ulatio n of i n ter

sar co st yl i c sarcoplasm m u st accordin gly for a seco n d reaso n
, ,

be discarded as i n adeq u ate .

Still an other explan atio n of this phen omen o n of stripe reversal

as an optical ill usio n might be based o n Schaefer s diagram ’ 27

of the i n t r asar co m eri c chan ges d urin g co n tractio n (fi g This .

diagram p u rports to explai n co n tractio n as the res ult of the

absorptio n of the hyali n e s ubstan ce of the light disc by the
‘ ’

‘
sarco u s s ubsta n ce of the dark disc If this be accepted as
’
.

represe n ti n g act ual co n ditio n s d uri n g co n tractio n the n the ,

appare n t reversal of striatio n co u ld very plau sibly be explai n ed

as the res ult of a relative co n de n satio n of the area abo u t the
telophragma an d a relative rarefactio n of the area abo ut the
mesophragma that is i n the origin al dark disc But the direct
, ,
.

evide n ce poi nts unmistakably to a moveme n t of fl uid i n exactly

the opposite directio n from the on e here ass u med n amely from , ,

the mesophragma agai n st the telophragma We are accordi n gly .

u n able i n View of the microscopic evide n ce to escape from the

, ,

co n cl usio n that an ac tu al reversal of striatio n as regards the ,

dark s ubstan ce of the sarcoplasm occ urs d uri n g co n tractio n

,
.

Fig ure 5 shows a reprod uctio n from Schaefer of En gl em ann s ’

photomicrographs of the same fi x ed fi b er said to be co n tracted

27
,
 18 H . E . JOR D AN

toward the middle as it appears with un crossed (B) an d with ,

crossed (A ) n icols respectively But the ill ustratio n does n ot

,
.

give co n vi n ci n g proof that the b ulged middle portio n is act u ally

co n tracted This area s u ggests rather a distortio n Skepticism
. .

regardi n g a ge nuin e co n tractio n is f urther aro u sed by the fact

FI G 5 . Ph o t o m c ro i g phra s of p o rt o n i of a l e g -m u s c le fib e r of C h ryso m e l a
c oe ru l ea , i
w th
‘
fix e d c o n t ra c t o n i wa v e,
’
a s see n u n d er t he p o lar i z ing m l c r o sco p e .

Fib erA w ph t g ph d w i t h
as o d i l fib B w i t h
o ra e d i l c ro sse n co s , er u n c ro ss e n co s

(f m S h f
ro cf t E gl m ) T h
ae e r , a p h t g ph
er ni t p t d by
e ann . es e o o ra s a re n er re e

Sh f
c d m t t i g th t th i
ae e r as e h
o n s ra g f p iti f th i t pi
n a er e s no c an e o os on o e a n so r o c

su b t f t h Q di
s an c e o d i g t ti
e h - sc t v l f t i ti
ur n c o n ra c on , en ce n o ru e re e r sa o s r a on .

S h f l im t h t t h m f t
c ae er c a s b d ma t t d i
e sa tyle f w p ac c an e e o n s ra e n sa r c o s es o as
’
s

wi g m t d di g t R ll t t h i ( mp fi g Sh f ’
n l t u sc e rea e ac co r n o o e s ec n c co a re . c ae e r

di gl y m k t h
a c co r n w t d
a mpti
es th t th
e un d p ly t i i g b
a r ran e a ssu on a e ee s a n n su

t
s an ce f t h d k di
o i id t i l w i t h p ifi
e ar sc i t p i m t i l f th
s en ca s ec c an so r o c a er a s o e

sar co pl m as .

that the anisotropic ban ds i n the area i nterpreted as co ntracted

are q uite as sharp an d disti n ct as those of the relaxed termin al
regio n s whereas most in vestigators agree that the an isotropy
,

of a con tracted fi b er is relatively feeble However this ill u s .

,
20 H . E . JOR D AN

disc . This phe n ome n o n of st rat i fi cat i o n may readily be demo n

st ra t ed u n der favorable co n ditio n s i n i n sect leg m u scle I have .

also co n vi n ced myself by m i crop o lari s0 0 pi c st u dies that the

a nisotropic strat u m does n ot i n ge n eral chan ge its origi n al more
or less defi ni t e segregatio n approximately withi n the limits of
the dark disc at least i n n o reg ular and un iform mann er d urin g
, ,

co n tractio n But similar st u dies of co n tracted fi b ers show also

.

that the an isotrop y of the fi b er an d sarcostyle i n this co n ditio n

is relatively feeble .

Fig ure 6 take n from Schaefer is of prime importan ce becau se

, ,
28

u po n it are based diagrams ( fig 9 ) which are u sed i n most of the .

c urre n t text books of gross an d m i cro sc0 p i c an atomy to explai n

-

the mechani sm of m u scle co n tractio n This fi gure i n cl udes .

three sarcostyles of the win g m u scle of the wasp treated accordi n g ,

to R ollet s gold chlorid tech n ic Sarcostyle C is said to be

’
-
,

un co n tracted ; B stretched an d A co n tracted It becomes

, , ,
.

n ecessary to recall the steps i n R ollet s tech n ic This tech nic ’

.

i n volves fix at i on i n 90 per ce n t alcohol for twe n ty fo u r ho urs —

,

tran sfere n ce to stro n g glyceri n for several ho urs thoro u gh ,

washi n g i n water immersio n i n a 1 per ce n t gold chlorid sol ution

,
-

for from fi ft een to th i rty mi n utes an d fi n al tran sferen ce to a ,

sol utio n of o n e part of formic acid to three parts of water for

twe n ty fo u r ho urs or lo n ger It is importan t to n ote that this
-
.

tech n ic i n volves the prolo n ged u se of an acid ulated hypoto n ic

sol utio n .

A n ticipati n g i n part what will be more f ully developed below ,

the followi n g stateme n ts seem desirable here regardin g this

fi g ure : In fi b er C Schaefer lays stress u po n an d attaches great ,

importan ce to the small circles i n the upper en d of the fi b er

,
.

These he i n terprets as ope n i n gs i n to pores i n the dark disc H e .

assu mes that these pores are O pe n towards the hyali n e sub
‘ ’ ‘

stan ce closed withi n the dark sarco u s ele m e n t Bu t s u ch

,
’ ‘
.
’

pores cann ot be see n i n fresh sarcostyles exami n ed i n R i n ger s ’

sol utio n ; n o r are they discer n ible i n sarcostyles treated accordi n g

to man y other histologic methods They m u st accordin gly be .

i nterpreted as artifacts res ulti n g from the use of R ollet s tech n ic ’

.

Moreover gran ti n g that there act ually occ u r capillary pores

, ,
 STR IPE D MU S CL E OF WA SP 21

there is absol utely n o eviden ce to i n dicate that they are o n ly

ope n at o n e en d an d closed at the en d within the dark disc .

A lso the prese n ce of a bisecti n g n arrow media n disc i n the dark

,

disc shows that the sarcostyle is n o t act u ally un co n tracted b ut ,

that it is i n an early phase of co n tractio n .

Sarcostyle B is said to be stretched This mu ch cann ot be .

den ied Bu t we are f urther led to in fer that the occ urre n ce of
.

the bisecti n g median disc is a coi n cide n ce of stretchi n g It may .

be gran ted that extreme te n sio n may cau se a divisio n of the

dark disc as here illu strated especially at the very begi n ni n g of
,

co n tractio n before the i n itial median disc is clearly discer n ible .

But moderate te n sio n e ffects n o s u ch res ult at this f un ctio n al

phase Te n sio n exerted upo n the restin g sarcostyle res u lts
.

simply i n a le n gthe n i n g of the dark disc It seems more i n .

accord with the histologic data to i n terpret sarcostyle B as

o n e at an early phase of co n tractio n seco n darily m o difi ed by
,

stretchin g .

Sarcostyle C is described as i n co n tractio n It is i nterpreted .

as demo n strati n g that n o reversal of striatio n occ u rs d urin g

co n traction The deeply stai n i n g dark disc which Schaefer
.
,

i den t i fi es with the an isotropic s u bstan ce has n o t moved from its

,

origi n al positio n i n the un co n tracted sarcostyle o n either side

of the mesophragma But this sarcostyle is n ot act u ally fun c
.

t i o u all y co n tracted It is simply swolle n an d i n co n sequ en ce

.

shorte n ed that is art ifi ci ally co n tracted thro u gh the e n dosmotic

, , ,

actio n of the hypoto n ic formic acid water sol utio n The fun c
- —
.

t i o n ally co n tracted sarcostyle has a totally di ffere n t appearan ce ;

here a tr u e reversal of striatio n occ urs (fi g s 8 3 3 an d .
, ,

Moreover the un m o di fi ed co n tracted sarcostyle is n o t beaded

, ,

b u t mai n tain s a sharp lateral co n to u r Sarcostyle A is accord

.

i n g ly an artifact an d can n ot properly serve as a basis for the

,

co n str u ctio n of diagrams to ill u strate the mechanism of mu scle

co n tractio n If s u ch a sarcostyle is placed i n a 2 per ce n t sodium
.

chlorid sol u tio n for several ho u rs it ret ur n s to a co n ditio n very

,

m u ch like that of sarcostyle C In other words sarcostyle A is

.
,

simply sarcostyle C swollen thro ugh the actio n of a hypoto n ic

sol utio n Sarcostyles like that ill ustrated i n C are o n ly fo un d i n
.
22 H . E . JOR D AN

cl um ps T he close co nn ectio n of adj ace n t sarcostyles by the

.

telophragmata appare ntly sec ures protectio n to a co n siderable

degree agai n st the swelli n g action of the hypoto n ic sol utio n .

Sarcostyles like A are o n ly fo un d isolated an d are i n additio n ,

u s u ally somewhat fl att en ed o ut un der the cover slip -

.

Fi g . 7 (A B C
, , ,
and D) I ll i
u s t ra t o n o f f ou r sa r co st yl es o f t he w in g m u sc l e
ofH yd ro p h i lu s p i ce u s ( f ro m c h a e S f er, a fter R an v 1 er ) . m , t elo h ra p g m a ; h,
di
me a n d 1 s o ; 3 , a r d k di sc ( sa rc o u s
‘
be , l ig h t di sc (
‘

h y l in
a e su b

R anv 1 e r an d S f
c h ae e r i n t e r p r es t A as
‘
m o st con t ra ct e d ,
’
and D as
‘
x
m o st e t e n d e d Th e fib r i l s a re
.
’
co m p a ra b le t o t he sa r co st yl es of t he w in g
p
m u sc l e o f w a s , b e e , e la t e r , an d fl y . As suc h t he w r 1 t e r i n t er p ret s C a s a sa r c o

x
s t y l e i n t h e re l a e d c o n d i t i o n ; B a s a sa r c o st yl e i n wh i ch t he d k di ar sc h as b ec o m e
sl igh t l y sw o ll en in c o n se q uen c e o f t he sl i ght a c t oni of a h yp t i o on c so l u t o n i ; A
as a sa r co s t y le wh i ch ha s b ec ome sw o ll e n an d b ea d e d ,
an d in c o n se q u en c e

s h o r t en e d (t hat i s, a rt ifi ma ll y c o n t ra c t e d) in c o n se q u en c e o f the p r o lo n gd e act on i

of a h yp i
o t o n c s o l u t o n , an d sa r co sti y le D a s one i n m 1 d p h a se o f c o n t ra c t o n u i p on

wh i ch has p e rh a p b s een su p p
er o se d a s t r et c h e d c on d 1t 1on .

Schaefer cites also R anvier s ill ustratio n s of the wi n g m u scle

27 ’

sarcostyle of the water beetle H ydrO phi l us pi c eu s (fi g in ,

.

s upport of his i nterpretatio n of the wasp s wi n g mu scle sarcostyle ’

-
.

R a nvier likewise co n siders the shorte n ed beaded fi b er ( A ) as

co n tracted R efere n ce to R an vier s origi n al article o nly gives
.
’

the i n formatio n that he st u died these sarcostyles i n the body

fl uid of the i n sect i n white of egg i n pi cro carm i n at e of ammo nia
, ,

a n d i n a 2 per ce n t osmic acid sol utio n H e does n o t specify -

.

the partic ular fl uid employed i n the case of the m u scle from
which the ill ustratio n was drawn A t least two of these sol utio n s .

are hypoto nic to the livi n g sarcoplasm A ccordi n g to o ur i n ter .

 STRIPE D M US C LE OF WA SP 23

sarcostyle C is i n a co n ditio n of repose sarcostyle D

p ret at i o n , ,

is stretched at an early phase of co ntractio n B is a relaxed ,

sarcostyle slightly altered by e n dosmosis sarcostyle A is swolle n ,

an d beaded thro u gh the actio n of a hypoto n ic sol utio n .

In s u pport of his claim that the striae are n o t reversed d uri n g

co n tractio n Schaefer cites also the experime n ts of M acall um
, ,
12

design ed to determi n e the distrib utio n of the potassi um salts i n

the sarcoplasm d urin g the several fun ctio n al stages Schaefer .

selects certain of Macallum s fi g ures of the win g mu scle sarco

’
-

styles of D yt isc u s an d i n terprets these as demo n stratin g that

,

also the potassi um salts do n o t shift their positio n d uri n g c on

traction from their locatio n withi n the dark disc i n the relaxed
,

sarcostyle Schaefer wo uld th u s seem to ide ntify the dark disc

.

n o t o n ly with the a n isotropic strat u m of E n gleman n b ut also ,

with the potassi um co n tai n i n g strat um of Macall um An

—
.

exami n ation of these selected fi g ures from Macall um s paper ’

.
“
(fi g 2 8 9 of Schaefer ) reveals the fact however that the so called
, ,
-

co n tracted sarcostyles are act u ally o nly fi b ril s which have

become art i fi ci ally beaded thro u gh osmotic mo di fi cat i on These .

ill u stratio n s reveal o n ly that the distrib u tio n of the potassi um

salts withi n the sarcoplasm is ro u ghly coexte n sive with the dark
disc b ut give n o i n formatio n regardi n g this distrib utio n d uri n g
,

co n tractio n R efere n ce to Macall um s origin al work (p 1 1 8 )

.
’
.

reveals moreover that the distrib utio n of potassi um salts i n

, ,

un co n tracted a n d co n tracted m u scles is n o t a very precise matter .

In the un co n tracted m u scle the potassi u m 1 s said to be ge n erally

limited to the dim ban d b ut also the potassium reactio n is most
,

marked i n those zon es of the dim ban ds immediately adj ace n t

to the light ban ds The latter co n ditio n i n my opi n io n is
.
, ,

comparable with an early stage of co n tractio n In the c on .

tracted sarcostyle M acall um fo un d the most marked reactio n for

potassi um i n the ce ntral regio n s of the dark disc However .
,

whe n the pe n etratio n of the reage n t ( the hex an i t ri t e of cobalt

an d sodi u m ) wa s delayed the stain i n g reactio n revealed the
,

potassiu m sometimes i n the light discs an d sometimes alo n g the

level of separatio n betwee n the light an d dark discs .
24 H . E . JOR D AN

M e n te n ,
her exte n sio n of the experime n ts of Macallum
16
in ,

has show n that the chlorides the phosphates an d the potassi u m

, ,

salts have an an alogo u s distrib utio n A n d her beautiful

‘
.
’

colored ill ustratio n s show clearly that these salts chan ge their
positio n d uri n g co n tractio n an d that this chan ge of positio n
,

correspo n ds closely with the tran sfer of the deeply stain i n g

s u bstan ce from the dark disc of the relaxed fi b er to the c on
tractio n ban d of the co n tracted fi b er A ccordi n gly there occ urs .
,

d uri n g co n tracti on a reversal of striae as regards these salts as

well as regards a possibly sp eci fi c s ubstan ce of the dark disc In .

f ull co n tractio n these salts are segregated practically within the

limits of the co n tractio n ban d (M en t en s fi g s 1 7 1 8 2 2 2 6 3 1 ’
.
, , , , ,

3 6 an d
,
The co n cl u sio n is s u ggested that the deeper color
an d the deeper stai n i n g reactio n of the Q disc an d of the co n

tractio n ban d are act u ally due at least i n part to the segregation
, ,

of these salts within these levels at the fun ctio n al stages of

relaxatio n an d co n tractio n .

In fi g ure 8 is reprod u ced a photomicrograph from Meig s

’
,

paper of a livin g co n tracted sarcostyle of the win g mu scle of

,
15

the fl y mo un ted i n a medi u m of equ al parts of white of egg an d

,

a 2 per ce n t sodi um chlorid sol utio n The co n tractio n ban ds

—
.

are co n spic u o u s The sarcomeres have lost approximately half

.

of their le n gth an d attai n ed approximately do uble their diameter ,

as compared with the relaxed fiber A t a the co ntractio n ban ds .

appear si n gle at I) do uble Meigs i n terprets the do uble c on

,
.

ditio n as an optical i llu si on due to the obliq u e orien tatio n of the

‘

sarcostyle at However the co n tractio n ban d is by Virt u e of

,

its origi n essen tially a do uble str u ct u re comprisin g pri n cipally ,

t w o halves of adj ace n t dark discs It seems to me more probable

.

that these appare n tly do u ble con t ra ct i on b an ds i n this ill us w

t r at i o n sho uld be i n terpreted as i n completely f u sed co n tractio n

ban ds In spite of t h e sharp lateral co n to ur of this co ntracted
.

sarcostyle clearly shown i n the fi g ure Meigs n evertheless i nter

, ,

pr et s s u ch fibrils as slightly b ea ded w i t h co n strictio n s at the , ,

levels of the telophragmata ( co n tractio n ban ds ) ; an ass umptio n

requ ired by the imbibitio n hypothesis of co ntractio n b ut i n ,

co n trave n tio n of microscopic data .

 STR IPE D MU S CL E OF WA SP 25

Fig ure gives Schaefer s diagrams design ed to

9 ( A an d B)
’ 27

explain mu scle co ntractio n based u po n his i n terpretatio n of ,

certain win g mu scle sarcostyles of the wasp These diagrams

-
.

persist i n some of the leadin g text books of an atomy an d physi -

ology They are o n ly applicable to a mechan ical explan atio n

.

of co n tractio n ; an d even as s u ch misin terpret the m i cro s0 0 p i c

Fi g . 8 Ph ot o g ph
ra (by u l t ra vi o l et l1 gb t ) of a c o n t ra c t e d sa r co st yl fe ro m
fiy
’
s w i g n m u sc l e t ea se d ou t f resh in a x
m i t u re o f e u a l q p a rt s o f wh t e i of egg

an d a 2 per c en t so di u m -
ch lo r id so l u t 1 o n , m a g n i fi e d 1 3 0 0 di am e t e r s ( a c c o r din g
to Me i gs) . T he di a m et er of t h1s co n t ra c t e d sa r c o s t yl e IS a pp x i m ro at el y t h r ee
i
t m es t h at of t he re l a xd
e sa r co st y le ,
an d t h e l en t h g of t he c o n t ra c t e d sa r c o

m er e s i s a pp x i m
ro a t el y f
on e - o u r t h t ha t of t he re l a xd e sa r c o m e r es . The d k ar

( con t ra c t i on ) b a n d a t b a pp ea r s d o u b l e a t a s in g le M e i g s in t e r p ret s t h e d ou b l e
,
.

a pp ear an c e o f t h e b a n d s ( a t b ) a s a n o p t i ca l eff e c t d u e t o t h e o b l i qu e p o s i t i o n o f

t h e fib e r Th e w r1 t e r i n c l i n e s t o i n t e r p r et t h e d o u b l e b an d s i n t e rm s o f t h e i r
.

d o u b l e o r i g i n a n d an i n c o m p let e f u s i on o f p a i re d c o n st i t u en t s N
.

data D iagram A is s upposed to represen t two sarcomeres of a

.

sarcostyle i n con ditio n of relaxatio n The dark disc ( sarco u s .

‘

elemen t ) is con ceived to be do uble con tain in g medially a bisect

’
,

i n g median ( H ) disc a n d is described as porifero u s T he ‘ ’

.
,
‘
pores are said to be o n ly open toward the light disc ( hyalin e
’ ‘

s u bstan ce ) D urin g co n traction the alleged more fl u id hyalin e

’
,
‘
,
’

s ubstan ce of the light disc is ass umed to flow i n to the pores of ‘ ’

26 H . E . JOR D AN

the dark disc an d i n co n seque n ce ca u se it to decrease i n le n gth

,

a n d i n crease i n width ( diameter ) prod uci n g th u s co n tractio n of

,

the fi b ers by reason of the beadin g and shorte n in g of the com

po n e n t sarcostyles represen ted i n diagram B
,
.

I have added diagram C to complete the series of chan ges i n

the sarcostyle d uri n g co n tractio n i n accord with the data to be
,

prese n ted i n the descriptive portio n of this paper In order to .

u se diagrams A an d B i n this series w e m u st disregard the ex ag

gerated le n gth of the sarcomere i n A My in terpretatio n of .

these diagrams is then as follows : Sarcostyle B is i n the relaxed

co n ditio n ; sarcostyle A i n an early stage of co ntraction as i n di ,

c at e d by the prese n ce of a media n disc a n d sarcostyle C is i n

,

the f ully co n tracted co n ditio n The co n tractio n ban d

.

of sarcostyle C in cl udes fu sed opposite halves of adj ace n t Q

discs bisected by the telophragma
,
.

The commo n ass umptio n that the s u bstan ce of the light disc
is relatively more fl ui d than the s u bstan ce of the dark disc is
also directly con trary to W hat the m i cro sc0 p i c data seem to in dicate .

The prin cipal tran sfer of s u bstan ces d urin g co n tractio n is n o t

from the telophragma to the mesophragma as is req u ired i n ,

the above diagram accordin g to Schaefer b ut i n the opposite ,

'

direction The distortio n e ffects of mechan ical an d osmotic

.

factors show their respective i n itial m o di fi cat i o n s fi r st i n the

dark discs as is to be expected if this portio n is relatively more
,

fl uid than the light disc A relaxed sarcostyle placed un der

.

te n sio n respo n ds fi r st by a le n gthe n i n g of the dark disc U n der .

the dehydrati n g effect of fi x at i o n with the higher grades of

alcohol the dark discs respo n d by a shorte n i n g ( co n de n satio n )
,

relatively far i n excess of that e ffected i n the light disc More .

over i n stai n ed preparatio n s it can readily be see n that a dark

,

stain i n g s ubstan ce of the Q disc act u ally moves toward the

telophragma d uri n g co ntractio n as i n dicated by a cleari n g of
,

the median portio n of Q an d the k n obbed character of the

termi n als of the Q portio n s of the sarcostyles .

The foregoi n g disc u ssio n sho uld have made it clear that the
prevailin g ideas of the morphologic chan ges s uffered by striped
m u scle d uri n g co ntractio n as based largely o n the ill ustratio n s
,
28 H . E . JOR D AN

co n tractio n as does i n sect leg m u scle E vide n ce will be pre .

se n ted for the co n cl usio n s tha t all varieties of striped mu scle

behave d uri n g co n tractio n i n an esse ntially ide n tical mann er ;
that the co ntractio n ban d of a co n tractin g fi b er is a ge nuin e
str u ct u ral e n tity n o t an optical ill u sio n represe n ti n g a n act u al
, ,

reversal of striae with respect of the deeply stai n i n g co n stit u e n t

of the st rat i fi ed sarcoplasm ; an d that this co n tractio n ban d
( disc ) is the primordi um of the several types of i n tercalated discs
of cardiac an d skeletal m u scle I n cide n tally m u st be co n sidered.

also the n at ure an d si g ni fi c an c e of the sarcomeres of the i n o ,

p h ra g m at a ( telophragma a n d mesophragma ) of the accessory ,

disc ( oi Merkel an d R ollet ) an d of the distrib u tio n of the alleged

,

an isotropic materials .

V . DES C RIPTI O N
a . L eg mu scle

The co n ditio n of the relaxed mu scle fi b er is ill u strated i n

fi g ur e 1 4
. This fi b er was fix ed i n 9 5 per ce n t alcohol an d stai n ed
with iro n hematoxyli n The un stai n ed fi b er has an esse ntially
—
.

ide ntical appearan ce The deeply stain i n g portio n s of the

.

stai n ed fib er simply appear more fai n t i n un stai n ed specime n s .

The fi b er is slightly distorted below The sarcolemma o n the .

,

left is slightly festoo n ed d u e t o the un eq u al co ntracti n g actio n

, ,

of the fi x i n g fl uid upo n the m u scle fi b ril s an d the sarcolemma .

Si n ce the fi bril s shorte n more than the sarcolemma un der this

i n fl uen c e a n d si n ce the sarcolemma is i n timately co n n ected with
,

the telophragmata the sarcolemma accommodates itself to the

,

shorte n ed co n ditio n of the fi b ri l s by separati n g from the fi b er

i n the form of arcades Q an d J are of abo ut eq u al thick n ess
. .

The telophragma appears relatively rob u st somewhat gran ular ,

a n d deeply stai n i n g The gran ular appeara n ce of the telo

.

phragma is d ue to swelli n gs at the poi n ts where the m yo fi b ril s

are attached to i t A n accessory ( N ) disc is very co n spic u o us
. .

It is formed by the horizo n tal align me n t of m o di fi ed mi nute areas

of the m yo fi b ri ls i n the Vici n ity of the telophragmata It has .

appare n tly m u ch the same chemical co n stit utio n as the Q discs .

 STR IPE D M US C LE OF WA SP 29

F ig u re 1 4 sho uld be co n sidered i n co n n ectio n with fi gure 1 8 .

The latter was fi x ed W ith Flemmin g s fl ui d an d lightly stai n ed ’

with iro n hematoxyli n The m yofib ril s are discer n ible b ut the
—
.
,

cross striatio n s do n o t appear O n ly the telophragmata are .

co n spic u o us However the fi x at i o n preserved the sarcosomes

.
,

an d they are see n to occ u r both i n the dark discs ( as Q gran u les ) -

an d i n the light discs ( as J gra n u les ) In alcohol fix ed tiss ue the —

—
.

granu les are dissolved an d so do n o t appear i n stai n ed sectio n s

(fi g. Their sol utio n i n alcohol together with certai n micro ,

chemical reactio n s i n dicate that their chemical co n stit u tio n is

,

at least largely lipoid The spherical J gran u les are smaller than
.
-

the oval Q gra nu les The differe n ce i n shape may be du e to

—
.

lateral press u re exerted by the m yo fib r i l s o n the larger origi n ally

spherical Q gran ules The simplest i n terpretatio n of the ge n etic
—
.

relatio n an d the segregatio n of these two varieties of sarcosomes

is that the larger develop from the smaller the latter origi n ati n g ,

alo n g the telophragmata probably by reaso n of the fact that the

,

telophragmata fur n ish effi ci en t pathways for the tran sfer of

n u tritive materials from the i n ter fi b er tiss u e spaces i n to the —

fi b er .

The locatio n of the smaller J gran ules close to the t el op hrag -

mata acco un ts for the co n fu sio n of these gran u les with the
co n stit u e n t eleme n ts of the accessory discs as well as with those ,

of the co n tractio n ban ds R etzi u s claims that the accessory

.
21

discs are act u ally aggregatio n s of J sarcosomes Holmgre n -

.
,
6

Heiden hain an d others claim that the co n tractio n ban ds of

,
5

co n tracted m u scle also are simply aggregatio n s of sarcosomes .

That n either of these i n terpretatio n s however is te n able is , ,

proved by the fa ct that i n alcohol fix ed tiss u e i n which the —

,

sarcosomes have disappeared thro ugh sol utio n both accessory ,

discs an d co n tractio n ba n ds still occ ur M oreover both of these .

,

str u ct ures occ ur also i n fi b ers i n W hich n o sarcosomes can be

demo n strated B oth accessory disc an d co n tractio n ban d are
.
,

largely at least composed of i nt ram yo fi b rillar eleme n ts The

,
.

sarcosomes are i n terfibrillar bodies .

The fi b er ill ustrated i n fi g ure 1 5 is at an early phase of c on

tractio n The Q disc has le n gthe n ed a n d its m yo fi br i llar elemen ts
.
30 H . E . JOR D AN

are pale medially an d darker stai n in g an d k n obbed termi n ally .

T here is obvio u sly goi n g on a tra n sfer of deeply stai n in g s ubstan ce

from the middle of Q toward the middle of J This process is
carried still f urther i n the fi b er of fi g ure 1 9 where relatively ,

pale thick co n tractio n ban ds have formed This stage corre .

sp o n d s to the so called homoge n eo u s phase of co n tractio n o f c ert a i n

-

writers The thick pale early co n tractio n ban ds co n de n se

.
, ,

i n to the typ ical defi n i t i v e thi n deeply stai n i n g co n traction

ban ds of the f ully co n tracted fi b er (fi g These same steps .

are show n i n lo n git u din al series within fo ur sarcomeres i n fi g ure

1 0 drawn from a preparatio n of leg m u scle fi b er of the grass
,
-

hopper .

Figure 1 6 is give n to ill ustrate the character an d distrib u tion

of the n u clei This fi ber is i n the relaxed co n ditio n an d o n ly
.

lightly stain ed The n u clei are ce n trally placed (fi g 1 7 ) an d are

. .

larger an d smaller spheroidal bodies with a delicate retic ulu m ,

an d relatively large irreg ular very chromatic n et k n ots The

, ,
-
.

nu clei occ u r arra n ged i n lo n g axial col u mn s ; i n deed i n certai n ,

fi b er s they seem to occ u py the core of the e n tire fi b er Where .

they occ ur i n shorter gro ups the termin al n u clei appear to be,

smaller than those more medially placed O ccasio n ally a reg ular .

progressive decrease i n size occ urs i n both direction s from the

middle to the termi n als of the gro up (fi g N o mitotic .

fi g ures were see n N u clear m u ltiplicatio n is here excl u sively

.

by the amitotic mode Cross sectio n s ( fig 1 7 ) show that the

.
—
.

fi b ers are of irreg ular cyli n dric form an d that the m yo fi br il s ,

( sarcostyles ) are of the lamellar type These lamellar sarco .

styles occasio n ally show a peripheral radial splittin g givi n g rise ,

to Y shaped sarcostyles as see n i n tran sverse sectio n

-
.

b . W i n g muscle

The wi n g m u scle di ffers markedly from the leg m u scle It .

con sists of rob u st fi b ers of irreg ular polygo n al form in tran s

,

verse sectio n (fi g The fi b ers are e n veloped by a delicate

.

sarcolemma A n occasio n al n ucle u s is peripherally placed the

.
,

maj ority are scattered i n discrimi n ately thro u gho ut the fi b er .

The n u clei are small oval bodies and are very chromatic ; i n
 STRIP E D M US C LE OF WA SP 31

tran sversely cu t and stai n ed sectio n s of fi b ers they are with

diffic ulty distin guishable from the very ab un dan t sarcosomes In .

ge n eral they are slightly larger than the large sarcosomes The .

fi b er is composed of relatively very rob u st sarcostyles amo n g ,

which the sarcosomes are scattered i n great prof usion In the .

photograph the sarcostyles appear white the sarcosomes an d ,

nu clei black ( fi g . The sarcostyles are circ ular i n o utlin e

i n tran sverse sectio n an d vary slightly i n diameter The sar .

c oso m es are i rr eg ul arl v stellate or f u siform bodies resembli n g

somewhat te n do n cells as see n i n cross sectio n (fi g s 4 5 an d -

.

L o n git u di n al sectio n s (fi g s 1 3 an d 2 3 ) show that the sarcosomes

.

are of irreg ular form b ut frequ e n tly appare n tly oval an d that
, ,

they are arranged i n si n gle an d do uble col umn s betwee n adj ace n t
sarcostyles Fig ure 2 1 a drawi n g of fi v e adj ace n t sarcostyles i n
.
,

lo n git u di n al sectio n shows a freq u e n t arran geme n t of the sar

,

co so m es n amely i n oval gro u p

, ,
.

Two other importan t facts are revealed i n fi g ure 2 1 First .

the regular align men t of the telophragmata i n adj ace n t sarcostyles

in dicates that they span also the i n t ersar co st yli c space This .

is den ied by Thuli n w ho claims that a telophragma is lacki n g

,
29

i n the wi n g m u scles of Coleoptera D iptera an d Hyme n optera , ,

.

Bu t his ill u stratio n s show clearly that his co n cl u sio n s are based
u po n seco n darily modified fi b er s that is fi b er s that have s u ffered , ,

distortio n an d r upt ure That the i n t er sar co st yli c represe n tative

.

of the telophragma is very delicate is demo n strated by the

ease with which it r upt ures i n fi x ed an d mechan ically han dled
tiss u e But that it is act u ally prese n t is fu rther proved by s u ch
.

examples as ill ustrated i n fi g ure 22 where a slight be n di n g an d

distortio n has simply drawn o ut an d i n co n sequ e n ce emphasized , ,

the i n t ersarco st yl i c portio n of the co n tin u o u s telophragm a .

Wh ere the sarcosomes are very ab un dan t however as i n fi g ures , ,

1 3 a n d 2 3 the telophragmata can n ot be traced amo n g the gro ups

,
'

of sarcosomes Here the telophragma mu st be either fe n es

.

t r at e d or have become r u pt u red a n d probably completely

destroyed In s u ch areas also the primarily reg ular (n ormal )
.

t ra n sverse alig n me n t of si m ilar portio n s of adj ace n t sarcostyles

has become Vitiated .

TH E A M ERIC A N J OU R N A L OF A N AT OM Y VOL , . 27 , NO . 1
32 H . E . JOR D AN

Certai n additio n al facts sho uld here be recorded regardi n g the

sarcosomes Whe n arran ged i n si n gle series betwee n adj ace n t
.

sarcostyles the lo n g axis of these sarcosomes is ge n erally at right

,

an gles to the lo n g axis of the sarcostyles ; whe n i n do uble series ,

their lo n g axis is i n ge n eral parallel with the lo n g axis of the

sarcostyles Certain irreg ular forms also occ ur wedge shaped
.
,
-

(fi g crescen tic (possibly collapsed ovals ) f u siform an d

.
, ,

stellate forms (fi g Tran sverse sectio n s reveal the fact .

that all types an d taken as a whole the vast maj ority of the, ,

sarcosomes have win g like processes which almost completely —

e n sheath the sarcostyles (fi g s 4 5 an d In this material n o .

very small spherical sarcosomes occ urred The sarcosomes of .

the wasp s wi n g mu scle were preserved i n all of the fix i n g fl ui ds

’

u sed ( Flemmi n g s fl ui d 95 per ce n t alcohol an d 1 0 per ce n t

’
,

formali n ) They m u st therefore co n tai n i n large meas ure

.

somethi n g i n additio n to lipoids Their irreg ular wi n ged form .

appears d ue to seco n dary mechan ical m o di fi cat i on s res ulti n g

from the press ure exerted by adj ace n t sarcostyles The evide n ce .
,

F ig . 10 L g i t di
on u i
n a l s ec t o n o f p o rt o n o f i l eg m u s c l e fi b e r
-
of a g r a ss ho pp er .

Th e fib e r p as se s a b ru p t l y f ro m a c o n di t i o n o f rel a xat i on t o on e o f c on t ra c t i o n
at p i o nt A, w h e r e a co n t ra c t i o n b an d can b e seen f o rm i n g b y p r o c ess o f f u s i o n o f o pp o
s it e v
hal es o f i v d k d s c s g i n t t h i n v l v d t l p h g m Th
t wo su c c es s e ar 1 a a s e o e e o ra a . e

t l e o p h gm i ra l g di n i b l t t h p h
a s no on f er nt t i n h wn
s ce r t A du
e a e a se o co ra c o s o a ,
e

p b b l y t i t b av i g b m t t h d b y t h h i z n t l t i n t t h i l v l t
ro a o s n ec o e s re c e e or o a en s o a s e e o a

d g e f d h
re e o y b y ed th limit f m i
ca c ep i c V i i n Th f m t i n f t h
on e s o c r o s co s o . e or a o o e

co nt ct i n b n d
ra b l wAh
o a ff t d t u v l f t t
s e o as e g d ec e a r e re e r sa o s r 1 a 1 o n s a s re ar s a

d p l y t i i n g n t t u t f t h d k di
ee s a n co F l mm i n g fi t i n i n h m
s 1 en o e ar sc . e
’
s x a o ,
ro -
e a

t y li n t i n
ox X 900
s a . .

Fi g 1 1 T n v . ti n f
ra w i n g m u l fib
s ers e s ec f th w p
o N u p iph
o a -
sc e er o e as . .
,
er

era l u l u ; th
n c enu l i s o tt d t h u gh u t t h di m t
er c e ar e f t h fib
sc a ; ere ro o e a e er o e er

S .
,
l mm
sar c o e Th l i gh t a . p n t w f dj c n t
e a r e astyl s ; th re r ese ro s o a a e sar c o s e e

d k ar i t v n ing
a r eas , n m er F l mm i n g fi x t i n i n h m t x y l i t i n
e sar c oso es . e a o ,
ro -
e a o n s a .

X 400 ( T h p h t m i g p h f fi g 1 1 1 2
. e o o d 13 w
cr o m d b y M Wi l l i m
ra s or s .
, ,
an ere a e r . a

S D u nn C
. l l Un iv
, ity M d l S h l N w Y k )
o rne ers e l ca c oo , e or .

Fi g 1 2 Sm ll . f m fi g u 1 1 m h i gh l y m g n ifi d T h i u l ou t
a ar ea ro re ore a e . e c rc ar

li n es o f th tyl l
e s ar c o s l y h wn ; l th nv l p i n g w i n g l i k p
es ar e c ea r e es
s o a so e e e o -
e ro c ss

o f th i t v nin g ie n er gu l e m N u nu l u
r re X 900
a r s a r c o so es . .
,
c e s . .

F i g 1 3 L n g i t u di n l t i f p t i n f fib f w p w i n g m u s l h w
. o a s ec on o or o o er o as
’
s c e s o

i n g t h l i g h t l y st i n i n g
e ty l nd t h
a int v ning d k c m
s ar c o s Th
es a e er e ar sar o so es . e

co n p i u ou
s t n vc lin
s p
ra t t h t l p h gm t
s er s e Z t el p h g m
es re r ese n M e e o ra a a .
,
o ra a

m p h gm ; S
e so ra m ; t
a tyl X 1 300
c , s a r c oso e s ,
sar c o s e . .
34 H . E . JOR D AN

which will be f urther disc ussed below seems to i n dicate that the ,

sarcosomes are n ot tran sie n t nutri tive reservoirs b ut are formed ,

d uri n g the early stages of m u scle histoge n esis to remain ,

thro u gho u t the life of the i n divid ual A ppare n tly o nly very .

few if any co n ti nu e to be added an d few if any e n tirely disappear

,
.

A s the sarcostyles i n crease i n nu mber an d grow i n girth probably ,

coin cide n t with the growth of the sarcosomes the sarcostyles ,

c rowd u po n t h e sar co so m e s an d by press u re alter their origi n al

x

spheroidal shape i n to irreg ular wi n ged forms .

We come n o w to the ce n tral poin t of this i n vestigatio n the ,

prese ntatio n o f the data W hich prove tha t the wi n g m u scle -

s arcostyle of the wasp passes thro u gh the same morphologic

phases d urin g co n tractio n as the leg m uscle fi b ri l s that is it —

, ,

u n dergoes a reversal of striatio n with the formatio n of a typical

co ntractio n ban d ; an d i n ciden tally that Schaefer s diagrams

, ,
’

of m uscle co n tractio n are based u po n the erro n eo u s i n terpreta

tio n of an artifact as a co n tracted sarcostyle .

Fig ure 2 5 shows the a ppearan ce of the W i n g m u scle sarcostyle -

o bserved i n the fresh co n ditio n i n R i n ger s sol utio n The in tra ’

.

sar c o st yl i c telophragmata are very co n spic u o u s as relatively

coarse stripes dark at the high level of foc u s light at low level
, ,
.

The dark disc occ upies almost the e n tire space betwee n s u ccessive
telophragmata the light disc appears as a very n arrow light area
,

o n either side of the telophragma In tiss u e fi x ed i n Flemmi n g s

’
.

fl ui d an d stai n ed lightly with iro n hematoxyli n the appeara n ce

—

of the sarcostyle is practically ide n tical ( fig The light .

disc appears slightly thicker the fi b er as a W hole seems to have

,

a slightly greater diameter probably the res ult of a slight swelli n g

,

actio n of the Flemmi n g s fl ui d ; an d there occ urs a slight c on

’

de n satio n of the deeply stain in g s ubstan ce i n the m i dreg i o n ,

perhaps i n dicati n g the locatio n of the mesophragma F i x at i o n .

with 9 5 per ce n t alcohol ca uses a shri n kage of the e n tire sarco

style a n d a disti n ct co n de nsatio n as the res ult of dehydratio n
, , ,

of the deeply stai n i n g s ubstan ce ( fig In s u ch a sarcostyle

.

the light disc appears somewhat thicker than the dark disc The .

d ehydrati n g a n d shri n kage e ffect of the alcohol shows mai n ly

i n Q demo n strati n g its relatively more fl uid co n siste n cy
,
Fix a .
36 H . E . JOR D AN

styles Sarcostyles occ urri n g i n gro ups held together by the

.
,

telophragmata are appare n tly fort i fied to some exte n t agai n st the
,

di storti n g e ff ects of the hyp oto n ic sol utio n an d so mai n tai n a ,

relatively eve n co n to u r That o ur interpretatio n of the sar

.

c o st yles of fi g ur es 2 9 an d 3 1 as swolle n relaxed fi b ers is correct ,

may be proved by tra n sferri n g s u ch sarcostyles to a 2 per ce n t

sodi um chlorid ( hyperto n ic ) sol utio n where they ret ur n to a ,

co n ditio n very similar to that seen i n R in ger s sol utio n ’

.

I n stead of h avi n g the special advan tage claimed for i t R ol ,

let s techn ic is i n fact very un favorable sin ce it leads to dis

’
, , ,

t ort i o n a n d the res u lti n g co n f u sio n i n i n terpreti n g res u lts It .

reveals n othi n g that cann ot be see n almost as well i n un st ai n ed

fresh material n or a n ythin g that can n ot be better see n i n
,

alcohol fix ed and iro n hematoxyli n stain ed preparatio n s an d it

— — —
,

is i n ferior to Toiso n s sol utio n i n which the fresh tiss u e is vitally

’

stain ed i n a n on distortin g isoto n ic sol utio n

—
.

The f un ctio n ally co n tracted fi b er is shown i n fi g ure 3 3 This .

drawi n g was made from a fresh preparatio n i n R i n ger s sol utio n ’

.

It correspo n ds with Meig s ill u stratio n of a co n tracted sarco

’

style of the fly s wi n g m u scle (fi g

’
an d with fi x ed co n tracted .

fi b er s i n stai n ed sec tio n (fi g Similar fi b ers may be see n i n

.

Toiso n s sol utio n an d occasio n ally also i n preparatio n s after

’
,

R ollet s tech n ic It sho uld be n oted that the sarcostyle i n this

’
.

co nditio n of f ull co ntractio n is no t beaded and that the c on ,

tractio n ban d i n places appears clearly do u ble Sarcostyles i n .

the co n tracted co n ditio n can be readily obtain ed i n fresh an d

fi x ed preparatio n s by drawi n g a sharp n eedle tran sversely across
a gro up of livi n g sarcostyles The mechanical stim ul u s ev i .

d en t l y is suffi c i en t to i n d u ce f ull co n tractio n .

Fig ures 3 4 3 5 an d 36 represe n t sarcostyles from the wi n g

, ,

m u scle of the elater beetle Al aus o c ul at us This m u scle was fi x ed

,
.

i n 9 5 per ce n t alcohol an d stai n ed with iro n hematoxyli n The -

.

sarcostyles di ffer from the wasp s sarcostyles o n ly i n bei n g sligh tly ’

coarser They ill ustrate two importan t poi n ts regardin g this

.

typ e of sarcostyle especially well The sarcostyle of fi g ure 3 4 is .

o n e of a small gro up at a n early stage of co n tractio n the median

, ,
 STR IPE D M US C LE or WA SP 37

( H ) disc bei n g co n spic u o u s The sarcostyle is slightly c on

.

st ri ct ed at the levels of the dark discs d u e to the dehydratio n

effect of the alcoholic fi x at i o n Beadin g is clearly a fi x at i on

.

artifact not an in dex of co n tractio n This co n cl u sio n is further

,
.

s upported by the sarcostyle of fi g ure 3 5 This w as an isolated .

sarcostyle lyi n g at the en d of a cl ump an d it was i n co n sequ e n c e ,

m o di fi ed to a greater degree It was moreover i n the relaxed

.
, ,

con ditio n The sarcostyle is sharply beaded du e to an ex

.
,

cessi v e dehydratio n an d co n seq u e n t co n de n satio n

,
of the dark ,

disc The osmotic e ffect of a dehydrati n g flui d or of a hyper

.
,

ton ic sol utio n shows itself fi r st an d to greatest degree i n t he

, , ,

dark disc This in dicates that the dark disc is more fl ui d than
.

the light disc which latter is i n additio n held open by the mor e
,

resistan t telophragma If the location o f the beads at t he

.

levels of the telophragmata were here excl u sively the res ult of
the relatively more resistan t n at ure of the membran e holdi n g
the fi b er open at this poin t rather than the res ult of a relatively
,

greater fl ui di t y of the dark disc the border of the bead wo uld b e

,

expected to be wrin kled i n stead of havin g a sharp co n to ur It .

might be obj ected especially i n co nn ection with fi g ure 3 5 that

, ,

the in vestigator might readily co n fuse telophragma and meso

phragma an d so misco n str u e the osmotic effects of the fi x i n g
,

fl ui d s . D etermi n atio n of telophragma levels however is i n fact , ,

a relat ively simple matter The sarcostyles are exten sively

.

fract u red especially i n the section ed material The levels of

,
.

fract ure i n the sectio n s an d i n the gold chlorid and fresh prepa
,
—

ration s occ u r almos t i n variably alo n g the telophr agmata The

, .

criterio n of level of fract ure j udicio u sly applied i n conn ection

, ,

with the observatio n of the order of stria tio n i n distan t parts of

the sarcostyle furni shes a precise test for the i dent i fi c at i on of
,

the telophragma i n do u btfu l cases .

The sarcostyle of fi g ure 3 6 was fi x ed with Flemmin g s fl ui d ’

an d lightly stain ed with iro n hematoxylin It appears to be at

-
.

a mi dph ase of co n tractio n It has special i n terest becau se it

.

shows the presen ce of fin e co n stit uen t s arcost yli c m et afib rils ‘

.
’

These are especially promi n en t at the u pper en d where they

stain ed deeply A fter R ollet s method of preservatio n an d
.
’
38 H . E . JOR D AN

stai n in g a similar phen omen o n occ urs Fig ure 46 ill ustrates an
,
.

en d view ( optical tra n sverse sectio n ) of a sarcomere from s u ch

a preparatio n Here the co n stit u e n t met afib ril s are largely

.

gro uped i n the mann er of more or less sharply co n to u red circles .

The latter s uggested to Schaefer the idea of a porifero us c o n

ditio n of the dark disc These pores certa m are fi x at i on arti
.
‘ ’

facts Whether the fib ril s (m et afi bril s) sho uld be similarly

.

in terpreted or whether this techn ic simply re n dered more con

,

S p i c u o u s co n sti tu e n t m et afi b ril s of the sarcostyle can n ot be ,

fi n ally decided These sarcostyles certain ly un der most c o n

.
,

d i t i on s both fresh an d fi x ed both i n lo n git u di n al an d tra n sverse

, ,

sectio n appear homoge n eo u s (fi gs 2 3 an d

,
This homo
.

g e n ei t y may be simply d u e however to a closely similar r efr ac

, ,

tive i n dex o n the part of these m et afi b rils an d the i nt r asar

c o st yl i c sarcoplasm so that the prese n ce of the m et afi b ri l s ca nn ot
,

be discern ed un til ren dered con spic u o u s by certai n fi x i n g a n d

stai ni n g processes .

Figur es 3 7 to 44 are twice the magn ifi catio n of fi g ures 2 5

to 36 n amely 2 600 diameters They are all of sarcostyles of
, ,
.

the win g m u scle of the wasp from tiss u e fi x ed with 9 5 per cen t
,

alcohol an d stai n ed with iro n hematoxylin Fig ures 3 7 an d 38 —

.

are both of relax ed fi b ers as in dicated by the un divided co n dition

,

of the dark disc The q u estio n arises as to why the dark disc
.

o f fi g ure 3 7 is more than twice the le n gth of that of fi g ure 3 8 .

It will be n oted also the dark disc of fi g ure 3 8 is more compact

than that of fi g ure 3 7 which latter is moreover slightly lighter
, , ,

i n color an d appare n tly lo n git u di n ally striped The short le n gth .

of the dark disc of fi g ure 3 8 as compared with this disc of the

,

relaxed fi b er i n fresh co n ditio n or after Flemmin g s fi x at i on ’

( compare with fi g s 2 5 a n d. is the res u lt of the dehydrati n g

( co n de n satio n ) e ffect of the alcoholic fi x at i o n The greater le n
gth .

an d so m ewhat di ffere n t character of the dark disc i n fi g ure 3 7 is

to be explain ed i n terms of a seco n dary stretchi n g of the fi b er .

These t w o sarcostyles ill ustrate the primary e ffects of exosmosis

an d mecha n ical te n sio n u po n the dark disc res u lts to be expected ,

if the dark disc is relatively more fl uid than the light disc .
 STRIPE D M US C LE OF WA SP 39

Figures 3 9 an d 40 ill u strate the same poin t i n co nn ectio n with

a sarcostyles at an early stage of co n tractio n However it does .
,

n ot seem to me that the above i n terpretatio n is q u ite adeq u ate

to explain all the variatio n s i n le n gth of the dark disc i n dif

fer en t sarcostyles . It un do u btedly acco un ts for a co n siderable
portio n of thi s variatio n b ut I feel co n strain ed to co n cl u de after
,

careful a nd prolo n ged st u dy of man y relaxed sarcostyl es that ,

the amo un t of deeply stain i n g s ubstan ce varies co n siderably i n

di ffere n t fi b ers at the same f un ctio n al stage i n the same micro
scopic preparatio n It seems to me fairly certai n that the
.

amo un t of dark an d light s ubstan ce varies withi n certain limits

i n di ffere n t appare n tly n ormal sarcostyles at the same fun ctio n al
, ,

stages an d un der the i n fluen ce of appare n tly ide n tical mechan ical
,

an d osmotic co n ditio n s .

Fig ures 40 and 4 1 at approximately the same fun c tio n al

,

stage (mi dpha se of co ntractio n ) sho uld be co n sidered together

,
.

O n e has a straight co n to u r the other is slightly beaded Fig u res

,
.

42 a n d 43 at a later fun ctio n al stage ill u strate the same poi n t

, , .

The sarcostyle of fi g ure 43 is at the same f un ctio n al stage as the

middle portio n of fi g ure 42 ; o n e sarcostyle is beaded the other ,

is n ot These fo u r fi gures (40 to 43 ) demo n strate agai n that

.

beadi n g is n ot an i n dex of a distin ct fun ctio n al co n ditio n that ,

is of degree of co n tractio n ; it is obvio u sly a fi x at i on artifact

,
.

This co n cl u sio n is re n dered certain by the co n ditio n of the

sarcostyle of fi g ure 44 T hi s sarcostyle is i n full co n traction an d
.

is characterized by co n spic u o u s co n tractio n ban ds b ut it still ,

main tain s a straight lateral co n to ur .

To ret u rn to fi g ure 42 the lower portio n of the sarcostyle is at

,

the same fun ctio n al phase as the sarcostyle of fi g ure 4 1 In the .

u pper an d mi ddle levels two co n tractio n ba n ds are begi n n i n g

to form The u pper o n e is at a later phase than the n ext

.

lower which shows thicker deep stai n i n g termi n al portio n s It

,
— .

is clearly see n here that the formatio n of the co n tractio n ban d

res ults from a movemen t of the deeply stain in g s ubstan ce of the
divi din g Q disc toward the telophragma But this material does
-
.

n o t here move en masse b ut i n s u ch mann er as to prod u ce

,
40 H . E . JOR D AN

tran sie n tly a thick less deeply stai ni n g ( dil uted ) co n tractio n
,

disc bisected by the telophra gma This disc ( co n tractio n ban d)

.

then co n den ses abo ut th e telophragmata to form the deeply

stain i n g relatively thi n co n tractio n ban d of the f ully co n tracted
, ,

fiber (fi g . The dil uted co n ditio n of th e co n tractio n disc

‘ ’

described above (fi g s 42 an d 4 3 ) represe n ts the so called homo

.
-
‘

g en eou s phase of co n tractio n It is o n ly rarely co n spic u o u s in

’
.

certain fi b ers an d seems to dep en d u po n the rapidity of the

,

co n traction process Whe n the process is rapid this very

.
,

tran sien t phase will ge n erally escape detectio n or may possibly ,

be abse n t an d the halves of Q seem to have moved en masse

,

aga in st the telophragmata .

In the co n tracted sarcostyle of fi g ure 44 an irreg ular clo u dy ,

li n e appears midway betwee n s u ccessive co ntractio n ban ds .

This may represen t a remn an t of deeply stain i n g material of the

orig i n al Q disc of the relaxed fi b er cli n gin g to the mesop hr agma
-
,
.

It is n oteworthy also that the telophragma is n o t discer n ible

i n this later co ntractio n ban d ( compare fi g s 42 an d Th e .

maskin g of the bisecti n g telophragma may be due to its havi n g

become closely adhere n t to an d so hidde n b y on e of the halves
, ,

of the deeply stai n in g co n tractio n ban d or to the fact that it ,

‘

has bee n stretched i n the shorte n ed wide n ed co n tracted sar , ,

co m ere to a poi n t where it is n o lo n ger co n spic uo u s un der the

ordi n ary powers of the microscope The paired co n stit utio n of .

the co n traction ban ds is a fact of prime importan ce i n con

n ec t i on with o ur i n terpreta t io n of the several varieties of the

sim ple type of i n tercalated disc as irreversible con traction ban ds .

These ill ustratio n s s upply the an swer to an other importan t

q uestio n regardi n g the str u ct u re of the sarcostyle n amely , ,

whether it is n aked or e n veloped by a sarcolemma Meigs ‘

.
’ 15

states that the sarcostyle of the fly s win g mu scle is homo ’

g en eo u s i n str u ct u re a nd is n o t s u rro un ded by a membran e .

The mo di fi cat i o n s produ ced by hypoton ic sol utio n s (fi g s 2 9 to .

3 2 ) an d by alcoholic fi x at i on (fi g s 3 4 3 5 4 1 an d 43 ) demo n strat e

.
, , ,

the fact that the sar costyle has a physically di fferen t peripheral
layer which performs the f un ctio n of an osmotic membran e .
 STRIPE D M US C LE OF WA SP 41

An d the beadin g of the sarcostyle i n hypoto n ic sol ution s is de

“
pen de n t u po n the prese n ce of the telophragma an d its attach
me n t to a peripheral layer with the properties of a membran e .

The same fact is ill u strated i n fi g ures 46 an d 47 Fig u re 47 shows .

i n tra n sverse sectio n s three s u ccessive stages i n the progressive

destain in g after applicatio n of the iro n hematoxyli n stai n The -
.

late phase shows a ce n tral deeply strai nin g gra nu le (fib ri l ? ) an d

a delicate peripheral membran e .

VI . DIS C USSI O N
a . Telop hm gma an d mesop hragma

In the fun damen tal morphology of striped mu scle there is

perhaps n o eleme n t more un iform an d defi n i t e than the telo
phragma Thulin recen tly p ublished a paper i n which he
.
29

claims that a telophragma does n o t occ ur i n the wi n g m u scles of

Coleoptera D iptera an d Hyme n optera His ill u stratio n s make
, ,
.

it clear that he is referri n g to the i nt er sarcost yli c portio n of the

telophragma for the i nt rasarcost yli c represe n tative is plai n ly
,

i n dicated T hul in s sole evide n ce i n s upport of his claim that

.
’

these mu scles lack a telophragma is hi s observatio n s o n sectio n ed

ma terial . His ill u stratio n s show that the material s uffered c on
si d er ab l e distortio n an d fragme n tatio n by his tech ni c as i n di ,

cat ed chi efl y by the relatively wide spaces betwee n adj ace n t

sarcostyles I have examin ed specimen s of fresh an d fi x ed win g

.

m u scle of represe n tatives of each of these gro ups of i n sects In .

fresh specimen s teased i n R in ger s sol u tio n gro ups of closely ’

adhere n t sarcostyles occ ur i n which the telophragmata lie at the

same tran sverse levels over as man y as from six to t en adj ace n t
sarcostyles The slight resistan ce o ffered agai n st dividi n g a
.

gro up of sarcostyles by teasin g is additio n al evide n ce that a

co n n ectin g telophr agma exists .

From a st u dy of well fixed tiss u es similar evide n ce accr u es

— .

E xami n atio n of s u ch areas as represe n ted i n fi g ur e 2 1 c an leave

n o do u bt that the telophragmata of the sarcostyles S pa n also the

extremely n arrow i nt ersar co st yl i c spaces an d hold the several

discs of the sarcostyles i n uniform horizo ntal alig n me nt T hi s .
42 H . E . JOR D AN

co n cl usion is re n dered in co n trovertible i n the light of s uch

eviden ce as is show n i n fi g ure 22 where the i nt ersarco st yli c ,

portio n of the telophragma can act u ally be disti n ctly see n b e

cau se slightly drawn o ut o i align me n t by reaso n of a be n di n g of
.

the two i n volved sarcostyles .

Thul in makes a similar claim of n on occ urren ce for the

29 -

mesophragma It is more di ffi cult to prove the existe n ce of an

.

i n t er sarco st yl i c portio n of this membran e du e to its extreme

delicacy It m ay be stated however that i n the section s
.
, ,

illu strated by Thu li n we wo uld n ot expect to see the i n t ersar

c ost yl i c portio n of the mesophragma This membran e eve n if .
,

prese n t wo uld n ot be able to withstan d the te n sio n exerted by

,

the widely separated sarcostyles ; an d eve n if suffici ent ly elastic

to be able occasion ally to mai n tai n an i n t ersarc o st yl i c i ntegrity ,

it wo uld have become too delicate to allow detec tio n by o ur

prese n t methods of microscopic examin atio n That the sarco .

styles themselves co n tai n m es0 phr ag m at a as well as telophragmata

cann ot be do ubted (fi g s 1 3 an d . That an i nt ersarco styli c
telophragma exists is also certai n ( fig This membran e is .

i n timate ly attached to the peripheral ser c olemm a cau si n g a ,

festoo n in g of the latter un der certai n art i fi ci al co n dition s (fi g .

A ttachme n t to n u clear wall may be i n ferred from o b

serv at i o n s o n the m u scle of other forms b u t I am u n able to ,
8

d efi n i t ely demo n strate s u ch co nn ectio n i n this m u scle .

Co n trary to co n ditio n s i n the mantis wi n g mu scle for ex ,

10

ample where the telophragmata form effi ci ent barriers agai n st

,

the lo n git udi n al moveme n t of the sarcosomes the telophragmata ,

of the wasp s wi n g m u scle m u st be either fen estrated or have

’

become seco n darily r upt ured an d destroyed i n certai n regio n s ,

i n order to allow for the acc umu latio n of large oval gro u ps (fi g .

2 1 ) an d compact col u m n s ( fi g 2 3 ) of sarcosomes The ex p eri

. .

me n tal evide n ce from treatmen t of the sarcostyles with hypo

to n ic an d dehydratin g solu tio n s i n dicates that the i nt rasarco
stylie telophragma is a relatively rigid an d b ut slightly exte n sible
membran e The rigidity of the membran e is i n dicated by fi g ures
.

2 9 3 0 3 1 3 2 3 4 an d 3 5; its slight exte n sibility by the sharp

, , , , , ,

un beaded lateral co n to u r of f un ctio n ally co n tracted sarcostyles

, ,
44 H . E . JOR D AN

b S arcosomes
.

Sarcosomes occ ur both i n the leg an d the wi n g mu scle of the

wasp In the m antis I co uld detect sarcosomes o nly i n the wi n g
.

m u scle In the leg m u scle of the wasp (fi g

. as i n the wi n g .

m u scle of the man tis they occ u r as two disti n ct gro ups ; a gro u p
,

of s m aller spherical eleme n ts (J granu les ) o n either side of a n d —

close to the telophragma an d a gro u p of larger oval eleme n ts

,

( Q gra n u les ) alo n g the midli n e of the dark disc Both i n the
—
.

leg mu scle of the wasp a n d the wi n g m u scle of the man tis these
gran ules are dissolved by alcohol b ut are preserved i n tiss u e ,

fi x ed with stro n g F lemmi n g s fl u i d or a 1 0 per ce n t formalin

’

sol utio n This microchemical reactio n s u ggests an essen tially

.

lipoid n at u re The most plau sible i n terpretatio n of thes e

.

granu les that all th e available data s upport is that they a re

formed i n close associatio n with the telophragma the latter ,

fur n ishi n g the pathway by which their co n stit u e n t eleme n t s are

carried from the i nt ersarc ost yli c tiss u e spaces an d that as they ,

grow i n size they become crowded toward the middle of t h e

sarcomere ( perhaps aided i n this moveme n t by m u scle con
tractio n ) an d here m odi fi ed i n to oval str u ct u res thro ugh t h e
,

m ut u al press ure of adj ace n t sarcostyles Their f un ctio n may b e .

ass umed to be n u tritive The above i n terpretatio n n ecessitate s

.

the fu rther i n fere n ce that the l arger sarcosomes are co n ti n u ally

u sed u p thro u gh the f un ctio n i n g of the m u scle fi b er a n d n ew ,

o n es co n ti nu ally formed i n the Vici n ity of the telophragma .

T he J sarcosomes lie at the levels of the accessory disc ( com

-

pare fi g s 1 4 an d . This spatial j uxtapositio n has led certai n

i n vestigators ( R etzi u s 9 0 ) to i nterpret the accessory disc as
,
’ 21

composed of J sarcosomes ; therefore a str u ct u re composed o f

-
,

i n t ersar c o st yl i c eleme n ts A gai n the overlappi n g of the c o n

.
,

tractio n ban ds i n co ntracted fi b ers with the two series of J

gra nules has give n a basis for the con cl usio n ( R etzi u s Holm ,
21

gren an d Heide n hai n ) that the co n tractio n ban d ( disc ) is

,
6 5

prod u ced by the aggregatio n of J granules The error of s u ch -

.

explan atio n of the accessory disc an d the co n tractio n ban d is

proved by the prese n ce of both of these str u ct ures i n fi b ers i n
 STR IPE D MU S CL E OF WA SP 45

which the sarcosomes have bee n dissolved by alcoholic fi x at i on

(fi g s 1 4 a n d
. Both accessory disc an d co n tractio n disc are
primarily an d esse n tially i n t rasarco st yli c (fi b rill ar) eleme n ts .

The sarcosomes of the wi n g mu scle of the wasp di ffer m arkedly

i n certai n respects from those of the leg m u scle In the fi r st .

place there are n o really smaller sarcosomes s u ch as correspo n d

, ,

to the J gran ules of the leg mu scle This wo uld seem to i n di

— .

cate that a n ew formatio n of sarcosomes does n o t occ u r or on ly ,

occ u rs to a slight degree perhaps o n ly i n certai n yo un g fi b er s

, .

In the seco n d place the sarcosomes are n o t gro u ped at d efi n i t e

,

horizo n tal levels b ut lie betwee n adj ace n t sarcostyles i n short

,

oval gro ups (fi g 2 1 ) or lo n g col umn s i n si n gle or do u ble fil e

.

(fi g .

The wi n g mu scle sarcosome is a primarily oval body which

-
,

becomes seco n darily modi fied thro u gh the operatio n of mechan

,

ical factors in cide n t to the mu t u al press ure of adj acen t sarco

styles i n to very irreg ul ar forms Tran sverse sectio n s of these
,
.

fi b er s show that the m o di fi ed sarcosomes have lo n g lateral , ,

wi n g like processes (the res ult of press ure ) which co n n ect with
—

similar processes from adj ace n t sarcosomes to form i n some

i n stan ces a complete sheath for the i n volved sarcostyles Frag .

m en t ed or un u s u ally pale sarcosomes were n ot see n i n this

tiss u e This s u ggests that the life of the sarcosomes i s n ot here
.

a tran sie n t o n e b ut that o n ce formed the sarcosomes persist

,

probably thro u gho ut the life of the m u scle fi b er Alcoholic .

fix at i o n c au ses collapse of certai n large sarcosomes i n dicati n g the ,

extractio n of fl ui d by the alcohol But the sarcosomes are i n.

gen eral al most as well preserved i n alcohol except for occasio n al ,

wrin kli n g of co n to u r as i n F lemmi n g s fl uid or 1 0 per ce n t

,
’

formali n .

These reactio n s to fix i n g fl ui ds i n dicate that the wi n g m uscle -

sarcosomes co n sist of some s ubstan ce i n additio n to lipoids The .

complete ge n etic history chemi cal co n ditio n an d fun ctio n al sig

, ,

n i fi c an c e of the sarcosomes remai n for the prese n t un k n ow n .

We are probably qu ite safe however i n ass umi n g tha t the

, ,

sarcosomes here described for the leg an d the wi n g m u scle of

wasp are esse n tially the same thi n g represe n ti n g simply less,
46 H . E . JOR D AN

highly an d more highly elaborated stages respectively i n the , ,

same esse n tially lipoid gran ules the complete series bei n g r ep re ,

se nted by the J gra nu les of the leg m uscle the large oval Q
-
,

granules of the leg mu scle an d the wi n g m uscle a nd fi n ally the ,

W i n ged sarcosomes of the wi n g m u scle The latter may possibly .

represe n t rem n a nts i n mechan ically m o di fi ed form of these prob

able reservoirs of reserve food material .

If o n e st u d ied the leg m u scle of the wasp o n ly with alcoholic

fi x at i o n o n e wo uld co n cl u de that this m u scle lacked sarcosomes
, .

R eports of lack of sarcosomes i n certai n i n sect m u scle may

perhaps be explai n ed o n the basis of faulty preservatio n by the
histologic tech n ic employed There may moreover most prob .
, ,

ably be wide variatio n s i n the ab un dan ce of these eleme n ts cor

respo n di n g to phases of maj or metabolic cycles of an i n divid u al .

The evide n ce to date s uggests that all m u scle co n tai n s the homo
l o g ues of the sarcosomes of i n sect m uscle i n at least some slight
degree of elaboratio n That the sarcosomes of the wi ng m u scle
.

are n ot mitocho ndria as has bee n s u ggested ( Th uli n B ullard )

, ,
29 2
,

is q u ite clear from their size shape an d resistan ce to the u s u al

, ,

mitocho n drial solve n ts It is d iffi c ult to see how M eigs was

.
15

led to the erro n eo u s co n cl usio n that i n the wi n g m u scle of the fly

“
the sarcosomes are almost certai n ly gra n ules of coag ulated
sarcopla sm ; there is n othi n g similar to be see n i n preparatio n s
of fresh m uscle (p ” . Co ntrary to this stateme n t n othi n g ,

co u ld be simpler than the isolatio n of these gran ules by teasi n g

fresh wi n g m u scle of fly i n R i n ger s sol utio n ’
.

c . The access ory di sc

This disc ( N stripe ) was fi rst see n by Bruck e i n 1 8 58 i n the

‘
-
’ ll

leg m u scle of H ydrO p hi l us p i ceus It has received careful st u dy .

at the han ds of R ollet who fi rst disposed of R etzi u s claim that

,
23 ’2
1

it co n sisted of J gran ules R ollet i n terprets it as co n sisti n g of

-
.

esse ntially the same s ubstan ce as the dark disc ; it is said to be

a nisotropic like the Q disc an d the telophragma b u t more
—
,

weakly an i st rop i c than Q .

The accessory disc is very co n spic u o u s i n the leg m u scle of the

wasp the elater an d the grasshopper It co n sists of mo di fi ed
, ,
.
 STRIPE D MU S CL E OF WA SP 47

portio n s ( spherical i n shape) of the myofi bri l s horizo ntally align ed ,

close to the telophragma It appears like a row of granules very

.
,

similar to the telophragma It stai n s like the Q disc an d has .

-

appare n tly a similar chemi cal co n stit u tio n Its ge n etic his .

tory an d its si g ni fi can ce however remain un k n own It becomes

, ,
.

involved with the dividi n g dark disc i n the formatio n of the

co n tractio n ban d .

O n e of the most i n teresti n g an d co n fu si n g facts regardi n g the

accessory disc 1 s i t s apparen tly tran sitory n at ur e By thi s I .

mean that it is n ot always prese n t It may be presen t i n on e .

fi b er an d abse n t i n an otherwi se similar adj ace n t fi b er R ollet .

23

also n oted its i n co n stan t n at ure i n arthropod m u scle This i n .

co n stan cy may be ge n etically related with the formatio n or ,

degree of developme n t of the J sarcosomes b u t defi ni t e i n

,
-
,

formatio n on this poi n t is e n tirely lacki n g .

d . The a n i sotrop i c su bstan ce

It w as fir st poi n ted ou t by Br u cke that the i ntra 1

fi b ri ll ar s u bstan ce of m u scle fi b er s occ urs u n der certai n c o n

d i t i on s i n alter n ati n g strata of isotropic an d a n isotropic materials .

M erkel an d R ollet i den t i fi ed this anisotropic s ub stan ce with

17 22

the material which gives the dark color to the dim disc of ‘ ’

striped mu scle whe n Viewed fresh with ordi n ary light These .

i n vestigators n oted a reversal of striatio n d uri n g co n tractio n

res ultin g i n the formatio n of co n tractio n ban ds They i n ferred .

that the co n tractio n ban d was composed of the an isotropic su b

stan ce of the dim disc of the relaxed fi b er T ourn eux an d .
30

R u therford also accept this i n terpretatio n of the co n tractio n

26

ban ds It is si gni fi c ant however that n o n e of these i n vesti

.
, ,

gators gives an ill u stratio n of the co ntractio n ban ds as see n i n

polarized light un der crossed n icols It has appare n tly bee n .

impossible to demo n strate an act u al reversal of striae as c on

cern s the an isotropic materials En glemann an d v an G ehu c h 4

“
.

t en - act u ally show as j u dged from their ill u stratio n s that the
? 32
, ,

a n isotropic s ubstan ce does n ot chan ge its locatio n d uri n g c on

tractio n Schaefer clai m s to have de m o n strated the same fact
.
28

TH E A M ERIC A N J OU R N A L or A N AT OM Y V OL , . 27 , N O . 1
48 H . E . JOR D AN

by u se of R ollet s gold chlorid tech n ic We have already show n

’
-
.
,

however that Schaefer as a matter of fact co n fu sed an arti

, , ,

fi c i ally swolle n co n ditio n of the fi b er res ulti n g from the actio n

,

of the hypoto nic formic acid water sol utio n of this techn ic with
- -
,

a co n ditio n of f un ctio n al co n tractio n The ass u mptio n that the

.

d arker q u ality of the dim disc of striped fi b er s is wholly the

res ults of the segregatio n of an isotropic materials withi n the
ge n eral limits of this disc un der certai n co n ditio n s has led to
m u ch co n fusio n an d has i ntrod u ced an eleme n t of unn ecessary
,

d i ffi c ul t y i n i n terpreti n g the morphologic chan ges s uffered by a

striped mu scle fiber d uri n g co ntractio n .

Moreover u n d ue emphasis u po n the prese n ce of anisotropic

,

materials i n this co ntractile tissu e has i n flu en ced some of the

leadi n g theories of co n tractio n c ulmin ati n g i n En gl em ann s hy ’

p o t h esi s that co n tractio n is esse n tially a matter of the ab sorp

4

tio n of isotropic by the an isotropic material However the .

,

prese n ce and segregatio n of sp eci fi c anisotropic materials cann ot

be held respo n sible for the dark color of the dim disc i n un ‘ ’

stai n ed fi b ers ; n o r are s u ch alleged an isotropic materials properly

i nterpreted as the s u bstan ce which takes the stai n i n the dark
disc i n stai n ed preparatio n s The dark disc is the resu lt of a
.

s ubstan ce apart from an d i n additio n to the alleged anisotropic

, ,

materials The stripi n g of the wi n g mu scle sarcostyles of the

.
-

wasp and other i n sects is q u ite as defi n ite if n ot more so both , ,

i n the un stai n ed an d stai n ed co n ditio n as i n the leg m u scle ; ,

yet o n ly i n the latter material can the st rat i fi c at i on of the an iso

tropic s ubstan ce un der certain co n ditio n s be clearly demo n
s t rat ed with the micropolariscope while i n the former it is
,

prese n t i n so small amo un t as to be practically impossible of

demo n stratio n i n the si n gle sarcostyle The s ubstan ce that .

moves d uri n g co ntractio n i n the formatio n of the co n tractio n

ban d is this additio n al deeply stai n i n g material I have c on .

vi n ced myself by a st u dy of the leg mu scle of the wasp that the

an isotropic material ( co n ditio n ) as segregated i n the relaxed

fi b er is n o t shifted i n to the light ( J disc d uri n g co n tractio n It .

is also clear however that the an isotropy of the fi b er is greatly

, ,

lesse ned by co ntractio n This latter fact is emphasized also by

.

Meigs .
15
 STRIPE D M US C LE OF WA SP 49

There is o ne ill u stratio n that calls for an explan atio n i n this

co nn ectio n It is fi g ure 7 1 by v an G ehu cht en
. which
r
31

shows a bisected an isotropic disc correspo n di ng to the m i dphase

,

of co ntractio n as represe n ted i n stai n ed preparatio n s If the .

an isotropic s ubstan ce does n o t divide an d move toward the telo

phragma to help form the co n tractio n ban ds as v an G ehu chte n ,

hi mself s upposed how the n is this illu stratio n to be i nter

, , ,

p r et ed ? Two expla n atio n s s uggest themselves n either how , ,

ever wholly satisfactory : 1 ) it may represen t a variatio n of the

,

more regular st rati fi c at i on whe n prese nt o r 2 ) it may be of a ,

fi b er stretched at an early stage of co n tractio n i n s u ch mann er

that the anisotropic s ubstan ce has act u ally become divided .

But q u ite aside from the erro n eo u s ass u mptio n of a strict

spatial correspo n den ce betwee n an an isotropic s ubstan ce an d ‘ ’

the Q disc of the stain ed fi b er e n tirely too great relative i m

-
,

portan ce has bee n ascribed to this an isotropic s u bstan ce i n

relatio n to the phe n ome n o n of co ntractio n In the fi rst place .
,

it may agai n be emphasized that man y n on co n tractile sub -

stan ces both i n organ ic an d organ ic are ani sot rO pi c C on

, ,
.

tractility cann ot be regarded as a fun ction of anisotropy More .

over as R an vier lo n g ago showed ani sotropy itself is a fun ctio n

,
20
,

of the un iform directio n of stress H e showed that i n dia r ubber

.

i n the u s u al co n ditio n is isotropic b ut whe n stretched it becomes

,

an isotropic H e s umm arized the results of hi s st udies on this

“
.

s u bj ect as follows : M on orefri ng ent bodies become birefrin ge n t

by V irt u e of a m odifi c at i on i n their molec ular state witho ut ,

chan ge of compositio n a body becomes birefrin ge n t

whe n its molec ules are orie n ted i n on e an d the same axial
directio n m u scle has the same stru ct u re i n repose ,

exte n sio n an d d uri n g ret ur n to repos e o n ly the

le n gth of the str u ct ures varies i n versio n of striae does n ot
—

exist .

There wo uld seem to be a parall el betwee n the di ffere n t

optical co n ditio n s of the i n dia r ubber i n repose an d exte n sio n
an d the co n tracted an d relaxed ( o r exte n ded m scle fi b
) u e r A .

similar phe n ome n o n of alteration i n optical con d u ct of a homo

g en eo u s s u bstan ce un der stress is e x hibited by glass whe n p ut
50 H . E . JOR D AN

un der press ure ; n ormally glass is isotropic b ut un der press ure it

,

develops temporary an isotropic striae The n otorio u sly feebler .

an isotropy of the co n tracted fi b er may res ul t from a dist urban ce

d uri n g co n tractio n of the more reg ularly orie nted particles of the
exte n ded fiber But the q u estio n still remai n s as to how the
.

st rat i fi c at i o n of the isotropic an d an i sot rO p i c s u bsta n ces of cer

tain fi b ers is to be explai n ed In the fi rst place it sho uld be

.
,

n oted that s u ch st r at i fi c at i o n i n otherwise appare n tly ide n tical

fi b er s may be prese n t i n o n e an d abse n t i n the other It may .

be j u stly claimed that failure to see this st rat i fi c at i on is due i n ,

a certain n umber of i n stan ces to un favorable orie n tatio n of the

,

fi b er un der exami n atio n . Whe n the stage of the microscope is

revolved un der the crossed n icols a certai n n umber of these,

appare n tly i n active fi b ers give evide n ce at a certai n portio n of

the fi el d of clear st rat i fi cat i o n But there still remain a certai n
.

num ber of fi b ers i n the relaxed co n ditio n which whe n Viewed , ,

un der the very best co n ditio n s still are apparen tly i n a ctive
, .

The most favorable co n ditio n for detecti n g the an isotropic striae

are fix at i on i n al c oh ol an d mo un ti n g i n glyceri n e R egardi n g
0
.

the appare n tly optically i n active resid ue of fi b ers it might be

claimed that their i n active co n ditio n is d ue to obliq u e lo n gi
t u di n al strai n s or distortio n s dist urbi n g the origi n al segregatio n
,
.

S u ch distortio n is almost un avoidable with fresh teased material .

Previo u s coagulatio n of the sarcoplasm with alcoholic fix at i on

gives to the an isotropic material a more stable co n ditio n an d a
sharper defin i t i on But it is also q uite probable that fix at i o n of
.

itself bri n gs abo ut a sharper segregatio n than is characteristic of

the n ormal livi n g sarcoplasm G ran ti n g however a more or
.
, ,

less sharp segregatio n of the anisotropic co n ditio n as we are i n ,

fact compelled to do for livi ng fi b ers of i n sect leg m u scle un der

certai n n ormal co n ditio n s the further questio n remai n s as to
,

why those segregatio n s correspo n d i n ge n eral with the limi ts of

the dark (Q ) disc The an swer to this q u estio n follows from the
.

fun dame n tal physical co n ditio n of the Q disc It w as shown — .

above that the actio n of alcoholic fix at i on an d the mechan ical

factors of te n sio n both i n dicated the relatively more fl ui d
n at u re of the s u bsta n ce of the dark disc D ehydratio n was.
52 H . E . JOR D AN

establish the histologic basis upo n which an adeq u ate theory

can be b u ilt The evide n ce disc u ssed above makes it clear
.

that co ntractio n is n ot a simple imbibitio n phe n ome n o n The .

f un dame n tal physicochemical or electrical processes upo n which

co n tractio n directly depe n ds are restricted to the i nt rasarc ost yli c
sarcoplasm How the alteratio n s i n the striatio n s are related
.

to these processes an d how both are fun dame ntally related to

co n tractio n can at prese n t o n ly be s urmised Pre n an t Bo u i n .
, ,

an d M aill ar d
é co n ceive of co n tractio n as an electrocapillary
l

process They describe the process as follows

.

W he n the ele ctri c al poten ti al of opp osite surfa c es o f c on t a ct between

the p arti cles o f the mu sc le fi b er be co m e s m o di fi ed i n so m e w ay as by ,

a n erv ou s sti m u l u s o r by e n er g y liber ate d by a c hem ic al re ac ti o n the ,

f o m o f these surface s ten ds t o be com e m o di fied

r resulting
The stri ated fiber where o c c ur tri ple
,

i n a co n tr acti o n o f the fiber .

,

c on t a c t s betwee n t he d a k d i s c the c le ar d i sc an d the sar c o pl asm is

r , ,

m u ch m o re sen sitive th an the sm o o th fi b e an d i t s d ivi si o n i n t o agg r e

r,

g at i o n s o f v ery sm all p arti c les i n co n se q u en c e o f whi c h it be c o m e s

,

very active fro m the poi n t o f v iew o f c apill ary attr ac ti o n g ive s t o the ,

whole c o n si derable en erg y S i n ce there is a re ci pro city between the

.

su f a c e d ef o rm ati o n s an d ele c tri cal v ari ati o n a si m ple sh o ck ( m e ch an

r ,

i c al stim ulu s) pro du ce s a v ari ati o n i n poten ti al whi ch by pro pag ati on , ,

a l so d eterm i n e s c o n tr ac ti on C o ntr a c ti on app e ars t o be a c co m p an ied

by a sl ow u sag e o f the alb um1 no id sub st an c e s o f m us cle with the form a ,

ti on of creati n b ase s b u t the pri n ci pal chem i c al chan g e c on si sts i n the

,

app e ar an c e o f a l arg e q u an tity o f l a c ti c a c i d whi c h either p arti ally or

,

t ot ally be com es con verted i n t o c arb onic anhydri de an d water Thi s .

l a cti c ac i d f orm s to the detrim en t o f mu scu l ar g ly c o g en an d espe ci ally

t o the g l u cose c o n ti n u ally su pplie d by the bl o o d ; the sug a r of the bl o o d
is then the p i m ary s our ce o f the en ergy b ut duri n g the wo k o f mu scle
r ,
r

c o n tr acti o n ,the reserve hydro c arb on s of the m uscle ti ssu e pl ay a

se c on d ary 6l e ( p
r .

The o utstan di n g morphologic mark of co n tractio n is the con

tractio n b an d T hi s str u ct u re is composed essen tially of f used
.

opposite halves of s u ccessive dark discs an d a bisecti n g telo

phragma The i n termediate phases whe n the Q disc is bisected
.
-

by a wide n i n g H disc has un til n ow bee n most diffic ult to explai n

-
.

The u s u al i n terpretatio n of these latter phases n amely as the , ,

res ult of a stretchi n g of the sarcostyle has led to m u ch co n f usio n ,

.

This i nterpretatio n has appare n tly adeq u ate observatio n al sup

port for sarcostyles with divided Q discs that is sarcostyles a t
,
-
,
 STR IPE D M US C LE OF WA SP 53

m i dph ases of co n tractio n freq u e n tly show relatively elo n gated

,

sarcomeres It m u st agai n be emphasized that a co n ditio n of

.

stretchi n g may be s uperimposed o n sarcostyles at any f un ctio n al

stage This fact m ust b e kept i n mi n d i n i nterpreti n g the
.
.

str u ct u re an d le n gth of sarcomeres of an y partic ular sarcostyle

or fib er The e ffect of stretchi n g is first shown i n an elo n gatio n
.

of Q If the sarcostyle is passi n g i n to co n tractio n that is to say

.
, ,

whe n it is i n a co n ditio n where a median (H ) disc is prese n t t h e ,

stretchi n g e ffect shows itself largely i n a wide n i n g of the median

disc that is cau si n g a wide separatio n of the two portio n s of Q
, ,
.

This res ult has led to the ge n eral opi n io n that stretchi n g cau ses
a separatio n of Q alo n g the mesophragma with the appearan ce
of an H disc This is the basis also for the i n terpretatio n of a
-
.

fi b er with an H disc bisecti n g the Q disc as o n e i n exte n sio n

- -
.

However the evide n ce is fairly complete that the relaxed c o n

,

ditio n of the fi b er is characterized by an un divided Q disc -

.

S uperpositio n of stretchi n g u po n a relaxed co n ditio n cau ses

c hi efl y a le n gthe n i n g of Q The prese n ce of an H disc i n dicates
.
-

n o t exte n sio n n or stretc h in g primarily b ut the i n termediate c o n ,

tractio n phases Stretchi n g of a sarcostyle i n this co n ditio n

.

prod u ces chi efly a len gthe n i n g of the H disc -

.

The unm odi fi ed fully co n tracted fi b er has a sharp lateral

co nto u r (fi g s 8 33 an d Beadi n g is n ot an i n dex of c on
.
, ,

tractio n as claimed by R an vier Schaefer Meigs M eD ou

, ,
20
,
28
,
15

gall,
an d others
1 3 ' 14
Co n tractio n is accordi n gly n o t simply a
.
‘

matter of the imbibitio n of fl uid by the sarcostyle It is a .

matter associ ated W ith the divisio n of the deeply colored sub
stan ce of the dark disc an d its s u bseq u e n t movemen t agai n st
the telophragma res ulti n g i n the prod uctio n of co n tractio n
,

ban ds .

M eigs has made o ne of the most co n siste n t attempts to

15

s upport by morphologic data the imbibitio n hyp othesis of mu scle

co n tractio n esse n tially as proposed by M c D oug all M eigs .

s ummarizes the res ults of his st u dy of the m u scle of the fly an d

of the frog by stati n g a series of obj ectio n s agai n st the opposin g
hypothesis n amely that co n tractio n res ults from fun damen tal
,

i n tr asarc o st yli c reactio n s

“
,

Meigs says that The latter hyp oph

.
54 H . E . JOR D AN

esis disregards almost all the facts that are k n own co n cer n i ng
m u scle ” . O n the co n trary it may be claimed that the hypo
,

thesis i n cl uded all the facts tha t are n ot artifacts " Meigs states
that the latter hyp othesis leaves u n explai n ed the divisio n of
the m u scle s ubstan ce i n to mi n ute sarcostyles for it is impossible ,

t o see wh y larger bodies of co n tracti l e s ubstan ce sho u ld n o t

‘ ’

perform their f un ctio n as well as smaller o n es ” To this the .

reply may j u stly be made that a divisio n of a fi b er i n to fi b rils

each s ur ro un d ed by semi fl ui d ex t rafi brillar sarcoplasm provides
-

a very m u ch more effi ci ent perhaps i n dispe n sable method of

, ,

providin g n utritive materials to a metabolically so active tissu e

“
.

Meigs claims that the n on imbibitio n hypothesis also leaves

-

u n explai n ed t h e divisio n of the sarcostyles i n to sarcomeres an d ,

the prese n ce of the Z and M membran es an d co n tradicts the -

,

well k n own fact that the Z membran e is more or less i n ex

”
- -

te n sible But the prese n ce of the Z an d M membran es whose

.
-
,

prese n ce imposes upo n the sarcostyles a divisio n i nto sarco

meres can be readily an d reaso n ably explai n ed as paths al o n g
,

whi ch travel the materials ( assimi lative and disassimi lative) of

metabolism Their i ntimate attachmen t to the sarcostyles and
.

the sarcolemm a speaks stro n gly i n favor of this i n terpretatio n

“
.

Meigs co n ti nues : It leaves un explai n ed the di ffere n ces i n ap

p earan c e betwee n the relaxed and co n tracted sarcostyles and is ,

f orced to ass u me that the appeara n ce of b ul g i n g s i n t he c on

tracted sarcostyles is a del usio n an d that the appeara n ce of the,

heavy lin es betwee n the b ulged areas is the res ult of the pro
d uctio n of a large amo un t of some n ew s ubstan ce withi n the
sarcostyles ” However the hypothesis which Meigs criticises
.
,

d oes n ot as u me that the appearan ce of b ulg i n g s ( beads ) i n

c o n tracted sarcostyles is a del u sio n It claims that they are .

artifacts The ill ustratio n of a co ntracted fi b er p ublished by

.

Meigs (fi g 8 ) does n ot act u ally show the b ulg i n g s whi ch he

.

attempts to impose upo n it i n his e fforts to s upport M c D oug all s ’

imbibitio n theory The heavy li n es spoke n of as betwee n the

.
‘ ’

‘
b ulged areas of his fi g ure are the co n tractio n ban ds formed by
’

the acc umul atio n of the darker s ubstan ce of the Q disc of the -

relaxed co n ditio n n ot the res ult of the prod uctio n of some n ew

,
‘
 STRIPE D M U S CL E O F WA SP 55

s ubstan ce withi n the sarcos tyles Fi n ally all of the defi ni t e

’
.
,

mo rphologic data are i n direct co n tradictio n of the facts upo n ‘ ’

whi ch the imbibitio n hypothesis is based In simplest terms .

,

as adopted by M cD oug all an d s upported by M eigs the hypoth ,

esis regards co n tractio n an d relaxatio n as phe n ome n a directly

comparable to the res ults of th e actio n of hypoto n ic an d hyper
to nic sol utio n s respectively O n the co n trary the evide n ce
,
.
,

stro n gly in dicates that co n tractio n depe n ds upo n (or is at least

a ssociated with ) esse n tially i n tr asa r co st yli c moveme n ts of the

dark s ubstan ces ( co n sisti n g i n part at least of chlorides phos ,

p h at es an d potassium salts) from the middle to the e n ds of the

sarcomeres that is from the mesop hr agma to the telophragmata
, , .

T hi s movemen t is probably associated with s urface te n sio n or -

,

e lectrocap ill ary ,

reactio n s amon g the ult rami cros0 0 p i c particles
of the i nt rafibri ll ar sarcoplasm .

Most previous i nvestigators have ass um ed that the s ubstan ce

of the light disc is relatively more fl ui d than that of the dark
disc an d that the i nt rasarco st yli c movemen t of fl ui ds i n con
,

traction is from the light disc toward the dark disc that is from ,

Z to M . T hi s is the View of E n glemann and of Schaefer But .

it was shown above that the res ults of the actio n of mechan ical
an d osmotic factors demo n strate the relatively more fl ui d n at u re

( f un dame n tally ,
that is aside from
,
the prese n ce of gra nu les i n

s u spen sion) of the dark disc An d the histologic preparatio n s

.

actu all y demo nstrate as shown by the stai n i n g res ults that a
, ,

deeply stai ni n g s ubstan ce passes from Q to J from M to Z ,

.

Moreover if a hyali n e s u bstan ce passed i n the opposite di rec

,
‘ ’

tio n as ass umed by Schaefer thu s dilu tin g the dark s u bstan ce
, ,

of the sarco u s eleme n ts and so cau si n g it to stai n less deeply

( th u s givi n g the ill u sio n of a reversal of striatio n ) the loss of ,

the deeply stainin g property sho uld appear first alo n g the
termi n al borders of the Q disc rather than alo n g the midli n e as
-
,

is act u ally the case .

f I n terca la ted
. di scs

The foregoin g bears directly upo n the qu estio n of the si g ni fi

can ce of the i ntercalated discs In a number of earlier p ap er s
.
s
56 H . E . JOR D AN

on this s ubj ec t I have prese n ted evide n ce i n s upport of my

hypothesis that these discs are esse ntially m odi fi ed irreversible
co n tractio n ban ds The simplest type of i ntercalated disc is
.

practically ide n tical i n str uct ure an d stain i n g reactio n with a

co n tractio n ban d The esse n tial do u ble n at ure of the c on
.

tractio n ban d s upplies the explan atio n of the several varieties of

the simplest type of i n tercalated disc with respect of their r e
latio n to the telophr agma Si n ce the two halves of a co n tractio n
.

ban d have a b i sarcom eri c origi n an d i n co n sequ e n ce a relatively

,

i n depen de n t relatio n ship it wo uld seem to follow that o n e half

,

might pass i n to relaxatio n whi le the other half remai n ed i n

capable of leavi n g the telophragma the latter half th u s b e ,

comi n g an i n tercalated disc of the variety bo un ded o n ly on o n e

side by a telophragma W h ere the e n tire co n tractio n ban d
.

failed to reverse the variety of i n tercalated disc which is bisected

,

by a telophragma wo uld take origi n The variety of disc bo un ded .

o n both sides by a telophragma co u ld arise by the s u bs q u e n t

e
f usio n of the opposite halves of two adj ace n t co n tractio n ban ds .

This process of fu sio n may act u ally co n sist essen tially of a

‘ ’

seco n dary m o di fi cat i o n of those portio n s of the sarcostyles i n

t er v en i n g betwee n the opposite irreversible halves of adj a ce n t
co n tractio n ban ds .

Besides the other more complex types of discs ( step an d

serrated forms ) still an other typ e of i n tercalated disc m u st here
,

be co n sidered This i s a relatively rare type i n c ardi ac m u scle

. .

It occ urs somewhat more frequ en tly i n the specimen of hu man leg
mu scle where it is scattered amo n g the predomi n ati n g simplest
,
9

variety with bisecti n g tel ophragma In a paper by J ordan an d

'

Banks o n the i n tercalated discs of the beef heart this type

11
,

was i n cl uded amo n g the ill u stratio n s (fi g b ut its i n t erp re .

t at i o n w as at that time n o t clear an d n o descriptio n was at ,

tempted It may be described n ow as esse n tially a thicke n ed

.

telophragma or portio n of a thicke n ed telophra gma It wo uld

,
.

seem that a co n tractio n ban d may disappear or reverse at the ,

time the fi b er passes i nto repose either as a whole or o n ly i n ,

half or fi n ally o n ly i n part In the latter circ umstan ce a mere

,
.

remn an t of the deeply stai n in g s ubstan ce of the composite con

 STR IPE D M US C LE OF WA SP 57

tractio n ban d might co nti n u e to adhere to a portio n of a cer

tai n telop hr agma i n an otherwise relaxed fi b er an d so i nitiate ,

thi s fo u rth disti n ct typ e oi i n tercalated disc Thro u gh sub se

-

q u e n t m o di fi c at i on by the pe n etratio n of tiss u e fl ui d Via the

telophragma s u ch a thicke n ed portio n of this membran e might
,

be c a u sed to persist as this typ e of disc .

VIII . S UMM A R Y

1 Co n tractio n i n striped mu scle is associated with a ge nuin e

.
-

reversal of striatio n s as regards a deeply stain in g s ubstan ce of

the dark disc of the sarcostyle T hi s reversal of striation s .

res ults i n the formatio n of co n tractio n ban ds i n the co ntracted

fi b er .

2 . co n tractio n ban d is composed essen tially of the fu sed

A /

opposite halves of two adj ace n t dark discs In fi b er s co ntai ni n g .

accessory discs the co n tractio n ban ds i n volve also two of these

,

discs an d i n fi b er s where sarcosomes occ ur ; some of these gran

,

u les may also become i n cl u ded wit hi n the i n t er sar c ost yl i c spaces

of the defi n i t i v e co n tractio n ban d an d co n trib u te to its deeper

color an d i n ten ser stai ni n g reactio n The method of formatio n.

of the co n tractio n ban d ex plai n s its bisectio n by the telophragma .

3 The stripi n g of the striated m u scle fiber res ults from the
.

segregatio n of darker an d lighter ( chr omatic an d achromatic )

s ubstan ces i n alter n ati n g dark an d light discs These discs are .

bisected by a mesoph ragma an d a telophr agma respectively ,

.

The deeper staini n g disc is n ot coexte n sive with n or the res ult ,

of the prese n ce an d st r at i fi ed distrib utio n o i sp eci fi c an isotropic ,

materials The darker appearan ce an d deeper stai ni n g reaction

.

of the dark disc an d of the co n traction ban d may be d ue i n ,

part at least to the segregatio n ( demo n strated by Me n ten ) of

,

chlorides phosphates an d potassi um salts i n the dark disc i n

, ,

the relaxed sarcomeres an d i n the co n tractio n ban d i n the c on

tracted sarcomeres .

4 A beaded co n ditio n of the sarcostyle is n o t an i n dex of a

.

phase of co n tractio n n or is it the res ult of co n traction It is

,
.
58 H . E . JOR D AN

an artifact due either to the dehydrati n g actio n of certai n

,

fi x i ng reage n ts like alcohol u po n the relatively more fl u id dark

discs or to the e n dosmotic action of hypoto n ic sol utio n s In the .

former case the co n striction separatin g s u ccessive beads is at ‘ ’

the level of the dark disc ( mesophragma) i n the latter at the ,

level of the telophr agma .

5 The co n ditio n of bisectio n of the dark disc by a median

.

disc is i dex of of stretchi n g as clai med

'

( H ) n o t an n ex t en si o n o r ,

by R oll et an d by Schaefer b ut of i n termediate phases of c on

‘

tractio n E rror i n the i nt erpretatio n of thi s co n ditio n has

.

res ulted thro u gh disregard of the fact that a co n tractin g fi b er

may be q u ite as readily stretched as a relaxed or a co n tracted on e .

Stretc hi n g produces its primary e ffect i n the re g io n of the dark

disc causin g a len gtheni n g of thi s disc A t the begi nn i n g of
.

con tractio n stretchi n g seems to produ ce the median disc whi le ,

i n reality it simply bri n gs i n to clearer View thro u gh exte n sio n , ,

an extremely thi n media n disc already prese n t as an a ecom

p ani am ent of the i n itial phase of co n tractio n These co n sider .

at i on s e x plai n the appare n tly parado x ical co n ditio n of occasio n al

greater le n gth of sarcomere i n a co n tractin g sarcostyle as com

pared with the sarcomer e of a relaxed sarcostyle Stretchi n g .

of a fi b er at the beginni n g of co ntractio n prod uces the ill u sio n

of the prod uctio n of an H disc by te n sio n
-
.

6 The segregatio n of ani sotropic materials ( co n ditio n s ) i n

.

strata altern atin g with isotropic levels an d correspo n din g more

,

or less sharply un der certai n co n ditio n s with the dark disc

, , ,

wo uld seem to fin d its explan atio n i n the relatively more fl u id

co n sisten cy of these dark discs I n terpreti n g the co n ditio n of
.

isotropy i n terms of si mi lar orie ntation of the sarcoplasmic

particles to li n es of stress (followin g R an vier s explan atio n) it ’
,

becomes appare n t why i n a str uct u re like that of a f un ctio n i n g

sarcostyle where the particles s uff er co n tin u al rearran geme n t
—

d uri n g co ntractio n an d relaxatio n an an isotropic arran geme n t

—

co uld be more readily ass umed i n the relatively more fl ui d semi

solid dark disc A lo n g the same lin e follows the explan atio n for
.

the more feebly an isotropic character of the co ntracted fib er i n ,

which the origin al orie ntatio n of the particles of the relaxed

60 H . E . JOR D AN
LITERATURE C ITED
1 B RU C KE ,
E . 1 858 U n t e r su c h un g en u b er d en Ba u d er M u sk e l f a se rn m i t
H ilf e d er p o l a r s e r t en ii Li c h t es . Wi en e r D en k s c h r m a t . .
-n a t . K la sse ,

Bd 1 5, S 69 . . .

2 B U LL A R D ,
H H . . 1 9 16 O n t he o c c u r ren ce an d phy i l gi s o o ca l s i g n ifi c an c e o f

f at in t he n o rm a l m y o c ar diu m an d a t r o - i v t i l en r c u ar s y st e m ( b un d le
of H i s) , in t er s t it i al g ran u l es ( m i t o c h ro n d r i a ) an d p h O S p h o l i p i n es i n
ca r di ac m u sc l e . Am . Jour . An at .
,
vo l . 19, p . 1 .

3 D A H L GREN ,
U .
,
A ND K N R EP E ,
W . A . 1 903 P i ip lr nc es o f an im al h i st o l o gy ,

p 85 .Ma c m ll an . i .

4 ENG L E M A NN , T W 1 893 . . Ub er d en U p rs r un gd er M u sk e l k r a f t . W . E gl n e

m ann , L i pz i g ( i t d f m H id h i ) e c e ro e en a n .

HEI D EN H A I N M 1 9 1 1 P l m d Z ll
,
G Fi h J . as a un e e . . sc er , en a .

H LM G R E 1 90 7 U b d i S k pl m k
O EN , . q g t if t M k l e er e ar o as a o rn er u er es re er us e

f a sern A t A Bd 3 1 S 60 9 . na . nz .
,
.
,
. .

7 J OR D A N H E 1 9 1 6 Th m i
,
. pi t t . f th l g m l f th e c r o sco c s ru c u r e o e e u sc e o e

p id A p l d t yl l t
sea - S A t R vo l 1 0 p 493
er, no o ac us en u s . na . ec .
,
.
,
. .

1 9 1 7 Th m i pi t t f t ip d m e l i L im l P b c r o sco c s ru c u r e o s r e u sc e n u us . u .

2 51 C g i I t i t t i f W h i gt pp 2 73 290
,
a rn e e ns u on o as n on , .
—
.

1 9 1 9 S t di t ip d m l t t uIV I t l t d di i
es o n s r e u sc e s ru c u re . . n erca a e scs n

v l t y t p d m l A t R vo l 1 6 p 2 03
o un a r s r1 e u sc e . na . ec .
,
.
,
. .

19 1 9 S t di t ip d m l t t u V Th mp t i v hi
es o n s r e u sc e s ru c u re . . e co ar a e s

t l gy f t h l g
o o d wi g m l f th m
o t i with p i l f e e an n usc e o e an s, s ec a r e er en c e

t t h N di
o d th e m Am J A t vo l 1 6 p 21 7
- sc an e s a r co so es . . o ur . na .
,
.
,
. .

J OR D A N H E A N D B A N K S J B 1 9 1 7 A t dy f t h i t
,
. .
,
l t d di ,
. s u o e n er c a a e sc s

f th h
o t f th b f e Am J A t vo l 22 p 285
ea r o e ee . . our . na .
,
.
,
. .

M A C A U M A B 1905 O t h di t ib t i
LL ,
. f p t i mi im l d
. n e s r u on o o ass u n an a an

v g t b l ll J
e e a P h y i l vo l 32 p 95 e ce s . ou r . s o .
,
.
,
. .

MC D O U G A L L W ,
. 1 897 O n t he s t ru c t u re of c ro ss st r a t e i d mu sc l e an d a

su gg t i es on as t o t he n a t u re of its c o n t ra c t o n i . J ou r . A n at . an d

Phy i s ol .
,
vo l 3 1 , p 41 0
. . .

1 898 A t h eo r of mu scu l a ry co n t ra c t o n i . Am . J ou r . A n at . an d

Phy i s ol .
,
vo l
32 , p 1 8 7 . . .

ME I GS E B , . . 1 9 08 Th e s t r u c t u re of t he e l em en t o f c r o ss- s t r a t e i d mu s c l e,
an d t he c h an g es of f or m wh i c h i t un d g er o es du rin g c o n t ra c t o n i .

Z e i t sc h r if t f ur A ll g em e n e i Phy i s o lo gi e, Ed 8 , S 8 1 . . .

M E N TE N M A UD , L . 1 908 Th e di t i b u t i
s r on of fa t , c hl o r id es , ph ph t os a es,

p o t a ss u m i an d i ro n in st r a e i t d mu sc l e . T r an s . C an a di I t i t t
an ns u e,

vo l . 8 , p 403 . .

M E RK E L ,
F 1 8 7 2 D er
. q g ue r est re if t e M u sk e l . I . D as p i m i t iv
r e Mus k el
e l em e n t d er A rt h r o p d o en . A rch . f m ikr
. . A nat .
,
Ed 8 , S 2 44. . .

1 8 73 D er q u er est re g if t e M u sk e l . II . D er C o n t ra c t i o n s -
v g g
or an im
p o l a r s e rt en ii Li cht e . S c h ul t z e s
’
A r ch iv ,
Ed 9 , S 293 . . .

P REN A NT ,
A .
,
BOUI N , P .
,
ET M A I L L AR D ,
L . 1 9 04 T it é d H ira
’
st o l o g i e , T 1, .

p . 440 .
 S TR IPE D MU S CL E OF WA SP 61

R A N V I ER , L . 1 88 0 L e go n s d An a t o m i e
’
G en era l su r le Sy st em e Mu scu l a i re .

P i ar s .

R E TZ I U s , G . 1 8 90 M u sk elfib r i l lae -
un d S k pl ar o as ma . Bi ol . U n t er s . v on

R tzi
e us .

R LL
O ET , A . 1 885 U n t e rs u c h u n g en u b er d en Ba u d er q g
u e r es t r e if t en Mu s
k el f a se r n . I und Wi n
II . e .

1 89 1 Ub e r d ie N S t e if n ( N b en h i b
- r e e sc e en ) das S k pl ar o a s m a un d die
Kon t ra c t i o n d er qu g t if t en Mu k l f
er es r e s e a se rn . A h f rc . . m ik r . A n at .
,

Bd 37 S 654
. . .

1 89 1 U n t er su ch u n g en u b er K o n t ra kt i on un d D pp lb
o e re ch un g de r
q g
u e r e st r e ift en M u skel fa sern . Wi en .

Ub
1 8 92 e er die K on t ra kt i o n s w e l le n u n d ih re B zi
e eh u n g zu d e r En t
z k gb uc un ei d en q u er est re g if t en Mu kel fa s s e rn . P fi u g er
’
s A r ch . Bd 54
,
. .

RU H RF RD W
T E O ,
. 1 89 7 O n t he s t r u c t u r e an d c o n t r a c t o n o f s t r i ip e d m us

c u l ar fib e r vo l 3 1 p 309
. Jour . An a t . an d Phy i s ol .
, .
,
. .

S C H A E F ER E A 1 91 0 E en t i al f h i t l o gy L n g m an G n
,
. . C ss s o s o . o s, r ee o .

1 9 1 2 T xt b k of m i p i n t my L n gm n G n C
e -
oo c r osco c a a o . o a s, r ee o .

T H U LI N I 1 9 1 5 I t di G un dm mb n in k on t n t v k mm n d Bi l
,
. s e r e ra e e s a or o e e

d un g i n d n que g t eif t n Mu k l f n ? A h f m ik An t
e r es r e s e a ser rc . . r . a .
,

Bd 86, S 3 1 8
. . .

TO U R N E U X , F . 1 8 92 S u r l e s m o d i fi c at i o n s st ru c t u r a l es qu e pé r sen t en t le s
m u sc l es j au n e s du D yt i q ue p d en an t la c o n t ra c t o n i . Jo u r . A n at . et

P yi
h s ol . n o rm . et p at h .
,
An n é 28e, .

VA N G E H U CH TE N ,
A . 1 8 86 Et u d e su r la st ru c t u r e int ime d e la cell u l e

m u s c u l a i r e st r i é e . La C el l u l e ,
T 2 , p 293
. . .

1 888 Et u d e su r la st ru c t u r e i in t m e de la c ell u l e mu sc ul a i re st r ié e

ch e z l es V téb é er r s . La C e llu l e ,
T 4 , p 247
. . .
62 H . E . JOR D AN

PL A TE 1

E X P L A N A TI ON OF F I G UR E S
L eg m u sc l e of w as p
( U nl ess o t h e rw s e i s p ifi d
ec e all su b quse en t d r aw n i g s ar e f
m t ssu e fi x e dro i
i n 9 5 p er c en t al c o h o l , s t a n e i d w ith i ro n -he m at o xy l i n an d m ag n i fi ed a o u t 1 300 b
di am e t er s .
)
14 P o rt o n of i l on g i t di u n al sec t o n o f a i fi b er , i n t h e rel a xd
e c on di t i o n , s h ow

i ng t h e Q an d id f t h t l p h gm ( Z ) t h N di
J- di sc s , an d on e ith er s e o e e o ra a e - sc s .

1 5 F ib in ly t g erf nt ti n e ar h w i n g t h t mi n l kn b b d
s a e o co r ac o ,
s o e er a o e c on

di t i n f t h Q gm n t f t h my fi b i l i n di t i n g p
o o e -
se e g f d p ly t in i n g
s o e o r s, ca a assa e o ee s a

sub t n s a t w d t h t l p h gm t n d t h pp
ce o ar en f t h bi
e o t i n g H dira a a a e a ear a ce o e se c -
sc .

16 F b in th 1l x d er n di t i n n l y l i ght l y
e re a t in d h win g th
e xi lco o ,
o s a e , s o e a a

a rr a n g m nt f th l ng
e e lu m n fo lti l y m l l nu l i
e o co s o re a ve s a c e .

17 T v ti n
r an s f fib e h wing
er se nt
se c l nu l u o th l m ll
o r,
“
s o a ce ra c e s, e a e ar

tyl nd t h nv l p i g l mm
‘ ’
ch t f th
ar ac er o e s ar c os es a e e e o n s ar c o e a .

1 8 F ib fi d i n F l mm i n g flu i d n d d t i n d t p i t wh t h Q di ’
er x e e s a es a e o a o n er e e -
sc

is n l ng
o n p i u us t
o er c osh w t h m ll
s ph i l J
c o m, n ith
o o e s a er s er c a -
sar c os o es o e er

sid f t h t l p h gm n d t h l g v l Q
e o e e o ra m a, a e ar e o a -
s ar c o s o es .

1 9 F ib t l t t g f
er a nt t i n i n w h i h pp
a a e s a it h lv f u e o iv co r ac o ,
c o os e a es o s c c ess e

Q di-
(sc s g d t h i d p l y t in i n g ub t
as r e ar s ) h v fu d w i t h
e r ee h th s a s s an c e a e se e ac o er ,

an d w i t h t h i n t v n i n g N di e g i n t t h t l p h gm t t f m n t t i n
er e - sc s , a a s e e o ra a a o or co r ac o

b an d s .

20 F u l l y nt t d fib e h ow i n g
co v n t ti n b nd
r ac e r, s se e c o n r ac o a s .

Wi n g m u s l c e

21 P or t on o f i l on g i t di u n a l se c ti n f fio dj o ve a acen t sar c os t yl es w ith si x g ro u p s

o f sar c o so m es , an d o n e n uc l eu s (n u ) . T h e t el o h r a m a t a p g of t he se v er al sar c o

st yl es ar e at t he s am e le v e l s, i n di c at n i g i nt er sar c os t y l i c c on t nu i ity . Fl emm n i g

fix at i o n ; l ht l ig y s t ai n e d .

22 Tw o a dj ac en t sar c os t yl es sh ow n i g a c on t nu i ity of p
t h e t el o h r a m a t a , i n g
t he re gi o n n ear t he m idd l e w h er e t h e fib r i l s h a v b n
e ee on l y v y l i gh t l y d
er s r aw n

a p ar t . F l mm i g fi e n x at i o n .

23 F i v dj t e a ac e n s ar c o st yle s w th i f u xt o r e en s iv g e p
ro u s of i nt v n i n g
er e s arc o

s om es . O n t h e sar c ost l es ar e see o e c o a rser , y n b th th d p ee er s a t i n i n g t l ph e o r ag

m at a an d t h e m o re el i c at e , d
a n t er m es o p h r ag ma t a f i . Th e n a rr o w er i n t er sar
c o st yl i c s p ac es c o n t a n s n i i gl e r o w s o f s ar c o so m es ; t h e w er , id d ub l w T h
o e ro s . e

l on g xi a s of t he s a r c o s o m es o f t h es e t w o g p
ro u s ar e i n g
e n er al p l d e p t iv ly
ace r s ec e

at ri gh t an gl es an d p arall el t o t h e l on g xi a s of t he sar cos t y l es F l mm i n g . e

fix at i o n . X 2 600 .

24 Th r ee c o mm o n f o r m s o f s ar c o s om es a s seen i n l on g i t di
u n al sec t on s o f i t he
w i n g mu sc l e . X 2 600 .
IPE D MU S C L E O F WA SP P L ATE 1
H . E J O RDA N

m:
wisttfigzfliu
l
 S TRIPE D M US C LE OF WA SP 65

PL A TE 2

E XP L A N A TI ON OF F I G UR ES

25 F r esh r e l a xd e sar c o s t yl e as s een in Ri g n er s

’
or T i o so n s
’
so l u t o n s i . T he
p
t el o h r a m a t a g ar e c o n s pi cu ou s, t he d k di ar scs a r e t hi k c , t he l i gh t di sc s v y er

th i n .

26 Rl xd e a e i g fl u id d l i gh t l y t i n d w i t h
s ar c o st yl e fix ed in Fl emm n
’
s an s a e

i n ro - h m t x yl i
n e hn i
a o u nly v . T hi t
y l i gh t m d i fi t i n s m ec c ca s es o er s o ca o as c o

p d w i t h t h f h fib i l i n R i n g
ar e lut i n
e r es r er s so
’
o .

27 R l x d t yl fi e ad i n 95 p e nt l h l nd t i n d w i th i n
sar c os e x e er ce a co o a s a e ro

h m t xyl i n
e a o T h fib i l wh l i .h u k n ; t h d k die l i g
r tly as a o e s s r n e e ar sc a so s re a

sh u n k n d u t d h yd t i n
r e ,
nd t in d p ly
e o e ra o , a s a s ee .

28 R l x d ty l fi d i n
e a 10 pe t f m l in
s ar c os l u t i n n d l i gh t l ye x e a er c en or a so o a

t i
s a ne d w i t h i n h m t x y l i n B t h t h t l p h gm t ro — d t h f int
e a o m . o e e o ra a a an e a er eso

p h gm tra v i i b l T h fib i l pp
a a ar e w ll n t th
s l v l f th
e . t l e r a e ar s s o e a e e e s o e e o

p h gm tra T h i pp n t w ll i n g i du t g t h i n k g i n t h g i n f
a a . s a ar e s e s e o rea er s r a e e re o s o

th m e p h gm t eso nd l t iv ly g ra t i g idi t y f t h t l p h gm t
a a a a re a e r e a er r o e e o ra a a .

29 F h l x d r est yl n f t b i f t t m n t w i t h di t i ll d
re a e sar c o s e as see a er r e r ea e s e

w t a er T h fib i l b. m b d d n d l i gh t l y h t n d du t t h
e r ec o l t iv es ea e , a s s or e e , e o e re a e

ri g idi t y f t h t l p h gm t n d t h o d m i uff d by t h
e e o ra m a a a e en os os s s ere e s ar c o e re s .

T h Q di e i l t iv ly thin
-
sc A f t p l n g d i mm i n i n di t i l l d w t t h
s re a e . er ro o e e rs o s e a er e

Q di-
pp sc t fill t h
a nt i m
e ar s d u t i t di l u t i
o nd th e e r e sar c o ere e o s on , a e s ar c o

m e r es v n tu l l y up tu
e Th ph n m n d m n t t t h p n f
e a r re . ese e o e a e o s ra e e r ese ce o a

p er i sar c o st yl i c m em b r an e .

30 ar c ost l e m S y i p f
y t in d f m p p t i n di n g t R l l t
er e c t l s a e ,
ro a re ar a o a ccor o o e
’
s

m th d
e T h i fib i l h
o . su ff d t h m m difi t i n
s r n p l d i n di t i l l d
as er e e sa e o ca o as o e ac e s e

w t a er o r o th hyp t n i l u t i n b u t i n x gg
er t d d go o du t t h c soti n o , e a er a e e ree e o e ac o

o f th f mi e id or c ac .

31 R t in g t yl es p p ly t in d f m t h m p p t i
sar c os e, ro er s a e ,
ro e sa e re ar a on as

fi gu 30
re T h i fib i l h m v b n l i ght l y m p
. s r d u d th v a s, or eo er , ee s co r esse n er e co er

gl as s It i . i m i l t t h fi b i l fi g u 2 9 n d 30 ; l t S h f
s s ar o fib i l A
e r s, res a a so o c ae er s
’
r ,

fi gu 6 re nd p nt ,
a t yl t i fi i l ly
re rese nt t d th t i w ll n nd
s a sar c os e ar c a co rac e , a s, s o e a

co n qu n t ly h t n d t h u gh t h m t i t i n ( n d m i ) f t h h y p
se e s or e e ,
ro e os o c ac o e os os s o e o

t nio qu u f m i id l u t i n m p l y d i n R l l t t hn i
c a eo s or c ac-
so o e o e o e
’
s ec c .

32 S t yl t m wh t l t
ar c o s t g f n d m i t h n t h t f fi gu
e a so e 31 a a er s a e o e os os s a a o re ,

in wh i h t h d k c Q ub t n h b e ar m di l u t d t
er n xt n t w h i t l i ght l y
—
s s a ce as ec o e e o a e e ere s

di l t h n t i b d d fib i l T h l t t h fi gu d m n t t t h t t h
sc o o r s e e re ea e r . e as r ee r es e o s ra e a e

t l p h gm
e o wh i lra l t i v l y i n x t n ib l m m b n
a, h
e a re a l i ght m u n t f e e e s e e ra e, as a s a o o

l t i ity
e as C mp i n f fi gu
c . 25 n d 2 6 w i t h fi gu
o ar so 29 n d 32 h w t h t o re s a r es a s o s a

th b e d d n di t i n f t h
ea e co tyl p n t n t if t o o e sar c os e re r e se s a ar ac .

33 C n t t d t yl
o f m f h p p t i n i n R in g
rac e sar c o s lu t i n e ro a r es re ar a o er s
’
so o

( mpco w i t h fi g 8 20
ar e nd T he nt t i n b nd
s . pp d u b l Th
, ,
a co r ac o a s a e ar o e . e

t l p h gm t
e o ran t n pi u u p b b l y i n n qu
a a ar e o f th i cot t h d s c o s, ro a co se e n ce o e r s re c e

co n di t i n i n t h nt o t d th t i di l l y w id n d
e co t yl
ra c e ,
a s ra a e e ,
s ar c os e .

34 S tyl f y d l t (Al u u l t u ) t n
ar c os e o ly ph
e e f n e a er a s oc a s a a e ar ase o co

t ti n
r ac Th o m . i n t i t d in th
e s ar c o g i n f t h H di i n n
er e s co s r c e e re o o e -
sc co se

qu n e f t h d h yd t i n g
ce o ti n f th e l h li fi
e ti n raI n h m t xy l i n ac o o e a co o c x a o . ro -
e a o

t i
s a n .
 H . E . JOR D AN

35 . S a rc o s t yl e o f el at er, fix ed i n 95 p e r c en t al c o h o l an d s t a n e i d wi th i ron

h em a t o x y l m , s h ow i g
n a g rea t e r d g e r ee o f c o n s t r i ti n c o an d b ding
ea du e t o al c o

i d yd i
h ol c eh ra t o n T h fi b i l lm t . k at t h l v l f t h
ese r s a os i v i bl y b
n ar a r ea e e e o e

t l ph
e o gm t T h n t i t i n i t h g i
ra a a . f th Q d
e co u d th n
s r c o n e re on o e -
l sc s n er es e c o

d tii w uld
ons m t i di t o g t dgsee f l t i v flu i d t y h
o n th i
ca e a r ea er e r ee o re a e i er e an n

th J di ; n d t h b ul g i n g t t h l v l f t h t l p h g m t d m n t t
sc s a e a e e o e e o a a o s r a es a
-
e e ra e

l tiv ly g
re a t i g idi t y n t h p
e re a t f th r m mb o e ar o ese e ra n es .

36 S tyl f l t
ar c o s t l t ph e f nt
o ti n
e a er ,h wing p i l ly a a er ase o co r ac o ,
s o ,
es ec a

w ll t t h u pp
e a en d t h u l t i m t fib i ll
e er l m n t (m t fib i l l ) f w h i h
,
e a e r ar e e e s e a r ae o c

th se tyl i
ar c os mp d F i x d i n 95 p
e s co t l h l t i n d w i th i n
o se . e er c en a co o , s a e ro

h m t xy l i n
e a o X 1 300 . .

37 S tyl f w p w i n g mu l i l u di g t h m l x d
’
ar c o s e o as in s sc e nc n r e e s ar c o er es , re a e

co n di t i n Th d ply t i i g Q d i
o . i u nu u ll y l
e ee g f m t i l fi d in
s a n n -
sc s s a on or a er a x e

l h l p
a co ib ly du t
o ,
t t h d
o ss di t i n f t h fib i l 9 5 p
e o a s re c nt l h l e c on o o e r . er c e a co o

fi t i n i n h m t xy lin t i n
x a o ,
ro X 2 600
— e a o s a . .

38 S tyl f w p w i g mu l i l u di g f u m l x d
’
ar c os e o as in th s n sc e nc n o r sar c o e re s , e re a e

co n di t i n w i t h u nu u ll y h
o ,
t Q di X 2600 s a s or -
sc s . .

39 T h m t ly t g i n
re e s ar c o nt ti A v y n w H di
er e s a e ar s a e co ra c on . er arr o - sc

b i t t h u nu u l l y l g Q di
se c s e X 2600s a on -
sc s . .

40 d 41 S u
an iv ly l t p h f nt
c c essti X 2600 e a er ase s o co r ac on . .

42 S t i l l l t ph f nt ti n
a er Tw ly nt
as e ti n b nd o pp co r ac o . o e ar co ra c o a s a e ar

t w d t h u pp
o ar nd f th e tyl er h emp d f pp i t h l v f
o e s ar c o s e, e ac co o se o o os e a es o

su i v Q di f u d b u t t h t l p h g m X 2600
c c e ss e -
sc s se a o e e o ra a . .

43 L t p h f a et ti n h w ing as e o t g in th c on v l f t i ti n rac o ,
s o a s a e e re er s a o s r a o s .

T h H di e f th - li ph
sc o f nt t i n n w f m t h l i gh t t i p
e e ar er f th as e o co r ac o o or s e s r e o e

fib ri l . X 2 60 0 .

44 S a r c os t yl f u l l c o n t rac t i o n
e in ,
s h ow n i g d
fi v e ( o u b l e ) c o n t r a c t o n an i b d s .

Th e se fi b er s al w a y s b r e a k a l o n g t h e m idd l e o f a c o n t r ac t o n b an , t h a t i s , i d at

t he l e v el o f t h e t e l o h ra m a p g . T he f ai n t st r ip es b l se c t i n g t h e s p ac e b etw een
s u c c es s iv e c o n t r ac t o n i b an d s m ay b e t h e m e s0 p h r ag m at a . X 2600 .

45 Sm al l ar e a of t r an s v e r se l y cu t w ing m u sc l e of w as p . Th e w i n g l ik e
-
‘

p r oc e s se s o f t he sar c o so m es a r e see n to en c rc e i l the sar c o s t y l es A b ov e i s s h ow n

.

a nu c l eu s . Fl e mm i n g ’
s fix at i o n , i ro n - h e m at o xy l i n st a n i . X 1 300 .

46 S ar c o s t yl v i e ew e d on en d , f rom a p p re ara t o n i ac c o r di n g to R o ll e t s
’

metho d wing p Th p if d mon g

‘
sh o at i o n a r t T he
’ ’
fix
‘
, o res . e se or es ar e ac t s . ots a

t he p ore s re p r ese n t t h e m e t a fi b r i l l a e . X 1 300 .

47 T ran s v er se se c t o n i of t h re e sar c o s t yl es of w as p ,
fix ed in F l mm i g
e n
’
s

s o lu t o n i an d i d st a n e w th i i ro n - h e m at o xy l i n ,
sh o w n i g r es u l t s of p g iv ro re ss e

d es t a i m n g w i th t h i e r o n - al u m so l ut o n i .

48 Sv e en s arc o st y l es an d fiv e sa r c o s o m e s f
m t he w n m u sc l e o f t he w as
ro i g p ,

cu t t ra n s v e rsel y . Fl em m ing ’
s fix at i o n , i r o n - h em at o l n st a n X 2500 xy i i . .
R esumen por la a utora Christiann a Smith , ,

Un iversidad Corn ell Ithaca ,

.

E st u dio del lipoide con ten ido en c l t 11 b ul o del ri non

’

El presen te est udio sobre el lipoide del rinon co n d u ce a las

sig uien tes co n cl usion es : 1 L a presen cia de lipoides e n mascara .

dos o libres es c aract eri sti ca de las cél ulas n ormales del r i non ,

y p u eden pon erse de m an i fiest o desp ués de la fi j aci 6n median te el

bicromato en calien te 2 En ciertas especies (por ej emplo el . .
,

gato ) diferen tes porcio n es del t ub ulo presentan formacion es

lipoides caract eri st i cas que p u ede n in dicar un a di feren cia f un
cion al 3 L os basto n es m i t oc on dri al es de la rama ascen den te
. .

del asa med ular son in ten samen te li p oi deos por n at u raleza re ,

sol vi én dose en gran os de lipoide baj o ciertas co n dicio n es 4 . .

L as g ot i t as li p oi deas q ue con tien en un tan to por cien to elevado

de olein a n o se con servan c on el mé todo de Bell 5 L a pre . .

sen oia di stri bu ci on y en algun os casos la di st ri b u ci on car

A

, , ,

act eri st i ca de los lipoides en las cél ulas del ri non i n dican q u e

p eu den estar i n t i mamen te relacion ados con procesos m e tab o

licos adem as de la posible f un ci on que se les atrib u ye por
,

algun os autores como agen tes que i nfluyen en el estado fi sico

del protoplasma L a i nt ro duc ci on del presente trabaj o con tien e
.

un a corta revisi on del trabaj o ya efect u ado sobre el or i ge n y

prese n cia de lipoides en los tej idos y especialmen te en las células ,

del rifi on L as propiedades hi sto qui mi cas de los lipoides se de

.

scr i b en y tambié n el material y métodos empleados en el prese n te

trabaj o L a mayor parte de las observacion es f ueron hechas

.

sobre las cél ul as ren ales del gato pero las del perro con ej o y , ,

rata fu ero n también examin adas El trabaj o termin a con un a .

disc u sion gen eral de la si gn i fi caci on de la presen cia de lipoides

en los tej idos .

Tra n sl at i o n by J osé F . N o n i d ez
C ar n e gi Ie n st i t ut i o n of Wa s h i gt
n on
A U T H O R S A BS T R A C
’
T OF T H I P AP ER
S IS S U E D

BY TH E BI B I O G R AP H IC
L S E R V I C E J A N U A RY 19
,

A ST U D Y OF TH E L IP O I D C O N T E N T OF TH E KI D N EY
T U B U LE
CH R I S TI A NN A S M IT H
Dep ar tmen t of H i s tol og y an d Embr yol og y, Corn el l Un i vers i ty, I thaca , N ew Yor k

F O UR TE E N F I G UR E S (T W O P L A TE S )

C O N TENTS

I t d ti
n ro uc o n an d h i st o r y.

Hi t h mi
s oc e c al c h ar ac t e r s i ti cs

M at er i al an d m e t h o d s
M o r p h o l o g y o f t h e u r i n ar y t u b u l e
O b ser v tia ons

kidn y
O n the e of t he c at

k idn y
O n t he e of t he do g
O n t h k id n y e e of t h e r a bb i t
O u t h k id n y e e of t h e r at
G n l di u i n
e er a sc ss o

I NTR O DU C TI O N
The omn iprese n ce of lipoids i n the tiss ues has bee n poin ted
ou t by chemists an d st u died by man y Cowdry i n his excellen t .
,

s u mmary of the fun ction al si gni fi can ce of mitocho n dria whi ch ,

are related to the ph o sph ol ipi n s qu otes Matthews apropos ,

of this w h o says that ph o sph ol ip i ns are fo un d i n all cells and

, ,

that it is un do u btedly their fun ctio n to prod u ce with choles ,

t er ol the pec u liar semi fl ui d semi solid state of protoplasm

, , .

However the prese n ce of lipoids has bee n overlooked by hi st ol

o g i st s i n ge n eral un til q u ite rece n tly becau se they are dissolved ,

i n man y fi x er s an d req u ire special tech n iq u e to preserve them .

That lipoids are presen t i n kid n ey cells has bee n k n own b ut ,

almost always it has bee n co nn ected with pathological chan ges .

Believi n g that microscopically lipoids co u ld be demo n strated i n

n ormal cells this st u dy of the lipoid co n te n t of re n al epitheli u m
,

69
70 C H R IS T I ANNA S M ITH

w as un dertake n at the s u ggestio n of D r B F Ki n gsb u ry w hose . . .

,

ki ndly i n terest an d ge n ero u s help is gratef ully ack n owle dged .

The co n cl u sio n s reached were that lipoids co u ld be demo n strated

i n kid n ey cells by special tech n iq u e that the distrib u tio n an d ,

formatio n s of the lipoids were more characteristic i n some species

than i n others an d that some lipoids may be i n tracell ular i n
,

origi n.

A ltho ugh it is agreed that lipoid is a p oor term chemically

“
,

speaki n g ( L eat hes i t is u sef ul i n histology to i n cl u de fats

, ,

fatty acids ph o sp h a t i ds cholesterol etc s ubsta n ces which

, , ,
.
,

have the same sol ve n ts an d wh i ch are fo un d associ ated i n cyto

plasm ( Ki n gsb u ry
,
a n d as s u ch it will b e employed i n

this paper .

If these lipoids are un iversally prese n t i n cells as it is believed ,

they are by chemists an d histologists ( Fischer an d Hooker 1 7 ; ,

’

Ki n gsb ury ,
the qu estio n arises as to the forms i n which
they exist an d their relatio n to the other cell co nte n ts i n t h e cell .

F i scher an d Hooker believe that i n n ormal cells an d flu i ds liqu ids ,

( Fischer an d Hooker use the term fat ) are prese n t i n fi n ely

‘ ’

d ivided form kept so by vario u s hydrated protei n s an d that the

, ,

amo u n t of lipoid varies greatly n o t o n ly i n a give n ce l l or

body flu id u n der d iffere n t physiological an d pathol o gical circ um
stan ces b ut also at all times i n di ffere n t cells an d fl ui ds It is
, ,
.

becau se of these fi n e em ulsio n s they say that large amo un ts of

, ,

lipoids may be prese n t i n certai n tiss u es an d n ot betray them

selves optically or by ordi n ary fat stai n s A ccordin g to this .

i n terpretatio n fatty dege n eratio n is merely a coarse n i n g of the

,

n ormally fi n e em ulsio n s d u e to the separatio n of the lipoids

becau se of an i n terfere n ce with the hydratio n of the protei n s .

This i nterfere n ce is accordi n g to them acid prod u ctio n cau sed

, , ,

by s ubstan ces s u ch as phosphor u s phlorizi n alcohol and c o n , , ,

d i t i o n s of an aemia or ge n eral circu latory dist u rba n ces .

That some of the lipoids i n cells are prese n t i n i n timate m ixt ures
has already bee n poi n ted out ( Ki ngsb ury an d that o n e ,

o f these mixt u res is represe n ted by mitocho n dria seems at prese n t

un q u estio n ed Cowdry . i n the paper to which refere n ce has

a lready bee n made gives the followi n g s ummary of the chemistry
,
 LIP O I D C O N TE NT OF THE K I D N E Y TU BULE 71

of mitocho n dria Mitocho n dria are sol uble wholly or partially

.
,

i n fat solve n ts alcohol ether chloroform an d dil ute acetic acid

, , , , ,

a n d the par t which is n o t sol u ble is a protei n ( alb u mi n ?) They

a re re n dered i n sol u ble by ch r o m at i o n Mitocho n dria do n ot
“

stai n with s udan III or scarlet red an d are blacke n ed o n ly at

times with osmic acid Becau se of their lipoid n at ure some
.
,

have so ught to fi n d i n them the so u rce of lipoids i n the cell while ,

o thers believe they themselves arise from lipoids ( Cowdry ,

The problem of the so urce of visible lipoids i n tiss u es is on e

that has aro u sed m u ch i n terest an d has bee n very mu ch i n volved
i n the q u estio n s of fatty i n filtratio n an d fatty dege n eratio n .

O n e explan atio n that Visible fat is d ue to the coarse n i n g of a

‘ ’

fi n e emulsio n of lipoid has already bee n give n O thers believe .

that fatty i n filt rat i on is an excessive depositio n of fat an d that

fatty dege n eratio n is a co n versio n of cell su b stan ce i n to fat .

(Virchow from Fischer an d Hooker ) E T Bell refers

“
. . .

to two diverge n t opi n io n s on this s ubj ect i n his paper o n The

D i ffere n tial Stai n i n g of Fats ” R ose n feld
. an d Krau s

believe tha t kid n ey fat is derived mai nly from the destr u ctio n of
i ntracell ular lipoids or from str u ct ural chan ges i n the cytoplasm
whereby fi n ely divided fat becomes Visible R ibbert says .

that lipoids are mai nly extracell ular i n origi n an d that the origin
of i n tracell ular lipoids is comparatively u n importan t Bell
'

s u ggests that oleic fat is extracell ular i n origi n an d other lipoids

are i ntracellular From t hi s brief acco un t of the problem of

.

lipoids i n tiss u es it can be readily see n that the qu estio n is o n e

,

o f importan ce ,
an d the si g n i fi c an c e of the prese n ce of lipoids

will be taken up i n the ge n eral disc u ssio n at the en d of the paper .

J u st as it has bee n bro ught o ut i n the fi rst part of the i n tro

d u ctio n that tiss ues possess lipoids which are di ffi c ult i n ge n eral
,

to demo n strate microscopically so is it tr u e co n cerni n g the lipoid

,

co n te n t of the kid n ey Krau ss i n 1 9 04 makes the stateme n t

“ ,
.
, ,

that tissu es that are microscopically fat free an d look n ormal ,

i n ge n eral for example kidn ey cells may co n tai n 2 0 per ce n t of

fat .
” , ,

A s late as 1 909 1 9 1 0 Mayer an d Bathery state that n o

—
,
,

fat disclosed by u s u al reactio n s is fo un d i n th e kid n ey cells of

mammals .
72 C H R IS T I ANNA S M ITH

his st udy of the lipoid co n te n t of epitheli um cartilage an d

In , ,

m u scle fi b ers of the ox E T Bell makes some observatio n s , . .

o n the lipoids prese n t i n the kid n ey an d fi n d s that the re n al ,

cells of some ox foet u ses co n tai n ed lipoid droplets an d that the ,

cortical re n al cells of two large steers were loaded with lipoids .

1

O ther observatio n s n ot his o wn are recorded by Bell i n this, ,

paper A scho ff . an d Pfeiffer described lipoids i n the

kid n ey an d other organ s of n ew bor n childre n ; Han semann —

fo un d fat 1 ) i n the h uman kid n ey an occ urre n ce which he

, ,

co n sidered u s u ally pathological b ut which might be n ormal ; 2 ) ,

i n swi n e especially whe n fatte n ed whi ch he tho ught ab n ormal

, ,

an d a n alogo u s to obesity i n m an ; 3 ) i n cats an d dogs In 1 9 1 0 .

,

Bell exami ned the tiss u es of calves cats dogs rats and frogs , , , , ,

an d fo u n d u s u ally a large nu mber of lipoid droplets i n the c o n

vol uted t ub ules a fewer nu mber i n the collecti n g t ub ules an d

, ,

i n some t ub ules n o n e H e observed also that the amo un t of .

lipoid i n the kid n ey varies greatly the cat showi n g the gr eatest
”
amo u n t an d the ox le a st In his work O n the D iffere n tial
,

”
.
, ,

Stai n i n g of Fats Bell u sed the kid n ey tiss ue of h u man

an d rat as o n e ki n d of material .

Polic ard me n tio n s i n most cases i n his work o n the histoge n esis
an d histology of the re n al epitheli u m o n ly the prese n ce of lipoids
“ ”
.

In his paper on the F un ctio n i n g of the Kid n ey of Frog

he makes however the followi n g s ummary of his observatio n s
, ,

i n regard to the lipoids i n the co n vol uted t u b u les The forma .

tio n s of lipoids which red u ce osmic acid b ut which stai n with ,

copper hematoxyli n after the method of Weigert R egau d are ,

-
,

of three ki n ds : First there are flakes of material which stain ,

gray that are e n closed i n vac u oles These are by him con .
, ,

si der ed artifacts Seco n dly very fi n e granules are prese n t which

.
,

stai n bl ue black and which correspo n d to the i nn ermost sub

—

c utic ular vac u oles described by vario u s observers b ut whether ,

it is the wall of the vac uole or the s ubstan ce co ntai n ed which

stai n s he does n ot k n ow Thirdly i n the regio n of the nu cle u s .
, ,

1 T h e t erm l
‘
i p id
o
’
h as b ee n su b s t i t u t e d h ere w h erev er B el l h as u se d th e w or d
‘
f at , ’
a l t h ou gh he u ses
‘
f at
’
i n t he g e n er al se n se to in c lu de t ru e f a t s a n d o t h er

l i p o id s .
 LIPO I D C O NT E NT OF TH E KI D N E Y TU BULE 73

larger grai n s exist at the periphery of the sectio n which he c on

siders badly fi x ed mitocho n dria These flakes an d gran ules .

which Policard describes are qu ite clearly lipoids i n combi n atio n

that are stai n ed by the mitocho n drial tech n iqu e which he u sed .

Evide n tly Policard does n o t take i n to co n sideratio n the fact that

i n the preservatio n of lipoids the best fixatio n of a mass of tiss u e
is at its periphery while at the ce n ter of the mass the fix at i on
,

may be i n complete In disc u ssi n g other portio n s of the t ub ule

.
,

Policard me n tio n s that lipoids are rarely fo un d i n that portion

which correspo n ds to the asce n di n g limb of the med ullary loop
( loop of He n le ) .

M ac N i der i n his work o n n ep hropathic dogs refers ,

to the prese n ce of fat i n the asce n di n g limb of the med ullary loop
as of frequ e n t occ urre n ce .

In his i n vestigatio n of the fat i n fi lt r at i on of t he cat s kid n ey

’
,

Mottram observes that lipoids are prese n t i n the co n volu ted

t ub u les an d rarely i n the straight t u b u les H e checks up i n .

thi s paper an obse rvatio n made by L eat hes that the lipoid
, ,

co nte nt of the cat s kid n ey i n stead of bei n g more un sat urated

’
,

than that of of the liver an d lipoid tiss u e as it is i n the kid n eys

‘
,
’

of other an imals m an dog goat pig is sometimes more sat urated

, , , , ,

than lipoid tiss u e an d n ever as u n sat urated as that of the liver

‘ ’
.

HIST O C HE M I C AL C HARA C TERISTI C S

The observ atio n s made by Bell i n his several papers i n regard
to the histochemi cal properties of lipoids may be tab ulated as
show n o n page 7 4 Most of these observatio n s were made i n
.

the st u dy of mu scle b ut they are applicable as well to lipoids i n

,

the kid n ey Fats that are solid at ordi n ary room temperat ure
.

do n ot form an y part of the stai n able s ubstan ce altho u gh s u ch ,

fats may be prese n t i n small q u antity .

M A TE R I A L AN D M ETH O DS

The materi al u sed i n this i n vestigatio n was take n from cats ,

dogs rabbits an d white rats that were killed by ill umi n ati n g
, ,

gas an d the tiss u e fixed immediately or at di fferen t i n tervals

,
74 C H R IS T I ANNA S M ITH

S l bl i b l t
o u e n a so u e al c oho l o r e t h er

Rf e r ac t o n i S t gl y f t i v
ron re r a c e I n t erm ed 1 at e l y re F i tl y f
a n r e r ac t iv e

f r ac t i v e

E ff ec t of fix at i v es L ittl e aff ec t e d by G d u ll y ra a d i s ap D i pp
sa e ar i n m os t
an d p ost m o r t em fix a t i v es ( e x c ep t p e ar un d er t he fix ing a g en t s ,

c h an g es f o r m al ni ) o r p o st i n fl u e n ce o f fi x i n g su c h as f or m al in
m o r t em c h an g es g
a en t s , an d o st p an d p o st mo r t em
m o r t em c h an g es . c h an g es . P re
P r e ser v d
e by s er v d by B
e ell s
’

B el l s
’
10 p er 10 p er c en t di
c en t di c h rom at e c hro m at e m e t h o d
me th d o

St i i g
a n n an d fix in g i
r eac t o n s

1 . S c arl e t r e d , f r esh I n t e n se l y i d L
sta ne ess i nt en s el y F i nt l y
a st a n e i d
m at e r i al st a n e i d
2 . S d u an III a ft er A nnu l ar f orm of S l id
o re d dr pl o et

B el l d plro et

3 . O sm i c ac id St i bl k
a n ac St i b
a n r ow n o r n o t N ot i d
st a n e

c ol o r e d
C h em i c a l c om p i Eos ssen t al li y i
ol e n O le in m ix d e w th i L ow m el t ing f at
t i on so m e o t h er f at t y (bu tyrin )
su b s t an c e , e .
g .
,

c h o l e st e r o l

after death Bell an d B ullard

. emphasized the
n ecessity of u si n g fresh materi al if al l fat prese n t w a s to be

demo n strated an d Policard an d G ar n ier ,

fo un d that re n al
cells un derwe n t postmortem chan ges fift een mi nu tes after death .

In the st u dy of the lipoids three ki n ds of material are u s u ally ,

employed : fresh tiss ue ; that which has bee n fix ed i n a dichromate

sol utio n where the red uctio n of the dichromate re n ders the fat
i n sol uble an d stai n able with hematoxyli n an d that which has ,

bee n preserved i n osmic acid w hich certai n lipoids will red uce ,

with s ubseq u e n t blacke n i n g of them In order to get as complete .

an idea as possible of the lipoid co n te n t of cells these three ,

methods sho uld be u sed o n e as a check for the other The ,

.
76 C H R IS T I ANNA S MITH

recomme n ded by H uber have bee n u sed i n this paper .

The parts of the t ub ule an d the kinds of epitheli um are b ri efly , ,

as follows : First comes the re n al corp u scle with its do uble

,

walled caps ule of flat epitheli um Co n ti nu o u s with the epi

.

theli u m of the o uter wall of the caps ule is the short ofte n i n dis ,

ti n ct n eck which un ites the caps ule with the fi rst portio n of the
,

t ub ule the proximal co n vol uted t ub ule with the med ullary
,

segme n t The epithelial li ni n gs of the proximal co nvol uted

.

portio n an d the med ullary segmen t are esse n tially the same
thro u gho ut The high cells with striated free border i n distin ct
.
, ,

cell bo un daries an d rodded protoplasm of the basal portio n s of

,

the cell are all disti n g ui shi n g feat ur es The med ullary segmen t .

of the proximal co n vol uted portio n is followed by that part of

the t ub ule k n own as the loop of He n le b ut for which H uber ,

s uggests the n ame med ul lary l o op The parts of this are the
,
.

proximal or desce n di n g limb the distal or asce n din g limb an d

, ,

the crest The epitheli u m of the desce n di n g limb is of a thi n

.

paveme n t type with relatively large n u clei which reach from

, ,

the top to the bottom of the cell an d may cause the cell to b ulge
i nto the l ume n The crest of the loop may be formed either by
.

the thi n desce n di n g limb if the loop is lo n g or if it is shor t by

, , ,

the thick asce n di n g on e The asce n di n g limb exte n ds to the

.

re n al corp uscle of its ow n t ub ule an d is followed at that poin t

,

by the distal co n vol uted portio n which exte n ds to the collecti n g

,

t ub ule Policard
. states that the type of epithelium
characteristic of the asce n di n g limb is co nti nu o u s i n the distal
co nvol uted t u b ule an d H uber does n o t believe the variatio n s
,

W hich may be prese n t i n the di ffere n t parts great e n o ugh to

warrant the recogn itio n of other types H uber however con .
, ,

siders the epitheli um as low col umn ar with i n disti n ct cell

, ,

bo un daries gran ular protoplasm i n disti n ct basal striatio n s and

, , ,

as possessi ng n o striated free border In his work on Batrachian .

an d mammalia n specime n s 10 Policard poi n ts o ut

’
,

that this segme n t possesses very thick rods or b at onn et s which ,

exte n d from the base of the cell to the edge borderi n g the l ume n ,

a characteristic which has bee n fo un d tr u e i n the forms st u died

d urin g this work A disc ussio n of this type of epitheli um will
.
 LIPO I D C O NTENT O F TH E K ID NEY TU BULE 77

be reserved un til the treatme n t of the asce n din g limb of the cat s ’

kidn ey whe n its lipo id co n te n t is take n up The tran sitio n

,
.

between the distal co n voluted an d collecti n g t ub ules is gradu al ,

an d the clear t ype of epitheli u m characteristic of this portio n

n eeds n o additio n al comme n t .

OB S E R VA TIO N S

In this st u dy of the lipoid co n te n t of the kidn eys cats wer e ,

u sed more e x te n sively becau se the lipoids prese n t were very

,

easily demo n strated by fix er s co n tai n i n g osmic acid D efi ni t ely .

c u t pieces of the kid n eys from thirty fo u r cats were fi x ed i n -

either Be n da s or Flemmi n g s fl ui ds A ll of these were fo un d to

’ ’
.

con tai n lipoids that wo u ld red u ce osmiu m tetroxid E leve n of .

these cats were from 4 1 to 52 cm i n le n gth meas ur i n g from .

occipital crest to base of tail fo urtee n were 30 to 38 c m n i n e

,
/
.
,

1 8 to 2 3 cm . that is grown cats half grown cats an d small

—
, ,
—
,

kitten s O f the se o n e died fo ur were killed by ether and the

.
, , ,

rest by ill umi n ati n g gas The older full g rown cats had kidn eys
.
-

whi ch were light i n color an d possibly pathological O f t he .

half grown cats an d kitte n s o n ly o n e pair of kidn eys were fo un d

—
,

to be pathological u po n gross exami n atio n the rest appeari n g ,

n ormal .E xcept for two kitte n s which had sore eyes the exter n al ,

an d i n ter n al appearan ces differed o n ly i n ge n eral thi nn ess or

fatn ess Takin g i n to co n sidera tio n the above observation s on e

.
,

co u ld almost always foretell how the sectio n of kidn ey wo uld

look after it had bee n placed i n Be n da s or Fle m min g s fl ui d A ’ ’
.

kidn ey of a very thi n kitten or half grown cat wo uld have a —

cortex with very characteristic black rays ; a fat kitte n wo uld

possess a kidn ey the cortex of which wo uld blacke n as a wh ole
, ,

while an old cat fat or thi n wo uld have a kid n ey with an i n ten sely
, ,

black cortex (fi g s 1 to . In this gross exami n atio n of the

kidn ey the tran si tio n betwee n the blacke n ed cortex an d lighter
,

med ulla w as very eviden t as a distin ct an d clear c ut lin e That - .

the cat possesses fat di ff eren t i n amo un t distrib utio n an d com , ,

positio n from other an imals was evide n t i n this very sup erfi ci al
exami n atio n an d may have some correlatio n with the
,
o b ser l -
78 C H R IS T I ANNA S M ITH

of M ottram
v at i o n s i n regard to the sat uratio n of the fats
prese n t i n the cat s kid n ey ’
.

A s a typical example of a kid n ey whose cortex is characterized

by blacke n ed rays o n e will be st u died from a 22 c m kitte n
,
-
.

which w as quite thi n The kitte n was killed by gas an d its

.

tiss u e fi x ed immediately The kid n ey appeared n ormal an d .

healthy A s the appearan ce of lipoids depe n ds u po n its preser

.

vatio n an d stain i n g the lipoid co n te n t will be exami n ed i n its

,

relatio n to t hemethods used .

Ten per cen t forma li n an d scar let r ed F ree ha n d sectio n s .

-

wer e made from material that had bee n i n 1 0 per ce n t formali n

for abo u t a week an d stai n ed with an alkali n e alcoholic sol ution
,

of scarlet red ( B ullard 1 2 F o rm ali n material was u sed

,
’

i n this case becau se the lipoids of the kid n ey of cat are little
affected by i t The rayed appearan ce of the cortex n oted i n
.

the gross exami n atio n of tiss u e fix ed i n Be n da s was appare n t ’

here an d the rays proved to be composed of t ub u les co n tai n i n g

,

heavily stai n ed lipoid granu les i n the med ullary rays The .

t u b ules of the labyri n th an d med ulla also co ntai n ed gran ules ,

b ut they were m u ch fi n er tha n those i n the rays .

P otassi um bi chr oma te a n d su dan II I This method was fi r st

u sed by Bell w h o said it wo uld stai n all gran u les see n i n a fresh
,

specime n an d wo uld also di ffere n tiate those co n tai n i n g olei n

from other lipoids The med ullary rays which were stai n ed
.

with Be n da s or scarlet red thereby stan di n g o ut promi n e n tly

’
,

becau se of their i nte n se stai n i n g were here very light i n color , ,

an d the t u b u les of the labyri n th were darker This variatio n .

w as d u e to the fact that those gran ules which were deeply stai n ed
by scarlet red or B e n da s were stai n ed o nly at the periphery or
’

n o t at all with s u da n III after c hr o m at i o n These droplets .

,

the n co uld be said to co ntai n olei n becau se they red u ce osmi c

,

acid an d beca use they were chr o m at ed o n the s urface or n ot at

all by Bell s method or are what he called an nular droplets
’
,
.

The med ulla w as also divided i n to two regio n s o n e correspo n di n g , ,

to the o uter zo n e of Peter which lies n ext to the cortex c o n , ,

t ai n ed more deeply stai n i n g t u b ules than the i nn er zo n e or

papilla .
 LIPO I D C O NT E NT OF THE K I D N E Y TU BULE 79

Whe n the cortex was exami n ed more closely it w as fo un d that ,

the cells of the proximal co n vol u ted t ub u le which had stai n ed

d eeply co n tai n e d a large n u mber of lipoid granu les some portio n s
, ,

of the t ub ules more than others A s stated by Bell whe n ther e .

,

was a small amo un t of lipoid prese n t the gran ules were collected ,

at the base of the cell an d whe n a large amo un t was prese n t

, ,

the granu les were scattered thro u gh the cell Where the lipoid .

was stain ed the mitocho n drial fil am en t s stood out amo n g them

,

i n co n trast very clearly A ltho u gh there was a te n de n cy for

.

o n e t u b u le to have the same ki n d of droplets it was very comm o n ,

to fi n d i n the same t u b ule both ann ular an d solid forms In .

some sectio n s of ann ular droplets a diff u sio n of the lipoid i n to ,

the s u rro un di n g cytoplasm co uld be see n qu ite plain ly It was .

also n oted that man y irregular shaped gran ules seemed to be -

composed of two or more droplets .

The t ub ules co n tai n i n g the very large droplets which stai n ed

i n te n sely with osmic acid an d scarlet red an d n ot at all or very ,

little with s u dan III were the med ullary segme n ts of the
,

proximal co n vol uted t ub ule A ltho ugh Bell describes the typ e
.

of droplets which co n tai n s olei n as ann ular from observatio n s ,

o n both n ormal an d pathological kid n eys where droplets which

stai n ed i n te n sely black were very ab un dan t it wo uld be see n ,

that the larger droplets were n ot preserved at all an d represe n ted

vac u oles That this co n ditio n w as n ot due to sectio n s where
.

the fix at i o n was i n complete w as shown by the fact that n eighbor

i n g proximal co n vol uted t u b u les possessed lipoid which stai n ed
with su dan III (fi g .

The tran sitio n from the med ullary segme n ts of the proxi mal
co n vol u ted t ub ule to the desce n di n g limb was very sharp and
the cau se of the clear cut li n e appare n tly betwee n the cortex and
—

the med ulla which was n oticed i n the gross e x ami n atio n The .

pave m e n t epithelial cells of this part of the med ullary loop or

the loop of He n le possessed solid lipoid granules quite n umero u s ,

an d large i n proportio n to the amo un t of cytoplasm i n the cells .

The asce n di n g limb of the loop had an appearan ce very differen t

from the other portio n s of the re n al t u b ule because of the presen c e
of the rods which Policard describes 10 12 A ltho ugh ’
,
’
,

TH E A M ERIC A N J OU RN A L OF A N A TOM Y VOL

, . 27 , NO . I
80 C H R IS T I ANNA S MITH

these are called mitocho n drial rods an d altho ugh Be nda con ,

siders them mitocho n dria properly speaki ng Policard does n ot ,

agree for the followi n g reaso n s : 1 ) Mitocho n dria are fl ex ous

fi l am ent s a n d these rods are like sticks arran ged i n b un dles o n
, ,

cross sectio n appeari n g like cardiac m u scle

-
2 ) these
bat o nn et s are more easily preserved than mitocho n dria altho u gh
‘ ’
,

mitocho n drial tech n iq u e bri n gs them o ut more clearly 3)

mitocho n drial fi lam ent s i n the proxim al co n vol uted segme n t
vary i n form becomi n g at times granular n ormally an d d ur i n g
,

autolysis while the rods of the third segme n t (which correspo n ds

,

to the asce n din g limb of the med ullary l o op ) n ever becomes

gra nular n ormally n or d ur i n g au tolysis A s to their n at u re
“
Policard describes them as protoplasmic rods the s urface o f
,

which see m s to be covered with a lipoid s ubstan ce ” ,

T hat these rods are lipoids or fatty i n n at u re is show n very

clearly i n these section s an d it is becau se s u dan III is sol uble i n
,

them that the regio n of the med ulla n ext to the cortex is stai n ed
more deeply red In thi n sectio n s where the rods were n ot so
.

closely packed that their i n divid u ality w a s hard to determi n e ,

two types co uld be disti n g u ished O n e form correspo n ded to

.

the homoge n eo u s rods of Policard an d they appeared like annular

,

droplets drawn o ut to form rods W hich varied i n le n gth an d wid t h ,

some bei n g slightly irreg ular It may be possible that the

.

method of fix at i on with the heat may have affected some of the

rods an d that they were atypical The other form was composed
.

o f small gra n u les arra n ged i n rows ( fi g This form of the

.

rods is q uite co n tradictory to P ol i card s descriptio n of them ’

.

S mall annular droplets an d solid gra nules were prese n t which

might be i n terpreted as sectio n s or tips of rods or distin ct i n
themselves .

The distal co n vol uted t ub ule had epitheli um of the same

heavily rodded character as that of the asce n din g limb of the
med ullary loop an d the cells of the collectin g t u b ules co n tai n ed
,

a few solid fat granules O n e portio n of the ren al t ub ule has

.

n o t bee n disc u ssed an d that is Bowma n s caps ule The fl at ’

.

epitheli um of the caps ule also possesses lipoid gran ules as do ,

the other segme nts of the t ub u le .

 LIP O I D C O NTENT OF THE K I D N E Y TUBULE 81

Ten p er cen t p o tassi um di chr oma te an d per os mi c aci d . T en

ce n t potassi um dichromate per ce n t osmic acid were u sed
an d 2

i n the proportio n s of fo u r to o n e an d 3 5 cc were a ci di fi ed by .

t hree drops of glacial acetic acid The tiss u e was s ubj ected to a
.

temperat ure of 51 for forty eight ho urs M aterial treated i n

°
—
.

this way showed the fat which red u ces osmic acid more clearly
an d ab un dan tly tha n tiss u e fi x ed i n either Flemmi n g s or Be n da s
’ ’

fl uid In imbeddi n g both p araffi n an d p arl o di o n were u sed for

.
, , ,

altho ugh more fat is preserved i n p arl odi on sectio n s paraffin o n es ,

were fo un d very usefu l i n defi ni t ely locati n g the gran ules i n the
di ffere n t regio n s of the t ub ule .

The rays which were described as stai n i n g black i n pieces of

tissu es fix ed i n Be n da s fl uid stood ou t very clearly an d in te n sely
’

an d the labyri n th was left un stai n ed except for some very fi n e

,

granules (fi g s 5 to . The same regio n s of the med ulla were

presen t as were described for material fix ed i n 1 0 per ce n t di chro
mate an d stai n ed with s udan III Here the o uter zo n e appeared
.

gray black an d the i nn er zo n e was left un stain ed The large

-
.

black glob ules were present i n the med ullary segme n t of t he

prox imal co n vol uted t u b ule an d were the same as those which
stain ed in ten sely red an d n o t at all or on ly at the periphery with
s u dan III O ther parts of the proximal co n vol uted t ub ule had
.

very small gran ules at the base of the cells which did n ot stai n
an i n te n se black In the co n vol uted t u b u les of the kid n ey of a
.

2 3 c m kitte n there was the same li n ear arran geme n t of fi n e fat

-
.
,

gran ules tho u gh n ot so regularly placed as may be fo un d i n the

, ,

asce n di n g limb of the med ullary l o op These formatio n s i n the

.

asce n din g limb will be described i n the n ext paragraph This .

lin ear arran ge me n t of the lipoids i n other portion s of the urin ary
t u b ule besides the asce n din g limb may be due to the positio n
of the lipoids i n relatio n to mitocho n dria either passively or i n
co n sideratio n of a m ore i n timate co nn ectio n or it may be an ,

early stage of fat formatio n i n depe n den t of mitochon dria (fi g .

No lipoid was visible i n the desce n di n g limb of the medullary

loop which stain ed black with osmic acid In the asce n di n g .

limbs those rods which at times appeared gran ular an d at other

, ,
82 C H R IS T I ANNA S M ITH

times homoge n eo u s whe n fix ed an d stai n ed by B ell s method ’

,

were here very d efi n i t ely composed of gran ules which stai n ed

black ( fig . These gran ular fil am en t s starti n g at the base of
the cell exte n ded to the edge borderi n g the l ume n an d occ upied ,

the space betwee n the n u cle u s an d periphery of the cell They .

r an parallel to each other an d their beaded appearan ce w as

absol utely clear In most t u b u les these granular fi l am ent s so

.
,

very disti n ct i n the med ulla were n ot preserved i n the cortex .

This w as probably due to imperfect preservatio n for i n the ,

dog s kid n ey which will be described later they were well

’
, ,

preserved i n the cortex The co n cl usio n seems j u st ifi ed there

.
,

fore that these homoge n eo u s rods which Policard describes as

“
,
”
protoplasmic rods with a lipoid s urface coveri n g n ever b ec o m
i n g gran ular are lipoids an d do become gra n ular fi l am en t s which
,

stai n black with osmic acid whe n treat ed W ith certain fi x ers as ,

a mixt ure of potassi um dichromate an d osmic acid with or ,

witho ut the applicatio n of heat .

The collectin g t ub ules did n o t co n tai n lipoids that redu ced

osmic acid .

Ben da s flui d Material fix ed i n Be n da s fl ui d showed the

’
.
’

same characteristics as that fix ed i n potassi um dichromate an d

osmic acid The preservatio n of the lipoids as it has bee n
.
,

s t ated before w as better i n the bichromate sol utio n s

,
7
.

O n e p er cen t osmi c a ci d In a simple sol utio n of os m ic acid

.
,

the same details were eviden t that were presen t i n the other
osmic acid fixers The rods of the ascen di n g limb of the m edul
-
.

lary loop appear granular upo n foc u si n g b ut homoge n eo u s W he n ,

i n packets .

The disc u ssio n of the tissu es fix ed i n H elly s fl ui d Ze nker and ’

, ,

R egaud will be omitted i n this paper an d take n up i n an other ,

whe n the effects of di ffere n t fix er s and the lipoids of the kid n ey

will be st u died i n their relatio n to mitocho n dria .

A s an example of a very fat kid n ey with a very black cortex o n e ,

will be take n from a cat which meas ured 4 6 cm occipital crest .

to the base of the tail This kid n ey possessed a very yellow

.

cortex i n the fresh co n ditio n an d w as probably pathological as ,

the liver appeared to be also .

 CH R I S TIA N N A S M ITH

From these observatio n s of the lipoid co n ten t of the kid n ey

of the cat it c an be see n that there is a characteristic distrib ution
,

of the lipoid granules which varies un der differe n t co n ditio n s ,

the n ormal an d pathological n ot appearin g the sam e Bec au se .

of the ab un dan t s upply of lipoids their characteristic formatio n s

, ,

s u ch as the gran ular forms of the rods of the asce n din g limbs ,

an d distrib utio n i n the cat s kid n ey it sho uld be a favorable

’
,

place i n which to st udy the relatio n of lipoids to mitocho n dria .

The lipoid co n te n t of the cat s kid n ey may be s umm arized as

’

o n page 85 .

The res ults ob t ai n ed from the st udy of the kid n eys of other
animals are added here with the realizatio n of their in complete
n ess an d the n eed of f u rther st u dy E T Bell s umm arizes the
. . .

factors which have caused co n fu sion i n the st u dy of fat i n mu scle

fi b er s They might well be repeated here i n regard to the
lipoid co n te n t of the kid n ey They are the followin g : 1 ) the
.

character of the fat stain employed ; 2 ) the species of animal ;

3 ) the character of the an imal s food ; 4) the ge n eral nu tritive
’

co n ditio n of the animal ; 5) the e ffect of fix at i ves ; 6) the e ffect of

postmortem chan ges That the character of the fat stain and
.

fi x at i ves employed an d co n ditio n s of the an imal are very i m

portan t is easily see n from the st u dy of the lipoid co n te n t of the
cat s kid n ey That the species of the animal an d e ffect of post
’
.

mortem chan ges m u st also be co n sidered is evide n t from the

followi n g observatio n s on the dog rat and rabbit , ,
.

Dog

A ltho u gh it might be expected that the dog an d cat wo uld

have kidn eys very similar i n their lipoid co n te n t the reverse ,

was fo un d to be tr ue O f the six dogs exami n ed n o n e prese n ted

.
,

the rayed appearan ce of the corte x so characteristic of the

kitte n s and half grow n cats b ut looked more n early like the
-
,

very fat kitte n (fi g

'

Two of these six dogs were fi v e day old

.
- -

p uppies on e a p uppy n ot yet wea n ed b ut whose exact age was

, ,

n o t k n ow n two were t w o weeks an d fi v e days old and o n e a

, ,

half grow n dog w ith the man ge O n the kid n ey of the p uppy
—
.

of un k n own age the followi n g observatio n s were made .

LIPO I D C O NTENT OF TH E KID NEY TU BULE 85

a a
c bE So
m& a
c
e o s
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$ y
o
fi
9 fi
r
au

m
o
5
x
s

a fi 3 8 E
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a
o 5 ?
w
1
3
S
8 o

E 5
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m
2
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86 C H R IS T I ANNA S MITH

Ten p er cen t forma li n gran ules were

a nd scar let red . No
preserved i n the proximal co n vol uted t ub ules i n most regio n s .

Some red stai n in g droplets were fo un d i n the t ub ules borderi n g

-
‘

the med ullary ray an d i n the med ullary loop an d collecti n g

t ub ule .

Ten p er cen t p otassi um di chroma te a n d su dan III The cyto

plasm of the cells of the proximal co n vol uted t ub ule stai n ed
di ffu sely with re d an d co n tai n ed irreg ular masses which might
be called sud an ophi le precipitate by Bell or perhaps ,

lipoids which have become dissolved i n the cells d uri n g the fi x at i on

with applicatio n of heat The lipoid granu les i n the asce n di n g
.

an d desce n di n g limbs of the med ullary loop an d i n the collecti n g

t ub ules showed clearly .

Ten p er cen t p otassi um di chroma te a n d osmi c a ci d The lipoids .

Whi ch blacke n ed with osmic acid were prese n t as fi n e gran ules

i n the asce n di n g limbs of the med ullary loop an d the lower ,

portio n s of t ub ules which were i nterpreted as the med ullary

segme n ts of the proximal co n vol uted t ub ules an d were fo un d i n
the labyri n th borderin g the med ullary ray In the asce n di n g .

limbs of the med ullary loop i n the cortex the granular rods ,

w hich were so clear i n the med u lla of the ca t were prese n t

'

, ,

tho ugh n ot preserved i n the medulla ( fi g In 1 per ce n t .

osmic acid the rods i n the asce n di n g limb appeared homoge n eo u s

, .

R a bbi t

As the kid n ey of the dog differed from that of the cat i n its
lipoid co n te n t so also is the kid n ey of the rabbit differe n t from
,

both of them Kid n eys of eleve n rabbits were exami n ed which

.

h a d bee n fi x ed i n Be n da s fl u id an d n o lipoids which wo uld red u ce

’
,

o smic acid were observed Fo ur of these rabbits were from seve n

.

days to seve n an d o n e half weeks old and observatio n s made o n

-
,

the ki d n ey of the seve n and o n e half weeks old rabbit will be give n
-
.

The older rabbits whose ages were un k n ow n were affected with

coccidiosis b ut the yo un g rabbits were n ormal an d healthy
,
.

F r esh ti ssue The cells of the t ub ules co n tai n ed fi n e red

.

g ran ules whe n stai n ed with scarlet red an d some t u b ules c o n

 LI P O ID C O NTENT OF THE K I D NEY TU BULE 87

cells with granules which stai n ed brown with

t ai n ed 1 per ce n t
osmic acid .

Ten p er cen t p ota ssi um di chroma te the

an d su da n III U n like
kidn ey of the cat the cortex did n ot possess any differe ntiatio n
,

in to rays b ut like i t possessed the two regio n s of the med ulla

, , , ,

the o uter zo n e stai n in g more deeply G ran u les were presen t i n .

the cells of the pro x imal co n vol uted t u b ules which Bell called
sud an O p hil e precipitate an d for which he did n o t co n sider the
,

evide n ce s ufficie n t to co n sider as diffu se lipoids A ccordin g to .

Bell sud an op hi le precipitate was n ot fo un d i n cells co n tain in g

,

lipoid gran ules A s droplets were fo un d i n the t u b ules of the

.

cortex which stai n ed with scarlet red an d brown ed with osmic

acid the co n cl usio n was reached that the gran ules here were
,

lipoids an d n ot precipitates The fatty n at ure of the rods i n.

the asce n di n g limbs of the med ullary loops was eviden t altho ugh
they stai n ed very lightly .

Ben da s flui d
’
Material autolyzed or n ot showed n o fat
.
, ,

whi ch wo uld redu ce osmic acid when Ben da s fl uid was u sed ’

witho u t the applicatio n of heat Also tiss u e which had n ot .

,

un dergo n e a u tolysis b u t was preserved i n Be n da s fl u id kept at

’
,

a temperat ure of exhibited n o blacke n ed gran ules Sectio n s .

which had autolyzed an d were fix ed i n the way j ust me n tio n ed

presen ted a q uite differen t appearan ce Man y black gran ules .
,

short granular fi l am en t s an d rods were presen t i n the cells of

,

the cortical t ub u les an d i n the i n terstitial tiss u e of the cortex

an d med u lla Therefore from these observatio n s it wo uld seem
.
,

that lipoids which red u ce osmic acid are fo un d i n the kid n ey cells
of the rabbit o n ly after autolysis an d fix at i on with heat Ziegler
“
.
,

i n his disc u ssio n of fatty dege n eratio n (p says that a .

process similar to that takin g place within the body occ urs
d uri n g autolysis of tiss u e preserved aseptically i n the i n c ubator ,

fat droplets becomin g visible i n s u ch tissu es ( He n sen W ent cher , ,

Krau s M uller and

, ,

To s um m arize fat is prese n t i n the kid n ey of the rabbit i n

,

th e fresh co n di t io n which stai n s with s u dan III after fi x at i o n with

dichromate : A fter au tolysis an d fi x at i on i n an i n c ubator if ,

os mi c acid is prese n t there are some lipoids which will stai n black
,
.
88 C H R IS T I ANNA S M ITH

The rods of the asce n di n g limb of He nle s loop agai n show their ’

fatty n at ure both i n dichromate fix at i on an d 1 per ce n t osmic

,

acid .

The kid n eys of rats were also exami n ed and the res ults i n
ge n eral were like those obtai n ed after the st u dy of the re n al
cells of the rabbit The kid n eys of n i n etee n rats were cut an d
.

fi x ed A ll of these rats were i n fected with lice b ut the i n ter n al

.
,

organ s appeared n ormal except for on e case of a cystic liver an d

o n e kid n ey that proved to be pathological whe n exami n ed

mi croscopically Kidn eys of rats which had bee n fed o n fat

.

diets were st u died an d both these and co n trols were s ubj ects
,

to au tolysis with fix at i on accompan ied by heat The resu lts .

i n brief were as follows : The rats possessed i n terstitial lipoid

gran ules which were especially ab un dan t i n the papilla The
,
.

presen ce of these gran ules made it di ffi cult i n the case of osmi c

acid fix at i on s to disti n gu ish always between i ntracellular an d
i nterstitial lipoid granules In the fresh tiss u e an d that preserved
.

i n formali n the i n terstitial were the o n ly lipoids that seemed

,

to be prese n t Tiss u e fi x ed i n 1 0 per ce n t dichromate with heat

.

an d stai n ed with s u da n III possessed gra n u les which Bell wo u ld

clearly call sud an op hi le precipitate for they were n o t observed ,

i n the fresh co n ditio n However if lipoids which stai n with

.
,

osmi c acid can be liberated after autolysis an d fix at i o n i n an

i n c ubator the n it does n ot seem un reaso n able to believe that
,

some lipoids i n combi n atio n i n the cell might be se n sitive to heat

an d liberated by i t c h ro m at ed a n d stai n able with s u da n III
, ,
.

Bell admits that more fat is demo n strable whe n higher tempera
t ures are u sed b ut does n ot yet i nterpret the sud an ophile p reci p
,

i t at e as di ff u se fat The gra n u les were fo un d i n all the cells an d

.

the cytoplasm was un stai n ed The res ults with Be n da s were .

’

the same as those observed i n the rabbit .

E xperime n ts to show the e ffect of fat diets were also carried

o ut .This feedi n g w as mostly o n fat meat an d bread soaked i n
fat o ne rat was fed olive oil twice on the day it w as killed There
,
.

was n o i nfilt rat i on of annular droplets as described by Bell ,

 LIP O I D C O NT E NT OF THE K I D NEY TU BULE 89

altho u gh with s udan III the gran ules appeared more n umero u s
an d somewhat more i n te n sely stai n ed an d a few fai n tly stai n ed
,

lipoid droplets were prese n t i n tiss ues fix ed i n an osmic acid -

sol utio n These res ults did n ot check Bell s work This may
.
’
.

be because the feedi n g was n ot forced .

GE N ER A L DIS C USSI O N
As it was stated i n the i n trod u ctio n the co n cl usio n s reached
,

i n this st u dy of the lipoid co n te n t of the kid n eys were that

lipoids co uld be demo n strated i n kid n ey cells by special tech ni qu e ,

that the distrib utio n and formatio n s of lipoids were more charac
t eri st i c i n some species than i n others an d that some lipoids ,

appeared to be i n tracell ular i n origi n The very defin i t e di st ri

.

b ut i o n of the lipoids i n the kid n ey of the cat has bee n show n a ,

distrib utio n n o t so marked i n the dog an d still less so i n the ,

rabbit an d rat However i n o n e segme n t there were lipoid

.
,

formatio n s pec uliar to them all These were the rods i n the
.

asce n di n g limb of the med u llary loop A s it w as poi n ted o ut i n

.

refere n ce to the cat an d dog these rods are lipoid i n n at u re an d

,

are resolved i n to fi l am ent s composed of lipoid droplets un der

certai n co n ditio n s A s these rods were co n sidered by Be n da
.

mitocho n dria properly speaki n g an d were called mitocho n drial

, ,

rods by Policard altho u gh he de n ied their ide n tity with them

, ,

the s uggestio n that there is a close relatio n betwee n these rods

an d the formatio n of lipoid droplets i n the cell o u ght n o t to be

overl ooked In the granu lar forms which were see n after a
.

fi x at i o n i n 1 0 per ce n t dichromate an d s u dan III a shadow of a ,

rod co uld be see n betwee n the granu les A fter a fi x at i on i n .

potassi um dichromate an d osmic acid the droplets stood out ,

clearly and separately i n a defini t e li n ear arran geme n t The .

prese n ce of lipoid droplets i n a li n ear arran gemen t i n cells of

other parts of the re n al t ub ule is also very s uggestive altho u gh ,

the evide n ce of their origi n is n ot clear Scott i n his work

.

o n the e ff ect of phosphor u s poiso n i n g o n mitocho n dria i n p an

creatic cells says that after the mitocho n dria lose their fil am en
,

to u s form that they aggl uti n ate an d fuse to form droplets

possessin g the characteris tic properties of lipoids Mitocho n dria .
90 C H R IS T I ANNA S MITH

as a so urce of lipoids i n the cells is disc u ssed by Cowdry

who thi nks that it wo uld require n o great stretch of imagin atio n
to believe that t his tran sformatio n co uld take place The .

i n tracell ular origin of lipoids does n ot however excl u de an

, ,

extracell ular origi n also as show n by Bell i n hi s experime n ts

,

with rats (Bell 1 4 ,

’

F urthermore the prese n ce of lipoids i n re n al cells either

, ,

masked or free the prese n ce of large amo un ts of lipoids i n cells

,

of the proximal co n vol uted t ub ule where it is gen erally co n ceded

either secretio n or absorptio n takes place the characteristic ,

occ urren ce of typical lipoid formatio n s ( large droplets i n the

med ullary limb of the proximal co nvol uted t ub ule i n cat an d the ,

rods i n the ascen di n g limb of the med ullary loop ) do n ot i n dicate

,

that lipoids i n kidn ey cells are merely passive str u ct ures n or ,

simply droplets formed by the coalescen ce of fi ner granules i n

an em ul sio n b ut that they may be a direct expressio n of activity
,

i n the cell eco n omy B ullard 1 6) a n d Bell have c on

’
.

cl uded that lipoids i n n ormal mu scle cells are a reserve food

s upply In disc ussin g the r 6le of fats (lipoids ) i n Vital phe n om
.

en a ,
L eat h es says that they are most co n spic u o u s as a
reserve f un d of fu el for growin g an d workin g cells b ut that i n ,

Virt u e of their ge n eral chemical i n ert n ess are capable of bein g

p u t to ma n y u ses i n the orga n izatio n of pla n ts a n d a n imals for ,

i n stan ce bei n g essen tial to the cohesio n an d physical co n stit u tio n

,

of the protoplasm In co nn ection with the u se of fat as f uel

.

for worki n g an d growi n g cells Batai ,

fi n d s that a lipoid free -

ratio n di min ishes the n ormal rate of growth of the body i n albi n o
rats L eat hes also says that the un sat urated fats fo un d i n the
.

cells of the body are broke n down by s uccessive oxidatio n s un til

they are completely b u r n t to oxyge n an d water I m rie i n his
‘

.
,

disc u ssio n of the fatty chan ges i n the liver heart an d kid n ey
, , ,

says that the lipoids are oxidized i n them to s u pply e n ergy In .

other words the lipoids are red u ci n g an d their probable relatio n

, ,

to the red uci n g power of cytop l asm is disc u ssed by Ki n gsb ury
i n his paper o n cytoplas mi c fi x at i on In this paper it is
.

poi nted out that as early as 1 8 8 5 E hrlich called atte n tio n to the
92 C H R IS T I ANNA SM ITH

From this brief s ummary of the si g ni fi c an ce of lipoids i n the

cells the evide n ce seems very s tro n g that besides prod u ci n g the
,

pec uliar sem ifiui d semisolid state of protoplasm that the lipoids
, ,

are i n some way i n timately co nn ected with the metabolic

process es .

S UMMA R Y

1 . L ipoids are characteristically prese n t ormal kid n ey

in n

cells either m asked or free an d are shown after a dichromate

fi x at i o n with the applicatio n of heat .

2 In certai n species ( example cat ) di ffere n t portio n s of the

.
,

t ub u le have characteristic lipoid formatio n s which may i n dicate

a differe n ce i n f un ctio n .

3 The mitocho n drial rods of the asce n di n g limb of the m edul

.

lary loop are by n at u re stro n gly lipoid an d are resolved i n to

lipoid granu les un der certai n co n ditio n s .

4 L ipoid droplets co n tai n i n g a large perce n tage of olei n are

.

n o t preserved by Bell s method

’
.

5 The prese n ce distrib u tio n a n d i n some cases characteristic

.
, , ,

distrib u tio n of lipoids i n kid n ey cells s u ggest that they may be

i n timately co n n ected with metabolic processes besides the
possible f un ctio n attrib u ted to them by some of i n fl uen ci n g the
physical state of the protoplasm .

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1 99— 200 , 1 1 t h e di t i on .
 PL A TE 1

F I G UR E X P L A N A TI ON OF ES

1 Kid n y f k i t t en 2 2 m i n l n g t h w i t h l i tt l
e o ,
b d min l ub ut n uc . e ,
e a o a or s c a eo s

fa t. R di l t i n f t i u fi d i n B n d fl u id h w l i p id b l k d b y
a a s ec o o ss e x e e a s
’
s o s o s ac en e

os mi id i n m d u l l y gm n t f p x i m l n v l u t d t u b u l e P h t g p h
c ac e ar se e s o ro a co o e . o o ra ,

X 24 .

3 Kid y ne of kitt en , 19 c m i n l en t h ,
. g v yf
er at , n o t ye t w e a n e d . R dia a l sec t o n i
of i s su e fi x e d i n e n
t B d a s
’
fl idu s h ow s l i p id b l k
o s ac en e d by os m c a om i i n l ab yi r nth .

P h o t o g r ap h X 44 , .

4Kid n y f k i t t n me figu
o 3 T n g n t i l t i n h w l by i n t h
e ,
sa e as re . a e a sec o s o s a r

b l k n d y u n t i n d P h t g p h X 44
ac e e ,
ra s s a e . o o ra ,
.

5 Kid n y f k i t t n 23 m i
e o l n gt h f i m u n t f bd m i n l n d ub
e ,
c . n e , a r a o o a o a a s

cut n a u f t
eo R di l t i f t i u fi d i n 1 0 p
s a . a a n t di h
se c m t nd mi on o ss e x e er c e c ro a e a os c

ac id w i t h u t h t sh w b l k n d m d u ll y gm n t i n y i n t x d
o ea o s ac e e e ar se e s ra s cor e an

c n di n g l i m b
as e f m d u ll y l p i n u t z n f m dul l P h t g ph X 1 6
s o e ar oo o er o e o e a . o o ra , .

6 Kid n y f ki t t n
e m figu
o 5 R di l t i n m fi g u 5 P h
e ,
sa e as re . a a se c o ,
sa e as re . o

t o g r ap h , X 44 .

7 Kid n y e of kitt en , s am e a s fi g u re 5 . T g an i
e n t a l se c t o n sh ow s i me d ull y ar

se gm en t s o f p x im
ro al c o n v olu t e d t ubul e i n c r o ss- s ec t o n i . Ph ot o g ph X
ra ,
44 .
 PL A TE 2

E X P L A N A TI ON OF F I G U RE S
Kid ey f k i t t n m
2 n fi gu o 1 T ng t i l t i n h w
e , ti n
sa e as re . a en a sec o s o s c r o s s- s ec o

o f m dul l ye gm n t f p x i m l nv l u t d t u b u l e y P h t g p h X 44
ar se e s o ro a co o e ra s . o o ra ,
.

8 Kid n y f t 48 5 m i l n g t h v y f t k id n y
e o ca ,
R di l t i f t i u c . n e , er a e . a a sec on o ss e

fi d i n F l mm i g
x e fl u id h w l b y i n t h i n t n el y b l k m d ul l y y u n
e n
’
s s o s a r e s ac , e ar ra s

s t in d
a Ph t g ph X 1 6
e . o o ra , .

9 F m k id y f kitt
ro m ne fi gu 1 T n t i n f t u b ul f m
o en , sa e as re . ra sec o o es ro

ti u fi
ss d in 10 p
e n t di h
x e m t nd t i n d i n u d n I II h w g nu l
er c e c ro a e a s a e s a s o s ra ar

a nd h m g n uo d f o n di n g l i m b
e eo t in d m
s ro tly t p i ph y f w
s o l id a sc e s, s a e os a er er ,
e so

g nu l
ra g n ul i n d n di g l i m b n d ll t i n g t u bu l P j t i n d w
es , ra es esc e n a co ec e . r o ec o ra

i ng , X 800 .

10 Kid n y e of c at , 46 cm . g
i n l en t h , l t t l e i a bd o m n al i or su b c u t an eo u s f at ,
l iv p
er at h o l o gi
Kid n y l i ght i n l
c al . T n t i n f t i su fi d in 1 0 p
e c o or . ra sec o o s e x e er

ce n t dih m t c t i n d i n u d n I II
ro a e, s a S h w v u l i n p x i m l n v lu t d
e s a . o s ac o es ro a co o e

t ubul d n d g nul t i d i n
es , r o s, a n di g l i m b
ra f m d u l l y l oo p
es s a n e P asce n o e ar . ro

j tiec d w i n g X 8 00
on ra ,
.

11 T n ti
ra f tubuls ec f on d i n g l i m b f m d ul l y l p f kid y f
o e o asc en o e ar oo o ne o

p u pp y 2 3 m l n g T i u fi d i n 1 0 p
,
c . o . n t di h m t nd
ss mi
e id n d
x e er c e c ro a e a os c ac ,
a

s h w g
o s ul d t i i n g b l k P j t i d w i g X 800
r an ar r o s s a n ac . ro ec on ra n ,
.

12 S m fi gu
a 10 T n t i n f p x i m l n v l ut d t u b u l n d m d n l
e as re . ra s ec o o ro a co o e e a e

l y ar gm t t h w v u l n d g nu l e t i n i n g w i t h u d n III P
se en o s o ac o es a ra s s a s a . ro

j t i n d w i n g X 800
ec o ra , .

13 T ti n
r an s e cf m d u ll y o g m n t f p x i m l nv l u t d t u b u l n d
o e ar se e o ro a co o e e a

n di n g l i m b f H l oo p C m p fi g u 1 T i u fi d i n 1 0 p ’
a sc e l o en e s nt . o are re . ss e x e er c e

di h m t
c ro nd a e mi i d w i th h t T h
a os c h h w n g nu l
ac d ea . ere ar e er e s o ra ar ro s

s t i ni n g b l
a k i n th n di n g l i m b
ac P j t i d w i g X 1 200
e as c e . ro ec on ra n ,
.

14 T ti n
r an se cf tub ul f k id n y
o Th
o m figu 5 It h w l i n
e o e . e sa e as re . s o s e ar

a rr ang m nt e f l i p id g nu l
e o in th p xim lo nv l u t d t ub u l
ra P j ti nes e ro a co o e e . r o ec o

d w i g X 1 200
ra n , .
LI P O I D CO N T E N T O F T H E K I D NE Y T UBU L E ABL E 2
T
C H RI ST I A N N A M I H
S T
R esu me n por a u tor G eorge S H un tin gto n
el ,
. .

U n iver sidad Col u mbia N ew Y ork

‘

.
,

U nc cri tica de las teor i as de la evol u ci on p ulmo n ar de los

m am i fero s .

El au tor presen ta algun as co n sideracion es sobre la filoge n ia de

p ulm on de los m am i fero s c on especial m en ci on del desarrollo de
,

los tipos arqu i t e ct on i c o s existen tes en los b ro n qui o s y su si gn i fi c a

ci on evol utiva L as co n cl u sio n es co n d u ce n a un a revisi on de las
.

opi n io n es act u ales sobre la o rg an i z aci on i n trap ulmo n ar En el .

prese n te t rab ej o se disc u ten : 1 L a teori a redu c ci on al de A eby

.

( 1 88 0 ) y D H ard i vi ll er ( 1 8 9 7 ) en la c ual comien za el problema

’

modern o de la i n t erpret aci on de los b ronq ui o s 2 L a teor i a de . .

la em i g ra ci on de los compo n e n tes bro n quiales prop u esta por ,

W ill ach ( 1 8 88 ) y N arath ( 1 8 9 2 1 8 9 6 y 3 El res u me n

,
.

de los resultados obten idos por el autor qu e co n stit uye n lo q ue ,

breveme n te p u diera d efini rse como la teor i a selectiva de la

esp ec i ali z a c i on p u lmo n ar de los mam i fero s .

T ra n s l a t i on by J é F
os N o m d ez
g
C a r n e i e I ns t l t u t i o n of Wa s h m g t o n
 I LI O GR A P H I C S
BY 1 1 1 13 B B ER V I CE , A P RI L 19

A C R ITIQU E OF TH E TH EO R I ES OF P U L M O N AR Y
EV O L U TI O N IN T H E M AM M AL IA l

G EO . S . HU N TIN G T O N
Col u mbi a U n i versi ty

FI F TE E N FI G U RE S

The p u blicatio n of A eby s ( 2 ) f un dame n tal work on the ’

M a m mali an Bro n chial T re e i n 1 88 0 laid the gro un d work for the

phyletic i n terpretatio n of p u lmo n ary orga nizatio n D uri n g the ,

fo u r decades which have elapsed si n ce its appearan ce the problem

has attracted the at te n tio n of ma ny i n vestigators The reaso n .

for this i n terest rests on the fact that the morphological co n di

tio n s are complex an d di ffi c ul t to a n alyze from the stan dpoi n t of
their evol utio n ary si g nifi can c e .

The determi n atio n of the fi ve types of the mammalian bro n

chial tree by A eby very soo n drew atte n tio n to the followi n g
cog n ate t 0 p i cs :
1 The ge n esis of the diverge n t forms an d their m u t u al i n ter
.

relatio n .

2 Their possible derivatio n from a commo n a n cestral bro n

.

chial tree by vario u s m o di fi cat i ons of the same d uri n g the phylo
ge n y an d possibly d uri n g the mammalian o n toge n y .

3 The si g n i fi can c e of the n u merical prepo n dera n ce of p u l

.

m on ary asym m etry amon g mammalia with the right l un g the ,

domi n an t an d more highly developed organ .

4 The h om ol o g i z at i o n of the i n divid u al compo n e n ts of the

.

bro n chial tree on the two sides i n asymmetrical l un gs .

5 The phyloge n etic relation of the mammalian l un g to the

.

avian an d reptilian respiratory organs .

1
Ab s t r a c t r ea d at t h e 2 6t h i
se s s o n o f t he A i i
sso c a t o n o f Am i
e r c an A i
n a t om s t s ,

P i t t b gh P
s ur ,
en n .
,
Ap i l 1 9
r ,
1 91 9 .
 GE O . S . HUNTINGT O N

These q uestio n s materially i nfl u e n ced i n varyi n g degrees the

co urse of s u bseq u e n t i n vestigatio n .

A critical a n alysis of the work i n this fi el d d uri n g t h e fo rty

years followi n g A eby s p u blicatio n is therefore perhaps best
’
, ,

u n dertake n by prese n ti n g i n place of a strictly chro n ological

,

sequ e n ce a gro u pi n g of the chief res ults obtai n ed un der provis

,

i o n al headi n gs s u ggested by the m ai n li n es of tho u ght co n cern

i n g the phyloge n etic history of the mammalian l un g which deter
mi n ed the tre n d of the i n vestigatio n s It the n appears that the .

fi v e f un dame n tal q u estio n s listed above elicited fo u r mai n hypo

thetical co n siderations of the problem The evide n ce on .
,

which these theories were based is commo n gro un d b ut the , ,

co n cl u sio n s reached are diverge n t O n the assumptio n of a .

commo n a n cestral bro n chial pattern of the mammalian l un g ,

from which all exta n t types are desce nded by evol u tio n ary modi
fi cat i o n s the q u estio n arose which of the existi n g types co n formed
,

most cl osely to this promammalian bro n chial tree an d he n ce ,

represe n ted the n earest approach to the hypothetical primitive

mammalian l un g It appeared probable o n ge n eral ge n etic
.
,

gro un ds that the an cestral l un g of the mammalia possessed

,

more or less complete bilateral symmetry and that he n ce the ,

choice lay betwee n the bilaterally symmetrical eparterial type

( fo un d i n some pi nnipede carn ivores an d cetacean s the Cebidae ,

amo n g primates certai n rode n ts an d perissodactyls elephas

, , ,

hyrax ,
a n d the bilateral symmetrical hyparterial bro n

chial tree of some Hystricomorphs and of T ax i dea These c on .

sideratio n s gave rise to three i n terpretatio n s .

I . REDU C TI O N THE O R Y
A by e ( 1 87 8 ,
1 88 0 , 1 88 2 )
D H a rd i v i l l e r ( 1 896, 1 8 97 )
’

B r em er ( 1 9 04 )

A eby ass umed ( 2 ) that at on e period i n the evol utio n ary his
tory of the mammalian l un g the organ possessed a bilaterally
symmetrical bro n chial g ro un d plan comprisi n g within its scope
-
,

all the compo ne nts fo un d amo ng the exta n t mammalian types .

He held th at this archeal gro un d plan was preserved un chan g ed

-
1 02 GE O . s . HUNTINGT O N

T his vesicle is s urro un ded by co n de n sed mese n chyme rep an d

re se nts i n d H ardi viller s opi n io n the anlage of a separate lobe

,
’ ’
,

of the left l un g Th e p u lmo n ary arteries desce n d symm et ri

.

cally o n each side ve n tral to both the right eparterial b ud an d

,

the proximal b u d from the left st emb ron ch us a n d the n t ur n ,

dorsad to follow the dorso lateral s urface of the st em b ro n ch i -

.

Both the right an d left cran ial b u ds tu r n dorsad after arisi n g

from their respective bro n chi Possessi n g ide n ti c al o rigi n di rec .
,

tio n an d relatio n to the p ulmo n ary artery th ey ap p ear as ab so

,

l u te morphological ho m olog u es D H ardi v i ll er he n ce co n cl udes .

’

that the l un gs of the rabbit begi n their developme n t as b i l at er

ally symmetrical organ s the bro n chial tree co n form i n g to the
,

bilaterally symmetrical eparterial form ( A eby s type I ) The ’ A

.

asymmetry is acq u ired d uri n g s u bseq u e n t developme n t The .

right eparterial a n lage co n ti nu es to exte n d b ut the left ep ar ,

t eri al vesicle becomes grad u ally red u ced an d fi n ally disappears

altogether In an embryo of 1 3 days 8 ho urs he co n ti n u es to
.

fi n d a vesicle attached to the cra nial part of the left st em b ro n

ch u s b ut its walls have become greatly thicke n ed A t 1 3 days
,
.

1 5 ho u rs the co nn ectio n with the left st em b ro n c h u s hollow i n the ,

precedi n g stage has bee n red u ced to a solid pedicle an d at 1 4

, ,

days there is left o nly a slight mesoderm al co n de n satio n at the

site formerly occ upied by this ephemeral left eparterial b ud .

D H ardi vi ll er bases the followi n g co n cl u sio n s o n the above

’

alleged observatio n s :
1 In the rabbit each l un g at fi rst carries a n eparterial bra n ch
.

of the st em b ro n ch us The left eparterial b u d follows for a

.

short period the same co u rse as the homologo u s b ud of the right

side It the n atrophies disappea ri n g e n tirely and the t w o
.
, ,

l un gs which were origi n ally symmetrical become seco n darily

, ,

asym m etrical .

2 The three characteristic bro n chial types established by

.

A eby i n the m a m malia have o n ly a seco n dary v a u e ( S i c "

’

)
‘ ’

The importa n t factor lies i n the existe n ce of a n eparterial ele

me n t o n each side This may persist i n developme n t or atrophy
.
,

o n the left side o n ly or o n both sides th u s givi n g rise to the

, ,

three pri n cipal mammalia n types Viz ,

 P ULMONARY EV O LUTI O N IN M M A MAL I A 1 03

Bilateral eparterial type

I . .

II R ight eparterial left hyparterial type

.
,
.

III Bilateral hyparterial type

. .

3 The left eparterial an lage is n ot a side bra n ch of the fi rst

.

ve n tral ( hyparterial ) bron chus ( N arat h ) .

4 The bilateral embryo ni c eparterial a n lage n explai n the

.

h uman bro n chial variation s

It is c urio us an d somewhat amu si n g to fin d D H ardi vill er so
, ,
’

c a rrl ed away by the importa n ce of his ow n discovery that he

assign s o n ly a secon dary val u e to A eby s bro n chial types

‘ ’ ’
,

from which his own work derives its chief si g ni fi can ce as c on

firm i ng the predictio n made by A eby seve n tee n years before .

But aside from this rather i n terestin g exhibitio n of sci en t i fic

temperame n t the embryological evide n ce presen ted by d H ardi
,
’

Viller wo uld have afforded absol ute an d co nvi n ci n g proof of the

correct n ess of A eby s thesis if s ubstan tiated by other observers
’
,
.

It is a matter of regret that the asto nishi n g revelatio n s of

d H ardi v ill er differin g from the fi n di n g s of n early all other i n
’
,

v e st i g a t or s w h o have st u died the problem of mammalia n p ulmo

n ary developme n t have n ot i n the twe n ty three years which have

, ,
-

elapsed si n ce their p ublicatio n received the atte n tio n which they

,

wo uld ab un dan tly merit if c on firm ed N arath ( 33 ) an d F li n t .

( )
2 1 are the o n ly i n vestigators who have serio u sly co n sidered
d H a rdi V i ll er s claims The reaso n for this is to be fo un d i n the
’ ’
.

fact that whe n N arat h i n his large mo n ograph p ublished i n

, ,

1 9 0 1 appeared as the chief s u pporter of W ill ac h s ( 3 7 ) M igratory

’
,

Theory s ubsequ en tly to be co n sidered i n detail (p

,
he .

fo un d himself co n fro n ted by d H ardi vil ler s observatio n s which’ ’

, ,

if valid demolished at on e stroke the mai n con te n tio n of the

,

theory he s upported and defini t ely proved A eby s hypothesis

,
’
.

N arat h s chief p ublicatio n embodyi n g the res ults of careful work

’
,

exten di n g over a nu mber of years ( 1 8 9 2 followed so closely

u po n the a nn o u n ceme n t of d H ardi V ill er s fi n di n g s that he co u ld
’ ’
,

n o t or at an y rate did n ot s u bj ect them to an i n depe n de n t r e

, ,

visio n H e he n ce n at urally accepted them at their face val ue

.
,

an d so fo u n d himself take n at an un fair adva n tage i n meeti n g

them H e based his chief arg ume n t again st the co n cl usio n s

.
1 04 GE O . s . HUNTINGT O N

draw n by the Fre n ch observer o n the cases of human bro n chial

va riatio n s reported by Chiari ( 6 7 8 9 ) an d D alla R osa ( 1 0) , , ,

with do uble right eparterial bro n chi i n which the additio n al ,

bro n ch u s arose either from the st em b ro n chus or trachea N arat h .

s uggested that an alogo u s varian ts if they developed on the left ,

side wo uld suffi ci ently acco un t for d H ardi vill er s left eparterial
,
’ ’ ‘

vesicles H e f urther cited two i n stan ces i n the ad ult rabbit i n

.
’

which the upper lobe of the left l un g n ormally supplied by the ,

asce n di n g b ran ch of the fi rst ve n tral hyparterial bro n ch us re ,

c ei v ed an additio n al bra n ch arisi n g from the st em b ro n ch u s n earer

to the tracheal bif urcatio n and dorsal to the p ulmo n ary artery ,

th us repeati n g i n atypical ad ults the temporary co n ditio ns de

scribed by d H ardi V i ller for the n ormal embryo
’
.

In a series of 7 0 ad ult rabbit l un gs which I exami n ed by cor ,

r o si on for the occ u rre n ce of bro n chial a n d arterial varia n ts I ,

fo un d i n o n e i n divid u al the cran ial pole of the left l un g s upplied

by an atypical fi rst bran ch of the left st emb ron chus arisin g dorsal
to the p ulmo n ary artery an d correspo n di n g i n position to the
,

larger eparterial bro n ch u s of the right l un g I have examin ed .

with the u tmost care all the series of rabbit embryos of the c ri t i
cal stages betwee n 1 2 an d 1 5 days an d from 8 to 1 1 mm i n
, ,
.

le n gth which were available to me These i n cl ude the series of

,
.

the Col umbia collectio n an d a nu mber of preparatio n s le n t by

my colleagu es i n other i n stit u tio n s N ot i n a si n gle i n stan ce .

has it bee n possible to detect eve n an approach to the con di

tio n s described by d H ardi V i ll er A very large an d closely graded
’
.

series of cat embryos i n the Col umbia U n iversity Collectio n ,

with numero u s i n divid u als of the same stage an d a similar , ,

tho ugh less exte n sive series of embryos of the albi n o rat which ,

I owe to the liberality of The Wistar I n stit ute have given the ,

same n egative res ults I do n ot believe it possible that these

.

preparatio n s can fail i n represe n ti n g correctly the tru e morpho

logical co nditio n s obtai ni n g i n the critical stages of these mamma
lian embryos and I he n ce do n ot hesitate to pro n o un ce d H ardi
,
’

viller s co n cl usio n s erro n eo u s an d his ge n eralizatio n un warra n ted

’

o n the evide n ce D H ardi vi ller does n o t me n tio n the number of

’
.

embryos i n which he fo un d his eva n esce n t vesicle The co n text .

1 06 G EO . s . HUNTINGT O N
Th at on l f t si de i s always sm aller and sli ghtly l ower pl a ce d
t he e
,

a n d t he a i ch am b o s su ppl ie d by it d o n o t f orm t h
r— r a pex o f the l u n g e ,

s t ill i n s pite of i t s m all size an d re l atively l ow p o siti o n it i s d i sti n ct l y

s ,

ab o ve the fi st ve n t al b o n ch u s an d behi n d the artery an d so c o rre

r r r

s p o n ds t o t h e e p arte i al b o n ch u s o f the ri ght l u n g an d m ay be c on s i d

r r ,

e e d as m akin g t he t w o lung s symm etri cal and re ptili an i n type as n o

r
,

p l ac e n t al l u n g s are .

the older po u ch stages ( 1 4 c m ) and i n the ad ult a n imals

In -

“
.

the oposs um l un g is described (p 7 2 ) as cha n gi n g from the rep

—
.

tilia n to the mam m alian also i n the loss of the left eparterial
‘

bro n ch u s Bremer is un able to state how this loss is bro u ght

.

abo ut owi n g to the lack of the req u isite i n terve n i n g stages i n

,

his material These observatio n s ra n ge themselves therefore i n

.

s upport of d H ardi V i ller s claim disc u ssed above that i n the

’ ’

m ammal (L ep u s) the left st em b ro n ch u s carries at o n e stage of

its developme n t an ephemeral eparterial bro n chial a nlage which
d uri n g the period of its temporary existe n ce re n ders the right
a n d left l u n gs symmetrical eq u ivale n ts They therefore call for .

caref ul co n sideratio n here si n ce they co n stit u te the o nly ev i

,

de n ce c onfi rm at ory of d H ardi V i ll er s observatio n s which has

’ ’

bee n offered i n the e n tire literat u re They he n ce fall withi n the .

scope of this paper i n a disc ussio n of the R ed u ct io n T h eory .

In his admirable acco u n t of the developme n t of the l u n gs i n

the pig p ublished i n 1 906 Fli n t ( 2 1 ) co n siders Bremer s fi n di n g s
,
’

from two importa n t sta n dpoi n ts :

H e calls atte n tio n ( p 22 ) to Bremer s stateme n t that n o pla
’
.

ce n tal l un gs are sy m metrical i n the possessio n of a bilateral

eparterial developme n t an d q u otes i n ref utatio n the lo n g list of
mammalian forms ( si n ce materially i n creased ) p ub i shed by ’

A eby N a rat h ( 33 ) an d myself ( 2 5) i n which the bilateral

eparterial bro n ch u s is n ormally fo un d .

Fli n t also q uestio n s the acc uracy of Bremer s ass umptio n that ’

the bilateral eparterial developme n t is a reptilia n character a nd

cites a perso n al co mm u n icatio n from Hesser ( 2 2 ) wh o was un able
to detect a ny eparterial bro n chial developme n t i n his reptilia n
material Fli n t co n cl udes his co n sideratio n of the q uestio n with
.

the followi n g stateme n t ( p 1 1 6) .

 P ULMONARY EV O LUTI O N IN MAMMAL I A 1 07

thu s h a dly po ssible i n these ob serv ati o ns o f d H a diV iller an d

It i s r
’
r

B emer th at w e a e de ali n g with a true reg re ssive p o c ess In f ac t

r r r .
,

i t i s m o e p ob able th at i n b o th c a se s w e a e eithe d e ali n g with a

r r r r

v i ti o n o a d orsal b on chu s whi ch i s pl a ce d hi ghe u p th an u su al

ar a r r r

u p o n t h e st em b ro n c h u s Thi s assum p ti o n i s m ade q uite prob abl e by

.

B em e s st atem en t th at h i s e p a teri al bro n chu s di d n o t su p ply t h e apex

r r
’
r

of t h e l un g .

I c an e n dorse this co n cl usio n absol utely Thro ugh the co u r .

tesy of Professor M c Clure I have had the opport un ity of st udyi n g

a n d reco n str u cti n g the l u n gs i n two u teri n e embryos of Dtdel

p hi s a n d i n o n e u teri n e e m bryo of D a s yu r u s co n tai n ed i n the ,

embryological collectio n of P ri n ceto n Un iversity N o trace of a .

left eparterial bro n ch u s was fo un d i n an y of these While these .

e m bryos an d their reco n stru ctio n s will be described an d figured

f ully i n a more exte n sive forthco m i n g p u blicatio n deali n g with
the m orphological details of the mars u pial l un g I appe n d here ,

two views of the model of the bro n chial tre e an d p ulmo n ary
artery i n the 1 0 mm u teri n e embryo of Di delp hi s marsu pi a lzs
’

. .

The architect u ral pattern of the l un g co nforms e n tirely to the

domi n an t place n tal type with the a n lage of the eparterial bro n
,

ch u s restricted to the right side The left st em b ron chu s does .

n o t carry a correspo n di n g compo n e n t b u t is e n tirely hyparterial ,

i n its distrib u tio n A fter givi n g o ff its cra n ial eparterial deriva
.

tive (Ep ) i n the typical sit u atio n the right st em b ron chu s carries
.

three well developed ve n tro lateral hyparterial bro n chi an d the -

less far adva n ced a nlage of the fo urth ( V V ) The cardiac l— 4

.

bro n ch us with three laterals an d the expa n ded en d b u d is

, ,

derived from the ve n tro medial aspect of the right st emb ro n ch u s

-

at the level of the caudal margi n of V The left st emb ron ch u s I

.

gives origi n to fo ur ve n tro laterally directed hyparterial bro n —

chi ( V V ) slightly less far adva n ced tha n the correspo n di n g

l— 4
,

b uds of the right side Th e A sce n di n g Bra n ch (A ) of left V

.
I

exte n ds cra n iad to the level reached by the eparterial compo n e n t

of the right side Each s upplies the ho m ologo u s apical portio n
.

of its l un g T here is n o i n dicatio n of a left eparterial b ud

. .

T he series of the dorsal bro n chi co n tai n s fo ur disti n ct compo

n en t s o n each side ( D
l
1 08 G EO . s . HUNTINGT O N

arises from the cau dal border of the eparterial

R i g ht lu n g : D 1

bro n ch u s It carries three mai n b uds directed mesad dorsad

.
, ,

a n d laterad each st u dded with several smaller evagi n atio n s

, .

Fig . 1 U t c rm e em by r o of Di del p hi s ma r s u pi a l z s , L . 10 mm . Pi t U
r n c e on ni

v i t y Emb y l gi l C ll t i
e rs r o o ca o ec on . R i
e c o n s t ru c t o n o f l un g an d p lm y
u on a r a rt

X 1 50 V t l v i w Ep e p a rt e ma l b ro n c h u s ; A a s c en di g b h of
“v
er y . . en ra e . .
,
.
,
n ra n c

fi t l f t v t l h yp t i l b
rs e en ra a r er a ro n c h u s ; V l-
V ,
en t ra l h yp a r t e r ali b ro n c h i; Cd .
,

ri gh t di b h
car ac ro n c us .

is the largest member of the dorsal series an d prese n ts the

D 2

same t ri p art i d un foldi n g .

D smaller shows a red u ced trefoil patter n

3
, ,
D is a si n gle .
4

dorsally directed b u d .
1 10 G EO . s . HUNTINGT O N

D idelp hi s specime n follows the same plan of bro n chial organ i

,

z at i o n ,
with the eparterial b ut limited to the right l un g .

In compari n g these reco n str u ctio n s with Bremer s acco u n t ( p

’
.

7 1 ) a n d with the diagram of his mm oposs u m l un g (p 7 2 . .

,

fig 1 0 ) I am obliged to co n cl ude that the left eparterial bro n

.
‘

ch u s described an d fig ured by him is the cran ial termi n al of the

’

fi rst left dorsal bro n ch u s a n d that the l un gs of Di delp hi s as well

, ,

as those of all other mars upials whose architect ure has bee n ‘

determi n ed do n o t depart either i n their o n toge n y or i n their

,

ad ult organizatio n from the pattern fo un d i n the domi n an t pla

ce n tal type O therwise it wo uld be n ecessary to adopt the
.

u n te n able View that the u teri n e embryo of Di delp hi s develops

the asymmetrical bro n chial an lage with on ly right sided ep ar ,

—

t eri al bro n ch u s a n d that i n yo u n g po u ch embryos betwee n

,
—

an d mm a left eparterial compo n e n t appears o n ly to

.
,

be lost i n older po u ch yo un g a nd the ad ult th u s agai n restori n g

—
,

the pri m itive asymmetry This is farther afi el d than I am will

.

i n g to follow the s u pporters of the R ed u ctio n Theory o n the evi

de n ce th u s far add uced However this as well as ma n y other
.
,

q uestio n s of i n ten se morphologic i n terest still await the opp or

tu n i t y of st u dyi n g large an d closely graded series of well fi x ed —

u teri n e embryos a n d early po u ch stages of the oposs u m I am -

.

still i n hopes that the rich embryo nic material of Di delp hi s

sec ured several years ago for The Wistar I n stit ute thro ugh the
e n ergy an d skill of D rs Hartman an d He u ser may eve n t u ally
.

become accessible to q uali fied American i n vestigators for the

p urpose of defin i t el y a n sweri n g the importa n t morphological
problems the sol utio n of which is locked up i n the mars upial
,

o n toge ny .

It is of co urse co n ceivable as i n the case of d H ardi viller s,

’ ’

p ublicatio n that all the i n divid uals composi n g the yo un gest

,

litter of Di delp hi s embryos st udied by D r Bremer possessed an .

aberra n t bro n chial orga n izatio n A gai n st this ass umptio n is the .

fact that cardi n al bro n chial varia n ts of the ad ult are excessively
rare thro ugho ut the e n tire mars upial s uborder as compared ,

with the place n talia N arath (3 3 p 3 2 6) fo un d n o i n sta n ce of

.
,
.

the left eparterial ( apical ) bro n ch u s i n 1 6 represe n tatives of 9

 P ULMONARY E V O LUTI O N IN MAMMAL I A

mars upial ge n era I obtai n ed correspo n di n g res ults n ot en

.
,

co unteri n g a si n gle left eparterial bro n ch u s i n my e n tire series of

mars upial l un gs the details of which are i n cl u ded i n a forth
,

comi n g p ublicatio n My material comprised 8 9 examples of

.

1 8 mars u pial ge n era distrib u ted amo n g 4 6 species a n d i n cl u di n g

,

2 9 i n divid u als of Di delp hi s assig n ed to 8 species with 1 7 S peci ,

me n s of D marsup i ali s ( vi rgi ni ana) I therefore believe that

. .

Bremer s i n terpretation of his

’
mm oposs u m embryo is at .

fau lt i n disregardi n g the fact that i n its typical co urse the left
p u lmo n ary artery of this f orm exte n ds cau dad i n the i nterval
betwee n the rows of primary ve n tral an d dorsal derivatives
from the st em b ron chu s i n all embryo n ic stages so that a dorsal ,

bro n ch u s is always placed behi n d a ve n tral always i n fro n t ,

of the artery at any give n level of the lun g st em .

In t h e p ass a g e q u oted above Bremer lays stress on the o b ser

'

vatio n that the left bro n chial eleme n t i n terpreted by him i n the
mm Di delp hi s embryo as an eparterial a nlage arose dis
.

t i n c t l y cra n ial to the derivatio n of the fi rst ve n tral bro n ch u s as ,

well as behi n d the artery This high origi n of a left first dorsal
.

bro n ch u s is very u n u su al In all of my reco n stru ctio n s of

.

yo un ger po u ch embryos (3 ) an d i n the corrosio n s of older stages

the bro n ch u s has i nvariably arise n disti n ctly cau dal to the level
of V Its more cran ial origi n described by Bremer m u st I
I
.
,

believe be co n sidered a varian t as already s u ggested by Fli n t

, ,

which may have wro n gly sim ulated a left eparterial an lage .

A s the co n cl u di n g li n es of the passage q u oted above i n dicate ,

Bremer regards the reptilia n l u n g as possessi n g a bilateral

eparterial system an d di ffere n tiati n g i n this regard from the
mammalian types Both of these ass umption s are i n correct
. .

Man y mammalian ge n era an d species are n ow k n own to have

typically a bilaterally symmetrical eparterial bro n chial organ i
z at i o n . O thers show i n i n divid u als i n additio n to the n ormal
right eparterial bro n ch u s a correspo n di n g derivative from the
left st em b ron chus as a varian t acqui sitio n In both grou ps the .

eparterial com po n e n ts are disti n ct mammalian n eomorph de

v el op m en t s a n d n o t reappeara n ces of a n archeal reptilia n char -

acter The assumptio n that the moder n reptiles possess a

.
1 12 G EO . s . HUNTINGT O N

symmetrical eparterial un foldi n g i n both l un gs is based on the

very few prophetic remarks with which A eby ( 2 ) co n cl udes his
mai n work o n the mammalian bro n chial tree H e there ( p 9 6) . .

merely expresses the expectatio n that the co n trast betwee n the

eparterial and hyparterial bro n chial derivatives determi n ed by

Fi g . 3 A s p i don ec tes sp s er , Le S ue u r . C o r ro s i o n of r i ght l un g an d pu l

mc n a r y a rt e r y . V en t r a l vi ew . C o l u m bi a M o r p h Mu s . .
,
no . 1 93 1 .

him for the mammalia wo uld be fo un d i n the reptilia represen ted

by a tran sitio n i nto parallel lo n git udi n al rows of bro n chi ,

placed o n the medial an d lateral side of the p ulmo n ary artery .

F urther s uggestio n s alo n g this li n e he co nsiders unwarranted i n

view of his lack of material .

Z umstei n one of the earliest i nvesti g ators to a nalyze A eby s

_
,
’

View critically reported (3 9 ) on several reptilia n l un g corro

,
1 14 G EO . s . HUNTINGT O N

de n t relatio n rests upo n the commo n morphoge n etic and fun c

t i o n al factors which li n k the reptilia n respiratory tract to the
early phases of its adaptatio n to mammalia n req u ireme n ts .

This does n ot i nvolve a hypothetical reptilian eparterial bro n chial

system which does n ot exist Whe n it appears i n the mamm a
.

lian li n e it does so as a n ew acquisitio n i n respo n se to very

d efi ni t e e n viro n me n tal an d f un ctio n al factors
With d H ardi V ill er s an d Bremer s positio n i n the matter
’ ’ ’

d efin ed it is possible to ret u r n to the critical co n sideratio n of

,

A eby s R ed u ctio n Theory on its ow n merits

’

This theory i n volves the acceptan ce of the followin g dogmata

1 The primitive a n cestral mammalia n l un g was i n its bro n
.
,

chial architect ure a bilaterally sym metrical organ with the

, ,

eparterial bro n chus developed i n both l un gs and arisin g from

the stem ( A eby s type I )’ A
.

2 The red u ctio n has c hi efl y affected the crani al portio n of

.

the left l un g e n tailin g the phyloge n etic s uppression of the left

,

eparterial bro n ch u s an d res ulti n g i n the establishme n t of the

,

domi n an t mammalian type with the eparterial bro n chu s lim

i t ed to the right l un g ( A eby s type H )
A ’
.

3 O n ly relatively few forms have retain ed the origi n al bilat

.

eral eparterial organ izatio n ( A eby s type I ) A ’

.

4 A still more limited nu mber of livi n g mammalia have

.

u n dergo n e red u ctio n also of the right l un g th u s establishi n g the ,

bilateral hyparterial distrib u tio n of A eby s typ e III ’

.

An a n alysis of these theses provokes the followi n g critical

co n sideratio n s
1 The ass umptio n of a bilaterally symmetrical bro n chi al tree
.

i n the most primitive mammalia or promammalia rests o n p u re

hypothesis an d is i n capable of direct proof With the elimi n a .

tio n of the evide n ce offered by d H ardi V i ll er an d Bremer t he’

,

o n toge n y of the mammalian l un g does n ot throw the light on

the problem which was c on fiden t ly anticipated by the au thor of
the theory The type of bro n chial architect ure characteristic of
.

a n y give n mammalia n species ma n ifests itself u nmi stakably i n

the earliest p u lmo n ary an lage of its embryos as soo n as the dif
feren t i at i o n of i n divid ual bro n chial compo n e n ts appears .
 P ULMONARY EV O LUTI O N IN MA MMAL I A 1 15

2 . A ss umi n g
that the domi n an t type of the l un g i n modern
mamm alia with elimi n ation of its left eparterial compo n en t re
, ,

cei v ed n o compe n satio n for this loss by a u gme n tatio n i n other

direction s the primitive l un g i n respect to expan se of respira

, ,

tory area an d fun ction al effi ci en cy attai n ed a higher degree of

,

developme n t than is exhibited by the vast maj ority of extan t

types .

3 A s far as act u al evide n ce is co n cer n ed this ded u ctio n from

.

the ten ets of the R ed u ctio n Theory is opposed both by p al aeon

t ol o g i cal an d comparative an atomical evide n ce The li n e sepa
.

ratin g the ge n eral repti lian an d mammalian organ ization s of

to day is n owhere more sharply drawn than i n the disti n ction
-

betwee n p oi kilo an d homoeothermal vertebrates which rests i n ,

the last an alysis u po n the di ffere n t ratio of tissu e comb u stio n i n -

the two groups This i n t u rn is determi n ed by the differe n ce

.

i n the rate of the respiratory metabolism an d is depe n de n t upo n

the degree of str u ct ural developmen t attai n ed by the respira
tory organ s It is scarcely co n ceivable that the desce n dan ts of
.

a reptilian an cestry with relatively low p ulmo n ary organiza

,

tio n sho uld e n ter the path of mammalian evol u tio n with s u ch a
,

s uperab un dan t respiratory e n dowmen t as to n ecessitate its re

d u ction i n the course of their f urt her advan ce O n the other .

han d the weight of the comparative an atomical data leads to

the co n cl usion that the bilateral eparterial bro n chial develop
men t is favored if n ot gen etically co n tin ge n t u po n certain en
, ,

v i ro nm en t al an d f un ctio n al factors s u ch as aq u atic habitat an d

,

res u ltan t periods of very active respira tio n altern ati n g with
s u spe n sion of the fun ction d urin g s u bmersion great b ulk of the
,

m u sc ul at u re rapid or lon g co n ti nued locomotio n etc All these

, ,
.

may have played an importan t part i n associatio n with a higher

p u lmo n ary organization evide n ced by the more exte n sive ep ar
,

t eri al developme n t of the cetacean s pi n n ipede carn ivores an d

,

un g ulates Were these or simi lar e nviro n men tal i nfluen ces de
.

oisive i n the evol u tio n of the a n cestral mammalia an d acco u n t ,

able for their high degree of p ulmo n ary expan sio n ? The R edu c
tio n Theory would deman d that the early mammalia were b ulky ,

aquatic forms wi th a p u lmon ary u nf oldi n g so far i n excess of the

116 G EO . s . HUNTINGT O N

requireme nts of their modern desce n dan ts that i n the maj ority ,

of these a very co n siderable red u ctio n of the respiratory area

was made possible an d act u ally e ffected i n the co u rse of ev ol u
tio n leavi n g the relatively small number of exta n t bilate rally
,

eparterial forms as represe n tatives of a precedi n g archeal phy

leti o stage The palaeo n tological evide n ce beari n g on the origin
.

of the mammalia declares the co n trary to have bee n the case and
characterizes the earliest mammalian or promammalian types as
exceedi n gly small a n imals abo u t the size of a shrew with ter
, ,

r est ri al
, probably arboreal habitat Ma n dib ular rem n an ts an d
,
.

teeth form practically the o n ly evide n ce of their presen ce dur

i n g the e n tire geological period of e n ormo u s le n gth exte n di n g
from the mesozoic to the upper cretaceo u s .

Palaeozoologists ascribe this lo n g co n ti nu ed s uppression of

the mammalian stem to u n favorable e n viro n me n t chi efly due to ,

the domi n ation of the mesozoic reptiles In the u pper limits of .

the cretaceo u s period man y of the large reptilian phyla appear

to have bee n s u dde nly destroyed an d i n co n seq u e n ce the mamma
,

lia began a very rapid co u rse of evol utio n ary developme n t an d

advan cemen t The earliest well k n ow n faun ae of place n tal
.

mammalia date fro m the begi nn i n g of the Tertiary period (basal

an d lower eoce n e ) In the i n sectivora primates car n ivora
.
, , ,

co n dylarthra amblypoda ede n tata the most primitive forms

, , , ,

leas t specialized i n de n titio n an d limbs are always small ani ,

mal s of t erresti al habitat The most primitive i n sectivores

.

were eve n mi nute an d shrew like i n form the most primitive

-
,

carn ivores an d archaic u n gulates abo u t as big as a large op os

sum . G regory an d Mathew hold that the immediately an ces
tors of the early E oce n e faun ae were arboreal types more or
less resembli n g the modern oposs u m i n appearan ce an d habits .

The palaeo n tological evide n ce as to the origi n of the M arsu

palia is sle n der b ut s upports the opi nio n s of H uxley D allo an d
, , ,

Be n sley that both the P olyp ro d ont an d Di prodon t types have

,

bee n derived from small arboreal forms resembli n g the Bi del

p hyi dae i n most characters The m arsup al i a as a gro up app ear
.

to represe n t en dfo rm s of a mesozoic pre place n tal stock of low -

brai n str u ct ure an d more or less arboreal habitat .

1 18 G EO . s . HUNTINGT O N

These are mostly characterized either by great bodily b ulk ,

heavy m u sc ulat u re rapid or lo n g co n ti nu ed locomotio n or by

,
-
,

aq u atic life with its res ulti n g effect on respiratory orga n izatio n
, .

The earliest mammalian or promammalian types as stated , ,

were exceedi n gly small lacki n g b ulk an d mu sc ular developme n t of

,

the modern un g ulate They were lan d forms the mari n e adap
.
,

t at i on s of the cetacea an d sire n ia were acq u ired i n later geologi

cal periods The pinn ipede carn ivora represen t a still later an d
.

very limited adaptation to aqu atic life .

3 Am o n g extan t mammalia the o n toge n y of the mo n otreme

.

an d mars u pial lu n g 3 0 3 2 3 5) fur ni shes exceedi n gly

, ,

importan t an d defini t e eviden ce of the phylogen etic desce nt of

the primitive mamm alian respiratory tract from a reptilian
an cestry Comparison of early embryos i n the egg or the uterin e
.

stages with the later p ulmon ary anl ages after hatchin g or dur
i n g the po u ch period clearly bridges the gap betwee n adul t
,

reptilian an d mamm alian organ izatio n ( cf p 1 9 2 ) b ut fails to . .

show any eve n tu al departure from the typical mammalian plan

i n the o n toge n y of the bro n chial tree i n these forms .

The mo n otremes i n partic ular stan d morphologically i n closer

relation to the promammalian reptilian an cestry than any other
extan t mammalian type If the earliest mammalia already pos
.

sessed the bilateral eparterial expan sion of the bro n chial tree ,

it is reaso n able to expect its reten tio n if an ywhere i n these , ,

most primitive members of the class O n the con trary the .

cran ial segme n ts of the mo n otreme l un g are striki n gly u n der

developed Both Echi dn a and P latyp us possess the domi n an t
.

mammalian asymmetrical type of bro n chial distrib u tio n The .

very small eparterial bro n chu s is c onfined except i n varian ts , ,

to the right side s upplyi n g a small apical process of the right

,

l un g which appears to be scarcely more than a r u dime n tary

,

appe n dage to the l un g stem The mo n otreme l un g i n stead of

-
.
,

s upportin g the R ed uction Theory s uggests rather the first un ,

foldi n g of the cran ial l un g poles leadin g eve n t u ally to the

exte n sio n of the n eomorph eparterial districts .

4 In completi n g this review of the R ed u ctio n Theory it is

.

fin ally of i n terest to co n sider the gro un ds on which A eby selected

 P ULMONARY EV O LUTI O N IN MAMMAL I A 1 19

the bilateral eparterial bro n chial tree as represen ti n g the primi

tive mammalian con dition from which he derived the remai ni n g
extan t types by redu ctio n The reason s for adopti n g t hi s vi ew
.

appear i n adequ ate an d n o t i n accordan ce with the facts It is .

,

however o nl y j u st to remember that A eby s pri n cipal effort w a s

,
’

directed toward the con stru ctio n of the firm morphological basis
u po n which fu rther i n vestigators wo u ld be able to b u ild up the

details of the p ulmon ary problem H e clearly recog nized the .

valu e of his discoveries to phyloge n etic i nterpretation b ut n ot ,

on ly refrain s himself from drawi n g premat ure detailed i n fere n ces ,

b u t distin ctly cau tio n s agai n st s u ch an attempt .

In disc u ssi n g the distrib u tio n of his three bro n chial types an d
their two s u bdivisio n s amo n g the mammalian orders he says (2 , ,

pp 9
. 1 1)
—

V on beson derem Intere sse sin d diejenigen O rdnurig en deren G lie der ,

v er schie den en L ag ern an g eb oren H ier i st w ohl au ch fur weitere

.

F orschun g en i n er ster L in ie der H ebel an z u setzen um u erf ahren i n

'

,
z ,

wel che Ve t hei lun g un d m i t wel chen Ab an derun g en der A n schluss an

r r

di e ein e o der an dere S eite erfol g t S o l an g e die s n i cht g e schehen

.
,

erscheint es v erfrii ht 1 11 allf all i g e phyl og enetische Erwag ung en ei n ut

,
z

reten u n d m uss es bei der einfachen Th at sache sein B ewen den h aben
, ,

d ass P fer d u n d T apir u n ter den P eri sso d act yl en ni cht ein i g g ehen u n d
dass d as L am a g eg en ii b er den an dern Ar t i od a ct yl en d as S t achel ,

s chwei n g e g enuber d en N ag ern das F aul t hi er g e g enuber sei n en z ahn

,

l osen G en ossen ein e S on derstellun g einnimm t L ei der i st es mi r trotz

.

Vielf acher Bem ii hung en ni cht g elun g en n ach dieser S eite hi n m ein
,

U n ter su chun g sm ateri al u erweitern z .

O cc upyi n gthis co n servative stan dpoin t A eby appears to have

li n ked up his fiv e bro n chial types almost at ran dom His choice .

of the bilateral eparterial tree as the primitive form which led ,

to the establishmen t of the R ed u ction T h eory was perhaps ,

determi n ed by the fact that the co n verse symmetrical bilateral

hyparterial type was k n own to him i n o nly one species of the
Hystricomorphs H ystri x cri sta ta
,
.

H e says p 1 2) .

Auf dem j etzig en S t an dpunkte der M orphol og ie erscheint es wohl

kaum al s zweif elhaft dass di e v erschie denen F orm en des B ron chi al
,

b aum s z u ein an der i n g e n eti scher B eziehu n g stehe n Wel che v on .

1 20 G EO . s . HUNTINGT O N
ihn en i st d i e p rim are ? N ach a llem
twi ckl ung sg eschi chte ,
w as d i e En
d er O g an e bi sher u T ag e g ef ci r d er t si cher ei n e d er sym m etri s che n
r z , .

Zwi schen d en bei de n di e ur Verfii g ung stehe n wird di e W ahl dadur ch

,
z ,

we sentli ch erlei chtert d ass i n de ein zig en Lung e wel che d er epar
,
r ,

t eri ellen S t o ckwerke e n tbehrt au ch di e hypart e i ellen ei n e h o chg r ad i g e

,
r

V erk um m eru n g erlitte n h abe n D a dur ch g ewi nn t d i e g an ze B il dun g

.

d en C h a akter ei n e R edu c t i on sf orm deren A usg ang spu nkt au f breiterer

r ,

G run dl ag e g esu cht wer d en m u ss Ei n e sol che i st der B r o n chi alb aum
.

m i t ep art eri ell en Br on ch en i n bei de n L u n g en Au ff alli g bleibt d abei

'

i m me hi n dass g er ade diejeni g en S aug et hi e e bei denen man am eb es

r ,
r ,

t en g en ei g t sei n m 6cht e d en B e sitz ei n er G r un df orm v o r aus zu setze n

, ,

n am l i c h di e M o n o trem e n u n d B e utler si e n i cht be sitze n S ollte da ,

.

n i cht v on d er O nt o ge n ie A uf schl u ss u erw arte n sei n ? z

It is di fficu l t to fin d an adeq u ate reaso n for A eby s assum p ’

tio n of a hochgradige V erkumm erun g of the bilateral hyp ar

‘ ’

t eri a l bro n chial tree i n H ystri x cri s ta ta It may have b ee n du e .

,

as his fi g u re s u ggests to the fact that he exami n ed this l un g by

,

d issectio n an d did n ot have the co n trol afforded by a good cor

r os i on preparatio n The l un g of Hystrix is un iq u e amo n g mam
.
I

m als i n its type of lobatio n The en tire organ is split u p i n to .

n u mero u s small polygo n al lob u les an d is likely to give the impres

s io n of a somewhat abortive str u ct ure whe n ha n dled It pos .

s esses the delicacy an d friability characteristic of man y rode n t

tiss u es an d is very liable to i n j ury d u rin g man ip u lation pre

, ,

ve n ti n g its complete diste n sio n The s u ggestio n of deficien cy .

which A bey received is also fostered by the mi n u te size of man y

o f its lob u lar s u bdivisio n s the organ rese m bli n g i n the plan of,

its organ izatio n e g the sea weed like liver of Cap romys pi le?
,
. .
,
“ -

a n d by the fact that the proximal segme n ts of the stem

'

t des ,

b ro n chi are i n cl u ded at least i n ma n y i n divid u als of H cr i sta ta

, ,

w ithi n a widely expa n ded tracheal b u lla i n place of the typical ,

bif urcatio n .

That the red uctio n of the l un g ass umed by A eby does n ot

act ually obtai n i n the bilateral hyparterial type is ab un dan tly ,

proved by the rich u nfoldi n g of the bro n chial tree shown i n good
corrosio n preparatio n s The foregoi n g review shows that Palae
.

o n tology Palaeozoology O n toge n y an d Comparative Mor

, ,

p h o l o gy are at o n e as far as the available

,
evide n ce goes i n ,

refuti n g the claim to primitive archeal character made i n behalf

1 22 G EO . s . HUNTINGT O N

This view is s upported by the followi n g co n sideratio n s

1 In the fi rst place it does away with the un te n able assum p
.

tio n of the R ed u ctio n Theory that the promamm alia had ac

q uired a higher an d more effi ci en t p ulmon ary organ izatio n than
the on e ex h ibited at the prese n t day by the vast maj ority of their
desce n dan ts i n whom p ulmo n ary red u ctio n on the left side and
, ,

i n a few forms o n both sides is s upposed to have establi shed the

,

hyparterial co n ditio n seco n darily .

2 It is very clear that the mam malian l un g is str u ct u rally o n

.

the asce n t rather than on the declin e It can i n n o sen se be

,
.

gro uped with the decaden t stru ctu res of vertebrate or g ani zatio n .

Morpholo g ically it does n ot represen t the direct co n ti nu atio n of

an old respiratory orga n izatio n b u t a n ew str u ct u re replaci n g the
,

archeal bran chial apparatu s which i n its tu rn was the termi n atio n
,

of the lo n g evol utio n ary path traversed by the vertebrate respira

tory tract i n its man y an tecede n t phyletic stages ( ci 29 p .
,
.

It is co n trary to all gen etic prin ciples un derlyi n g evol u tion to

co n sider a n eomorph of this character capable of e n terin g the
organi c complex i n a state of su ch advan ced developme n t as to
call for s u bsequ en t red u ction of the nu mber and exte n t of its
compo n e n ts The assu mption of mu tatio n is clearly n ot applic
.

able un der the premises for a mu tan t requires a pre existin g

,
—

stru ct u re i n which the chan ge becomes manifest .

3 The f un ctio n al i n terpretatio n of the comparative an a

.

t om i cal evide n ce beari n g on the q u estio n speaks for the p ro g res

sive un foldi n g of the mammalian respiratory area especially i n ,

its cran ial sectors i n direct ratio to i n creased fun ctio n al deman d
,

res ulti n g from sp eci fi c e n viron me n tal adaptation s leadi n g to the

establishme n t of the eparterial district i n the more highly
organized l un gs .

The u n g ulates i n ge n eral are represe n ted by animals of mas

sive b ulk an d heavily m u scled Man y are e n dowed with the
.

capacity for rapid and lo n g co n ti nu ed motio n calli n g for i n

,

cr eased tiss u e comb u stio n an d a rapid respiratory exchan ge .

The members of the gro up have acq uired a marked expan sio n
i n their p ulmo n ary developme n t .
 P ULM ONARY EV O LUTI O N IN A
M MM AL I A 1 23

In view of the recen t palaeo n tological evide n ce of the sepa

rate phyloge n etic derivation of the Perissodactyl an d Artiodactyl
types it is si gni fican t to n ote that the p u lmon ary extensio n pre
, ,

sum ab ly correlated with the same cau sative factors of body

weight mu scu lar b ul k an d active locomotion has been a ecom

, ,

p li sh ed i n the two gro u ps by di ffere n t m o di fi c at i on s of t h e

respiratory pattern The artiodactyl l un g is characterized by

.

the relatively e n ormou s developmen t of the cran ial lobe of the

right side with the supplyin g right eparterial bro n chu s arisi n g
,

from the trachea above the bifu rcatio n In many forms this .

exaggerated developmen t of the right crani al lobe amoun ts

fun ctio n ally to the i n trodu ction of a third l un g exte n di ng from ,

the right side of the thorax acro ss the median lin e an d cappin g
the apex of the left l un g A s a matter of record th e lun g of the
.

cetacean P on top oma blai n mllei has been described by Max Weber
’ '

( 3 6) as possessi n g a bron chial tree with triple division of the

trachea i n to three main bron chi of un equ al caliber .

Amo n g the artiodactyl un gulates the Camelidae ( Camelus ,

A u cheni a) an d G iraffes are characterized by the un foldi n g of th e

typical artiodactyl right eparterial bron chu s arisin g from th e ,

trachea above the b i furcation and i n addition by the develop ,

me n t of a left eparterial compo n en t from the st em b ron chu s of

the left l un g .

In the perissodactyls ( with the exceptio n of the Tapir ) on the ,

other han d the deman d for exten sio n of the respiratory area is
,

met by the bilateral developmen t of the eparterial bron chu s

derived from the st emb ron chus of each lun g .

A mo n g the aq u atic mammalian forms the developme n t of the

lun g i n most cetacean s follows the artiodactyl type while the ,

pi nnipede carn ivores have developed the perissodactyl plan of

p ulmon ary exten sion It is n oteworthy that the sole aqu atic
.

artiodactyl H i pp op otamus agrees with the aqu atic Carn i

, ,

y ores ,
i n the bilaterally symmetrical developme n t of the ep ar
t eri al compo n e n t .

The aqu atic carni vore adaptatio n s gen erally possess the power
of rapi d an d active motio n thro u gh the water exec u ted mai nly
‘

by the highly developed axial an d cau dal trun k mu sc ulat ure .

1 24 G EO . s . HUNTINGT O N

They f urther s u spe n d respiratio n for lo n ger or shorter periods

d u ri n g s ubmersio n In almost all forms spec i al provisio n is
.

made for the collectio n of the ve n o u s blood i n large reservoirs

d u ri n g the respiratory i n termissio n s s u ch as the h uge hepatic ,

caval si n u s of the seals an d the e n ormo u s retiform plex u s of the

,

abdomi n al visceral an d ge n ito u ri n ary vei n s of M acrorhi nu s —

.

Whe n the an imal emerges for breath the large amo un t of ac cum u
lated ve n ou s blood calli n g for oxyge n atio n req uires a rapid and
complete excha n ge In co n formity with this deman d the p ul
.

m o n ary architect u re of these forms prese n ts the most brillia n t

examples of the f ull bilateral an d symmetrical developme n t of
the eparterial system of both l un gs In attempti n g to eval u ate .

the si g nifi can ce of these facts they lead to the followi n g c on

c l u si o n s :

A . E specially
exacti n g f u n ctio n al dema n ds are accompan ied
by modificatio n s of the mammalian p ulmo n ary organ izatio n
thro ugh more exte n sive developme n t of the eparterial bro n chial
districts This exte n sio n of the u ltimate respiratory area may
.

occ u r o nly o n the more favorably disposed right side ( c p .

by u tilizin g a more cran ial poi n t of derivatio n of the right ep ar

t eri al bro n ch u s directly from the trachea cran ial to the bif u r
,

catio n of the mai n t u be ( artiodactyls sire n ia ma n y cetacean s ) , , ,

or on both sides by the additio n al developme n t of the left ep ar

t eri al tr un k ( perissodactyls pi nn ipede car n ivores aq u atic ro
, ,

de n ts arboreal primates Hippopotam us )

, ,
.

B E n viro n me n tal factors have so cha n ged the co n stit u tio n of

.

the germplasm as to tra n smit the res ulti n g alteratio n i n p ul

m c n ary type It is probable that the i n fl u en ce of e n viro n me n t
.

upo n the developi n g germ cells a n d thro u gh them o n the prog ,

ress of evol utio n has bee n u n derestimated i n the past ( Weis

,

man n ) V ariatio n of l un g str u ct ure especially the occ urre n ce

.
,

of m u tatio n s may have bee n the sole or at least the pri n ,

c i p al phyloge n etic factor which led forms possessi n g these modi

,

fi c at i o n s an d by Virt u e of them capable of greater p ulmo n ary

,

un foldi n g to follow li n es of e n viro n me n tal adaptatio n e n di n g i n

,

the evol utio n of the mari n e u n g ulate an d aq u atic types char,

a c t eri z ed i n their extan t desce n da n ts by the high developme n t

of the eparterial system .

1 26 G EO . s . HUNTINGT O N

from the remaini ng members of their respective gro ups is fo un d

i n their i n trap ulmo n ary organ izatio n This raises the important
.

qu estio n what is the evol utio n ary sig n ifica n ce of cardi n al di verg
,

e n ce i n the bro n chial architect u re an d what its val ue as a ge n etic

character i n determin i n g phyletic kin ship ?
To take the example of Tax i dea as a co n crete case i n poi n t .

The zoological gro up of the musteli dae i n which Tax i dea is ,

i n cl u ded is derived by Prof G regory from the lower O ligoce ne

,
.

Considerin g the facts obtaini n g i n the modern des een

'

P lesi ctzs .

dan ts of this commo n an cestral form the R edu ction T h eory would
hold that the P lesi cti s l un g has already developed the domi n an t
—

mammalian type with the reten tio n of the dextral eparterial bron
chu s an d the same had been tran smitted un chan ged to all of the
,

modern descen dan ts with the si n gle exception as far as k n own

, , ,

of Tax i dea In this form fu rther pulmo n ary red u ctio n oc c urred
.
,

res ulti n g i n the phylogen etic loss of the right eparterial lun g
se g men t an d the acqu isition of the tracheal b ulla It is di tfi .

c ult to co n ceive of e n viro n me n tal chan ges capable of i n du ci n g ,

this red u ction an d confined i n their operatio n to Tax i dea to

, ,

the excl u sio n of the other mu stelidae The s upposition ass u mes .

the existen ce of morphogen etic factors which we fin d n owhere

i n the mammalian series I do n ot k n ow of a sin gle i n stan ce of
.

eve n a probable red u ction of a formerly developed p ulmon ary

segmen t i n a mammalian l un g The phylogen etically acquired .

metameric redu ction of the trun k cavity i n mammalia with its ,

in cide n tal e ffect on the ple ural S pace wo uld i n itself n egative this
,

ass umptio n The in cl u sion of the right cardiac lobe i n the

.

p h re n i co m e d i ast i n al a n gle of the right lower lobe i n m a n an d

some other mammalia is merely the res ult of the clos u re of the
,

pericardiophre n ic space followi n g pericardiac fix at i on to the

diaphragm The morphological stat u s of the lobe is mai n tai n ed
.

by the cardiac bran ch of the right st emb ron chus .

If d H ardV i ller s alleged discovery of an ephemeral left ep ar

’ ’

t eri al bro n chial vesicle i n the embryo of the rabbit had bee n
c o n fi rm ed it wo uld poi n t far more directly to an attempt on
,

part of the left l un g to acqu ire additio n al respiratory territory

than to the evan esce n t appearan ce durin g o n togen y of a pul
 P ULM ONARY E V O LUTI O N IN MAM MAL I A 1 27

m on ary elemen t lost i n the phyloge n etic evol ution of the mam
malian l un g This is c onfirm ed by the occ urre n ce of the adu lt
.

bro n chial variation s of L ep u s cited above (p 1 04) i n this c on .

n e c t i on .I feel that we are j usti fied on the basis of all of the ,

act ual evide n ce i n co n cl u di n g that any mammalian l un g which

,

has on ce acqu ired a focal poi n t of e n todermal bro n chial prolif

e ra t i on ,
retai n s the same i n its stru ctu ral organi zatio n eve n ,

again st appare n tly un favorable local an d topographical co n di

tio n s as e g i n limitatio n s of the available thoracic space
, ,
. .
,
.

O n the other ha n d accepti n g P lesi cti s as the comm o n a n cestor

,

of the modern mu stelidae it is possible to assume that the ,

P lesi cti s l un g was orga n ized o n the bilateral s ymmetrical hyp ar

—

t eri al type retain ed i n o n ly on e of the extan t desce n dan ts of

,

to day Tax i dea while i n all the other recen t mu stelidae as far
-
, , ,

as determi n ed e n viro n me ntal chan ges have so affected the con

,

st i t ut i on of the germ plasm as to prod u ce the more advan ced

—

p ulmon ary type with the developmen t of the right eparterial

bro n chu s Al l of our available eviden ce proves the mamm alian
.

lun g to be extremely se n sitive i n its stru ct ural respo n se to en

v i ronm en t al cha n ges a ffecti n g respiratio n An y e n viro n me n t .

capable of produ ci n g chromomeric alteration i n the developin g

g e r m ce l l s of P lesi c ti s m u st have a ffected equ ally a n d u n iformly
all the desce n da n ts derived from the same a n cestor .

The l un g of Tax i dea is physiologically as effi ci ent i n s upplyi n g

the respiratory requ ireme nts of the organi sm it serves as are
the lun gs of the other mu stelidae O therwise Tax i dea wou ld .

e ither have become exti n ct or its l un g wo u ld have adapted itself

,

s tr u ct u rally to the co n ditio n s which i n d u ced the p u lmo n ary dif

f erent i at i o n i n all the other mu stelidae The hypothesis here .

un der disc u ssio n wo u ld therefore become un te n able u n der its

o wn terms were it n ot permissible to a n alyze the evol u tion ary

,

factor which we described as e n viron me n tal adaptati on more

, ,

c losely i n to its compo n e n t parts .

The sp eci fi c gen eric an d ordin al di fferen tiation s of the extan t

,

mammalia appear clearly as the ou t come of adaptation s to a

physical e n viro n me n t di fferi n g widely i n the variou s forms of
m amm ali an o rg an i z at i on i n respect to habitat character of food , ,
1 28 G EO . s . HUNTINGT O N

locomotio n etc The great number of the diverge n t type owe

,
.

their prod u ctio n to differe n ces i n the form of the alimen tatio n
adopted by each In the fin al an alysis the m odifi cat i on s of the
.

skeletal an d m u sc ular systems arose i n respo n se to the primary

n ecessity of obtai n i n g food an d the ma n ifold morphological
,

di ffere n tiatio n s of the i n testi n al tract are the stru ct u ral expres
sio n s of the variety e n coun tered i n the types of alime n tatio n .

The on e relation of the organ ism to the exterior which remai n s

fun dame n tally un altered un der all con ditio n s of mammalian
evol utio n is the respiratory fun ctio n The air breathed by the .

mammal save for slight variation depe n de n t u pon altit u de

, ,

temperat ure aqu eou s vapor an d other extra neo u s admixtu res
, , ,

remai n s physically an d che m ically the same for all forms n o ,

matter how widely their relatio n s to the milie u may di ffer i n

other respects .

The primitive i n testi n al can al of the mammal di ffere n tiat es

hen ce phyloge n etically alo n g two disti n ct li n es leadi n g to res ult s ,

which di ffer correspo n di n gly i n their morphological detail an d

physiological si g nifi can c e .

The strictly alimen tary portion of the t u be refl ect s l n i t s

str u ct u re the e n ormou s diversity of the physiological work
assign e d to it i n the vario u s types i n strict co n formity with the
c orrelated diversity i n the character of their f ood This prod u ces .

the vario u s types of glan d u lar derivatives the stru ct u ral m odi fi ,

catio n s i n le n gth an d caliber the divisio n i n to differe n tiated seg

,

men ts an d compartme n ts the provision of the valvu lar apparat u s

,

an d sphi n cters the variatio n i n the s u ppleme n tary str u ct u res of

,

the oral cavity to n g u e lips salivary glan ds an d teeth the pre

, , , ,

h en sil e m o di fi cat i o n s of the extremities a n d other parts of the

body etc The mammalian organ izatio n compasses the e n tire
,
.

ran ge of the digestible material i n cl u ded withi n the e nviro n

me n tal limits to which its members have become adapted b ut ,

all m ammals breathe the same air Co n seq u e n tly the respira .

tory e n todermal derivatives i n co n trast to the alime n tary modi

,

fi c at i on s prese n t a simple an d u n iform str u ct u re

,
The mam .

malian l un g co nforms to b ut a si n gle basic morphological type ,

n o matter to what a n imal it belo n gs The variatio n s en c ou n

.
1 30 G EO . s . HUNTINGT O N

of the exta n t mammalia n forms the l un gs are asymmetrical

,

o rga n s b u ilt o n A eby s type II with the eparterial developme n t

’
, ,

c o n fi n ed to the right l un g This e n ormo u s prepo n deran ce of

.

the domi n an t mammalian type can o nly sig nify that this degree
of p ulmo n ary exte n sio n meets the respiratory dema n ds of all the
varied sp eci fic forms which operate un der i t despite their ma n i ,

fold diversity i n other stru ct ural directio n s an d is amply ,

suffi c i en t to carry them alo n g their specialized developme n tal

paths .

The exceptio n s above me n tio n ed stan d o ut clearly from amo n g

the great mass of the average forms They are all associated .

with an i n creased respirator y deman d to which the res ulti n g pul

'

m on ary exte n sio n appears as the str u ct u ral respo n se .

In co n trast with these orga n izatio n s typifyi n g advan ce over

average mammalian co n ditio n s the small gro ups co n stit uti n g
,

A eby s fift h type of bilaterally symmetrical hyparterial di st ri b u

’

tio n can o n ly be i n terpreted as the rem n an t of an archeal mam

,

malian organ izatio n which has bee n i n co urse of evol u tio n re

placed by higher types of p ulmo n ary developme n t i n the vast
maj ority of livi n g forms b ut still persists i n a few isolated i n
,

sta n ces a n d gro u ps as i n Tax i dea amo n g the c a

,
rn ivores and i n ,

some of the Hystricomorphs amo n g the rode n ts .

I believe that the co n tradictory discrepan cy i n trod u ced i n to

the problem of ge n etic ki n ship by the atypical p ulmo n ary or
g ani z at i o n of Tax i dea as compared with all other m u stelidae of ,

the Hystricomorphs i n co n trast to the remai n i n g members of the

e n tire rode n t order an d perhaps i n the cetacean exceptio n s m en
,

t i o n ed fi n d s its u ltimate sol u tio n i n the fact that the e ffects of

,

e nviro n me n tal i nfl u en ce upo n the vertebrate orga n izatio n as a

whole mu st be j udged separately b ythe res ults show n i n the i n
,

divid ual compo n e n ts of the same In other words E nviro n .

‘

me n t is a composite co n cept comprisi ng a large number of physi

’
,

cal factors some of which are freq ue n tly a n tithetical to others

,

i n their i n fl uen ce o n orga nic evol u tio n The vertebrate body is

.

likewise a composite of a n umber of sp eci fi c str u ctu res which ,

while they act as a un it i n perform i n g the o ffi ces of the organ iza

tio n as a whole respo n d to their e nviro n me n t as i n divid u al parts
,
 P ULMONARY EV O LUTI O N IN MAMMAL I A 131

of this whole Co n fro n ted by a variety of mu t u ally i ncom p at

.

ible opport unities each poi n ti n g a diverge n t evol u tio n ary path
, ,

the choice of on e of these with the n ecessary excl usio n of the

,

remai n der will determin e the res ulti n g str u ct ural type Thu s
,
.

the compo n e n ts of the organism which come i n to direct relatio n

with a di v ersified e n viro nme n t will adapt themselves o nly to
certai n factors of the same with the res ulti n g produ ctio n of
,

varied and divergen t forms while the l un g which e n counters

, ,

the uniform and practically un varyi n g i nfl uen ce of its specifi c

factor the air will correspo n di n gly appear as a un iform stru ct ure
, ,

i n all the di ffere n t type s .

The followi n g co n cl usio n s c an be based on these co nsideratio n s

i n the special problem here disc u ssed :
While the mu stelidae as a gro u p i n cl u di n g Tax i dea adapt
, ,

themselves to their special e n viro nmen t i n regard to all parts

of the organism i n direct relatio n to the same the res ulti n g ,

adaptatio n s and diff ere n tiatio n s did n ot alter the stat us of the
l un g because the e nviro n me n tal factors did n o t i n cl u de the special
,

con ditio n s determin i n g sp eci fi c chan ges i n the p ulmo n ary archi
tecture They th u s become welded i n to the homoge n eo u s taxo
.

n omic gro u p of the modern m u stelidae i n all the exter n al char

ac t ers of body str u ct ure alime n tatio n habitat locomotio n etc

-
, , , ,
.
,

b u t the i n tri n sic p ulmo n ary orga n izatio n was left u n to u ched by
any e n viro n me n tal i n fl u en c e directed sp ec ifi cally toward str u e

tu ral chan ges i n the architect ure of the l un g beyo n d that com,

m o n to the mammalian type i n ge n eral It is co n ceivable that

.

the direct forebear of Tax i dea had retai ned a more primitive
type of bro n chial orga nizatio n tha n the remai nin g co n temporary
forms which had already adva n ced to the acq uisiti on of the
right eparterial bro n chus .

In the co urse of further evol u tio n n o additio n al factor was i n

t rodu c ed to alter the i n itial discrepa n cy betwee n the two p ul
m on ary types which has th u s become perpet u ated i n their
,

prese n t represen tatives .

This appears to be the most available i n terpretatio n acco unt

i n g for the existe n ce of the closely k n it zoological gro up of the
modern M u stelidae con sisti n g of forms which agree i n the de
,
1 32 GEO . s . HUNTINGTO N

tails of their ge n eral orga nizatio n b u t i n cl u de on e member

, ,

Tax i dea differi n g from the remai n der i n the str u ct u re of its
,

bro n chial tree .

These co n sideratio n s led me to regard the bilateral hyparterial

bron chial tree as the archeal type formi n g the basis of the ,

E xte n sio n T h eory u po n which the remai ni n g mammalian bro n

,

chial orga nizatio n s are b u ilt rather than to begi n at the top
, ,

an d by s u ccessive detachme n t of bra n ches arrive at the bottom , ,

as post ulated by the R ed u ctio n Theory .

I am more than ever co nvin ced by the additio n al phylogen etic

evide n ce which the i n terve n i n g years have bro u ght to the st u dy
of the problem that the basic co n cept o u tlin ed above con tai n s
the seed of the correct i n terpretatio n of the evol utio n ary process
respo n sible for the extan t types of the mamm alian l un g str u ctu re ,

b u t I have aban do n ed my origi n al v 1 ew s regardi n g the ge n etic

stages in volved In the earlier period of this study the idea of
.

the migratio n of p u lmo n ary compo n e n ts was beginn in g to make

headway and appealed to me as affordin g the most available
explan atio n of the steps i n p ulmon ary evol utio n u po n which to
base the E xte n sio n Theory S ubsequ e n t to the p ublicatio n of
.

the same con tinu ed further comparative an at omical an d on to

,

gen etic st udy of the problem led to a radical revision of my

con ception as to the morphogen esis i n volved an d I aban do n ed ,

the Migratory Theory for the View which impressed me as the

only logical ded u ctio n warran ted by all the facts an d which , ,

for wan t of a better term can be b ri efly design ated as the ev o

,

l ut i o n of bro n chial types by selectio n or adaptatio n This .

Selective Theory is f ully expo un ded i n the fin al sectio n of this

paper Before proceedi n g to its co n sideration it is proper at
.
,

this level of the historical review to prese n t the seco n d tho ught
,

li n ked to the theory of p ulmo n ary evol utio n vi z : ,

.
1 34 G EO . s . HUNTINGT O N

prese n tatio n the h om ol o g i z at i on of the right eparterial bro n ch us

with the asce n di ng bran ch from the fi rst left ven tral hyparterial
bro n ch u s to the cra nial pole of the left upper lobe .

Fi n ally N arath ( 3 1 3 3 ) i n a prelimi n ary paper p u blished i n

, ,

1 8 9 2 an d i n his large mo n ograph

,
became the chief an d
very e n thu siastic expon e n t of the theory offeri n g i n s upport of ,

the same man y addition al comparative a n atomical an d o n to

ge n etic obse rvatio n s and an i nval u able record and cl assi fi ca
,

tio n of bro n chial varian ts If i n the followi n g pages I appear

.

as an an tago nist to N arath it is solely because many years of

,

caref ul st u dy give n to the problem have co n vi n ced me that th e

ded u ction s which he has drawn from his material and the theo ,

ret i c al co n cl u sio n s based thereo n are erro n eou s These I feel

, .

obliged to oppose But I do thi s with a full realizatio n an d

.

appreciation of the perman e n t val u e of N arath s con trib ution ’

.

His mon o g raph is a veritable storehou se of n ew morpholo g ical

observatio n s an d records sple n didly arran ged clearly expressed
, ,

an d beau tif ul ly ill u strated It will remain for all time as one
.

of the classical records markin g the progress of an atomical

scie n ce .

A s elaborat ed by N arath the mi g rati n g theory of bro n chial

,

evol utio n if followed to its logical con cl usio n ass umes a simpl e
, ,

hypothetical phyletic groun d plan of bro n chial organi zatio n

-
.

The st emb ron chus gives origin solely to a sin gle ve n tro laterally -

directed series of primary bron chial derivatives L ateral .

bran ches of these alter their primitive positio n i n the bro n chial
system i n the co urse of evol ution by becomin g split off from ‘ ’

their origi n al pare n t stems an d tran sfer themselves thro ugh a

,

process of wa n deri n g or migration to the st emb ron chus ob

‘ ’ ‘ ’
,

tai ni ng th us n ew and i n depe n de n t poi nts of origin directly from

the same In the words of the theory as stated by N arath
.
, ,

the origin al ve n tro lateral bran ch of the st em b ro n chus has

—

‘
ceded ’
o n e of its lateral derivatives to the same .

The evol utio nary record of the domi n an t mammalian type

(A eby s type II ) wo uld read as follows :
’ A

The bro n ch u s desti n ed eve n t ually to s upply the upper lobe

of a f ully developed right l un g makes its first appearan ce as a
 P ULMONARY EV O LUTI O N IN MAMMAL I A 1 35

side bran ch of the first ve n tral derivative of the right stem

-

bro n ch u s ( A eby s first right ve n tral hyparterial bro n ch u s ) It

’
.

then detaches itself from the pare n t stem an d shifts or mi ‘

grates dorsad o n to the right mai n st em b ron chu s an d j oi n s a

’

gro u p of similar bran ches derived i n the same w ay from the pri
mary ve n tral bron chi arisi n g f urther down i n the row from the
st em b ron chu s It co n stit u tes th u s the most cra nially located
.

member (D ) i n the series of A eby s dorsal bron chi A s s u ch D

1 ’
.
l

comes to exte n d i n to the dorso cranial l un g segme n ts s upplyi n g-

,

its cranial pole as N arath s apical bro n ch u s It possesses the

,
’ ‘ ’
.

fac ulty of co n ti nui n g to wan der further crani ad alo n g the stem
bro n ch u s Whe n this mi gratio n has carried it above the level
.

of the i n tersectio n of st em b ron chu s and p ulmon ary artery it

becomes still as apical bron ch u s the equivale n t of A eby s
,
‘ ’
,
’

eparterial bro n chus It is able to exten d its forward march

.
,

leavi n g the domain of the right st emb ron chus and gai ni n g an
origi n from the right side of the tracheal bif urcation or from the
lateral wall of the trachea above this level as i n the artiodactyl ,
B
l un g ( A eby s type II ) In the left l un g the same compon e n t
’
.

( apical bro n ch u s) retai n s the primitive origi n from the fi rst

‘ ’

ve n tral hyp arterial bron ch u s appearin g as its asce n din g ,

bran ch does n ot emigrate an d supplies the dorso cranial seg

,
—

men t of the left upper lobe .

If it follows the co urse take n by its homologu e of the right

side and wan ders craniad o n to the left st emb ron chus A eby s ,
’

bilateral eparterial type I develops If no shift occ urs on either

A
.

side the bilateral hyp arterial tree is prod u ced ( A eby s type III ) ’

O ther lateral derivatives of the primitive ve n tral bran ches ,

u s u ally small an d of mi n or importa n ce wa n der from their pare n t ,

stem to the st emb ron chus an d the n appear as seco n dary bran che s
of the same ( accessory bro n chi A eby s ,
’

arisi n g from the mai n can al i n the in tervals betwee n the rows of
pri n cipal ven tral an d dorsal bro n chi .

O rdi n arily o n ly on e of these derived origi n ally from the fi rst

,
.

or sec o n d ve n tral bron ch u s migrates to the st emb ron chu s an d

,

attai n s a greater degree of developmen t s upplyi n g as right car , ,

diac (i nfra cardi ac) bro n ch u s the azygos or cardiac lobe of the,
1 36 G EO . s . HUNTINGT O N

right l un g A similar accessory bro n ch us may develop on the

.

left side as the left cardiac bro n ch u s to a left azygos lobe or to ,

the correspon di n g portio n of the left l un g stem .

The thou ght un derlyi n g this co n cept of mammalian bro n chial

evol ution is best expressed i n N arat h s own words : Es herrscht “
”
’

j a ii b e ral l i m Bro n chialba u m das G esetz der A bspalt un g .

This co n de n sed accoun t embodies the chief dogmata of the

Migration Theory It is psychologically of i nterest to consider
.

b ri efl y the reaso n s for the ready acceptan ce of this idea an d its

almost uni versal adoptio n i n sci ent i fi c an d ed ucatio n al circles
to day
-
.

In the fi rst place the e n ormo u s nu merical prepo n deran ce of

th e domi n an t type II amo n g livi n g mammalia imparts to the fo u r
A

remai ni n g architecton ic types the character of aberratio n s By .

r eason of their bilateral sym metry they give the impressio n of

bein g either earlier more primitive forms or they s u ggest a rela
, ,
A
tion to type I as represen tin g dedu ction s from the sam e or
,

additio n s to type III N o observer who has stu died large

.

series of p ulmon ary corrosio n preparation s c an fail to recogni z e

t h e close architect u ral gradatio n s of the bro n chial tree i n di ffer
en t forms Witho ut co n sideri n g the variation s within a sin gle
.

species which are often nu mero u s an d highly s u ggestive whe n

, ,

many i n dividu als are compared it is easy to form lo n g series ,

whose links pass i n to each other by almost in sen sible degrees ,

a n d th u s j oi n the extreme examples of a type at either en d of

the li n e It is n at ural to regard these tran sitio n s as poin ti n g

.

the w ay to the phyloge n etic derivatio n of a give n type from a

p recedin g stage by a shift or migratio n of some of its bro n
c hial compo n e n ts N arat h f u r n ishes an excelle n t ill u stratio n of
.

t his me n tal process In s upport of his co n te n tio n that the dorsal

.

d erivatives of the st em b ro n ch u s were origi n ally side bra n ches of

the ven tral bron chi which have migrated o nto the st emb ron chus
itself he says (33 p 3 1 1 )
, ,
.

M ankann si ch bei m an chen schon g elung enen Inj ecti onspraparat en

k aum des G e danken s erwehren d ass d er d or sale B ron chu s ei n A rt
g en osse der extern en S ei t en ast e d er Ven t ralb ro n chi en i st u n d d ass er
g ewi sserm assen al s V o poste n vo rg eschoben wurde auf d en S t ami n
r

bron chus .
1 38 G EO . s . HUNTINGT O N

cept of the early phyloge n etic type of bro n chial un foldi ng If .

the right eparterial an d cardiac bro n chi of the prevale n t type

were primarily derivatives of the fi rst ve n tral bro n ch u s the for ,

mer a dorsal the latter a ve n tro medial bra n ch an d if i n the

,
—
,

rest of the series all dorsal dorso medial an d ve n tro medial ,

-
,
—

compon e n ts of the st em b ron chus were origi n ally derivatives from

the primary ve n tro lateral bro n ch u s of their respective levels
—
,

then these premises wo uld define the most primitive mammalian

or promammalian stem bro n ch us as givi n g origi n solely to m on
-

op o di c ve n tro lateral derivatives

-
whos e seco n dary bran chi n g ,

Fi g . 4 A
H yp i
o t het c a l pl an o f t he p im it i v d iv t i v
r e er a es o f t h e st em - b ro n c h u s ,

as st ip u la t e d by t he M i g ra t io n Th eo r y .

carries their distrib utio n in to the remain in g dorsal dorso medial ,

-

an d ve n tro medial districts

—
The schematic cross sectio n of a
.

hypothetical reco n stru ctio n of this bro n chial type wo uld appear
as i n figure 4 A
Migratio n the n becomes respon sible for the re
.

B
distrib u tio n of the compo ne n ts shown i n figure 4 .

The p ulmo n ary organ izatio n represen ted i n figure 4 is not A

foun d i n the phyletic series .

If this is accepted as i n dicatin g the earliest phyloge n etic mam

malian type of bro n chial organ izatio n on e of two co n cl usion s ,

wo uld be i n evitable
a E ither the mammalia n l un g represe n ts the persiste n ce of a
.

more primitive p ulmo n ary type than that fo un d i n the embryo

or ad ult of any of the extan t lower vertebrates or ,
 P ULMONARY EV O LUTI O N IN MAM MAL I A 1 39

bThe mammalian l un g is a ce n ogen etic stru ctu re an d does

.

n ot represe n t the en d li n k i n a co n ti nu o u s evol u tio n ary process

-
.

There is n o phyletic relatio n betwee n it an d the l ungs of the lower

vertebrates .

N either of these hypotheses is te n able O n ge n eral ph ylo .

gen etic gro un ds the migratory theory n ot o nly receives n o sup

port b ut is positively co n tradicted
,
.

Fig . 4 B
Th ie r su b quse en t re di ib ut i
st r on t hrou h m g ig i
rat o n . V .
, ven t r a l ;
V—L .
,
v en t r o -l a t er a l ; D .
,
d o r sa l ; Dm d .
,
o r s o -m e di al ; M me di al ; Vm .
, v en t r o

me di al .

Hesser i n his importan t and thoro u ghly scien tifi c paper on

,

the developmen t of the reptilian lun g ( 2 2 ) voices this fun da

me n tal obj ectio n to the migratory theory admirably In dis .

c u ssin g the mu ch mooted qu estion of mon opody versu s dichotomy

-

i n the developme n t of the bro n chial system he says (p .

S tellt m an das ,
erschie denen S tellen g eaussert hat
was N arat h an v

u ber d en phyl o un d on t o g e n eti sche n U rspru n g d er v om S t amm b r on

chus au sg ehe n de n Aste z u samm en so k omm t m an u ei n em R e su l t at
, ,
z ,

wel che s schon an un d fur si ch unwahrscheinli ch erscheinen muss un d ,

wel ches irn Li chte des B au es der nie deren L ung enf orm en als fal sch
bezei chn et werden mu ss Er sag t n amli ch d ass ni cht nur di e dor .
,

sale n S ei t en b ron chi en i n ihrer e sten En t stehu n g m i t den ven tr ale n r

z u s amm en g el en k t si n d son d ern d ass au ch di e sog e n ve n tr alen N eben

,
.

br on chien ursprun gli ch auf diesen an g eleg t werden wi e au ch di e sogen ,

.
1 40 GE O . s . HUNTINGT O N
d or salen N eb en b ro n chi en primar Aste d er dorsalen S ei t enb r o n c hi en
sin d . S ek un dar wahre n d d es Ent w i ckel un g sg ang es komm e n d ie se drei
verschie denen A rten Aste ihren Platz auf dem S t amm bron chus z u
erh alten A ber , sag t er weiter , wenn ei n Ast auf diese Wei se an einen
.

an dern soll ab g e g eben wer d en k enn en , so mu ss er stere r sehr f rii h

an g ele g t wer de n , z u ei n er Zeit , w o d er M u tt erast si ch nu r i n sein er

ersten A nl ag e befindet und si ch an der Wurzel n och ni cht z usammeng e
z og en h at .

A ls o wii rde n a ch N arat h ein e v en trale K n ospe , wenn si e n o ch kein e

hoh ere En twi ck elun g errei cht h at , al s d ass si e di e F o rm ei n es n ie dri g en ,
ab g eru n d ete n K eg el s m i t breiter B asi s h at , A n l ag en z u Vier v ersc h i e

den en Asten ( ein em ve ntr alen un d ein em d orsale n S ei t enb ron ch u s wie
einem ventralen und ein em dorsalen N eb enb ron chu s) i n si ch schli es
sen , v o n d en e n di e d rei letzteren sekun dar den erstere n v erl assen u n d ,
au ch u n terei n an der i soliert , ihre n be stimm te n P l atz auf d em S t amm
bron chus einnehmen .

A u sser , d as so g ar N arat h selb st , d er d o ch die sem K apitel sei n e b e

so n dere A ufm erks am keit g ewi dm et h at , i n d en v e n tr ale n K n ospen
n iem al s die se Vier A n l ag e n differen ziert un d d e n n och i m Z u samm e n
han g unterein ander hat fi n den k 6nnen , liegt wohl a pri ori fur j e der
m ann etwas U nn at urli ches u n d U nw ahrschein li che s i n die ser Auffas
V erf ol g t m an N arat h s G e d anken g an g n o ch etw as weiter , so
’
sun g .

kommt m an z u dem v on N arat h selb st n i cht ausg esprochen en und von

ihm Viellei cht ii b erseh en en S chluss , d ass 1 n d er Phyl o g en e se d er L u ng e
ei n S t adium exi stieren s ollte , w o v om S t amm bro n ch us n u r ei n e ein z i g e
R eihe S ei t en ast e au sg eht , wel che wahren d d er f ort schreiten den phyl o
g en eti schen En twi ckel un g ei n en T eil i hrer N eb en ast e an d en S t amm
bron chus abg eg eben h aben , wo durch dieser bei den ho chsten Lungen
formen i n di e L ag e komm t , i n m ehr als einer R eihe Aste auszusenden .

No ch s ollte n d o ch au ch bei d en S éi ug et i eren diese l et z eren Zweig e auf

d en anf ang s allei n vorh an den e n ventr ale n S ei t enb ron chi en an g ele g t
werden u nd dadur ch ihre ursprii ngli che H erkuni t d okumentieren '

In the amphibia n an d reptilian lung the dorsal eva g i n atio n s

appear n ot o n ly as separate derivatives from the ce n tral pul
m o n ary space b u t i n some reptilia n ( lacertilian ) types they
, ,

form o n toge n etically as the earliest i n dicatio n s of the i n creasi n g

complexity of the orga n whereas the larger ve ntral compart
,

me n ts develop as b u ds from the pu lmo n ary circ umfere n ce

separately an d at a later stage .

Hesser describes (l c p 2 3 5) and fi g ures ( Taf 1 9 Fig 1 1 )

. .
,
. .
,
.

the p ulmo n ary reco n str u ctio n of an mm Tare n tola embryo .

—
,

i n which this co n ditio n is show n admirably i n the dorsal view of

the left l un g In t u rtles he fi nds (l c p 3 00) the maj ority of
. . .
,
.
1 42 G EO . s . HUNTINGT O N
X
7 may arise from the a ngle formed by the st em b ro n chus
. D
with the origi n of V Z
.

X
8 D a n d V may arise by a short commo n segme n t from the
Z
.

st em b ro n ch u s .

K
9 . D may default e n tirely .

5 6

Fig . 5 S c h em a o f v i ti
ar a on in or i g in of t he d o rs a l b r on c h i in i
re l a t o n t o t he
st e m an d t he
‘

v en t r o - l a t e r a l p im
r ar y b ran c h es . Vy .
,
p xim l v t
ro a en ro - l a t e ra l

b ro n c hu s ; Vz .
,
di st a l v e n t ro -l a t e ra l b r on c h u s ; D x . c o r re s p di g d l b
on n o rs a ro n c h us .

This comprises the s ubstan ce of N arath s comparative an a ’

t om i cal evide n ce for regardi n g the dorsal bro n chi as derived

phyloge n etically from the correspo n di ng ve n tral bro n chi H e .

lays stress on the fact that they show a ge n eral resemblan ce to

certai n of the side bran ches of the ve n tral bro n chi that they ofte n ,

arra n ge themselves i n series with them correspo n di n g i n size , ,

mann er of divisio n and directio n and that they appear so to

“
speak to be i n a way related to them (3 3 p ” ,

,
.
 P ULMONARY E V O LUTI O N IN MAMMAL I A 1 43

Cases i n which the dorsal bro n ch u s of a district defaults ( fig .

5,
or arises very n ear the ve n tral associate ( fi g 5 or where .
,

the dorsal an d ve n tral bro n chi of a give n level have a commo n

origi n from the st em b ro n chu s ( fig 5 or are j oi n ed i n the cor .
,

r o si on cast by a sort of ridge appeal to N arat h as yery co n vi n ci n g

,

evide n ce for his theory .

Fran kly I fail to see i n what respect these facts s upport his
co n te n tio n If w e regard the co n ditio n s show n schematically i n
.

fi g ure 5 W itho u t prej u dice or preco n ceived theory they simply ,

demo n strate the fact that the dorsal bro n chi are primarily char
a c t eri z ed by the extreme variability i n their origi n an d i n rela

tio n to adj acen t elemen ts of the bro n chial tree .

In disc u ssi n g N arat h s ass umed migratio n of b u ds from the

’

ve n tral anlage n to the st emb ron chu s to co n stitute the dorsal ,

series Hesser (l c p 3 03 ) makes the followi n g s u ggestio n :

,
. .
,
.

Viellei cht i st di e i n R e d e stehen de Erschein un g n u r al s ei n e v on der

g ew ohnli chen En t stehu n g swei se abwei chen de V ari ati o n aufz uf asse n .

Es er schei n t m i r n am li ch al s ob d i e d orsalen B r on chien bei den S éi u g e

,

tieren verschieden e V ari ation en hin si chtli ch des O rtes ihrer ersten
En tstehun g d arbieten sollte n In d er R eg el wer den si e auf d em S t amm
.

bron chus ang eleg t si e konnen abe au ch auf der G ren ze zwischen di e
,
r

sem u n d d en v e n t alen S eite n kn osp e n o de

r i n Au sn ahm ef all en sog ar r

g an z u n d g ar auf den letzteren er schei n e n A u ch i n die sem letzten .

F all k omm e n si e d o ch seku n dar auf d em S t amm bron chu s u sitze n z ,

Viellei cht weil di e den bei den K n ospen anf an g s g em einsam e M un dung
i n den S t amm br o n chu s hi n ein g ez og en wird wen n die ser wahre n d d er ,

En twi ckel un g an Um f an g z u n im m t .

Hesser co n cl udes this co n sideratio n of N arat h s developmen t ’

of the migratory theory with the statemen t ( 22 p 3 0 5) of his

“
,
.

co nvictio n dass die dorsale n Sei t en b ron chi en ihre n U rsprun g

direkt vom Stammbro n ch u s n ehmen ” .

Fli n t (2 1 p p 1 1 9 1 2 0) makes the followi n g comme n t :

,
.
,

In l o oki n g u pon the d orsal g r ou p as deriv ativ e s of the l ate al br on r

c hi N arat h h as the su pp o rt o f Bli sn i an sk aj a w ho arg u e s if the

‘
,
ep ar ,

t eri al i s a d o rso l ate al bro n chu s it i s e aso n able t o su p pose the re

’
r ,
r

m ai n d er of the se ies are sim il a ly d e ive d

r Neither of the se authors
r r .
,

however h av e foll owe d the wan dering ste p b y step eithe of the
,
— — r

eparteri al or the do sal bran ches o n t o the stem bron chus They are
r .
,

o n the c on tr ary i n de penden t de iv atives of the stem and like the

,
r ,

l ateral series are t o be c on sidered as a g roup of p in ci pal b onchi

,
r r .
1 44 GE O . s . HUNTINGT O N

He f urther states (2 1 , pp 1 1 2 ,
.

All of t he rg um en ts o f N a ath an d Bli sni anskaj a con cerni ng t he

a r

deriv ati o n of the ventr al d o sal an d m e di al serie s either pri m a ily

,
r ,
r

or se co n d arily fr om the l ater al br on chi are q uite u n co n vi n c i n g f o r li k e ,

the su pport which N arat h bri n g s from the com parative an at omy the
, ,

f act s are c apable of a sim pler expl an ati on i e a wi de v a i ati on i n the ,

. .
,
r

positi o n of the b u ds an d t h e power of on e br o n chu s sub stit uti n g f or

an o ther .

The comparative an atomical facts add u ced by N arath f urther

demo n strate that the dorsal p ulmo n ary areas fn ay receive their
co n d u cti n g t ubes from any accessi ble p oi n t of the adj acen t bronchi al
tree wi thi n thei r r an ge This is the importan t co n cl u sio n whose
.
,

si g ni fi c an c e will be disc u ssed prese n tly The Migratory Theory .

s u ffers from a grave co n stit utio n al defect It has become so .

imb u ed with the mechan ical aspects of its problem that the n aked
corrosio n cast of the bro n chial tree ass umes the proportio n s of a
morphological un it complete i n itself an d to be j u dged a s s u ch
,
.

It is a tree stripped of its fin er twigs and leaves the un foldi ng of ,

which is the primary physiological p u rpose of the en tire stru ct ure .

A l un g corrosio n i n which the i n j ectio n has bee n carried i n to the

respiratory bro n chioles an d alveolar spaces correspo n ds to the
tree i n f ull leaf In both the same degree of un foldi n g of an
.

equ al respiratory s urface c an be accomplished by gro u pi n g the

eleme n ts of the co n d u cti n g an d s upporti n g skeleto n i n a variety
of mechan ical pattern s The tree has the larger ran ge of varia
.

b ili t y i n the nu mber an d dispositio n of the i n divid u al leaf b u ds ,

b u t the bro n chial system tho u gh red u ced i n pote n tial exc ursio n
,

by the physical e n vi ro n men t follows absol u tely the same mor ,

p h o g en et i c plan .

It is at times diffic ult especially i n the cau dal p ulmo n ary areas
, ,

to select from amo n g the irreg ularly disposed dorsal bron chi the
proper i n divid u al eleme n t s u pposed to belo n g ge n etically to a
give n ve n tral bro n ch u s N arat h himself recog nizes this whe n he
.

says (33 p ,
. D i e O ri en t i run g wird dann u n gemei n erschwert
un d m a n ch esm al ist ub erh aup t ei n e sichere Besti m m un g des

Bro n ch u s unmogli ch .
1 46 G EO . s . HUNTINGTO N

N a ra t h s ums up the morphological characters

(3 3 , p .

which the eparterial an d dorsal bro n chi have i n commo n as

follows :
1 Both exhibit a great variety i n their poi n t of origi n from
.

the st emb ron chus and a marked capacity for wan deri n g
,
‘ ’

( W a n derun g sfah i g k ei t
‘ ’

2Both arise at a higher leve

. l from the st emb ron chus tha n the
correspo n di n g ve n tral bro n chi
. .

3 Both have the fac ulty of defaulti n g altogether i n certai n

.

cases .

4 Both can arran ge themselves i n a well ordered serial ro w

. .

5 Both agree i n their relatio n to the p u lmo n ary artery

.

6 Both s u pply o nly dorsal p ulmo n ary segme n ts

. .

7 They may be do u ble or m u ltiple

. .

8 A ccessory bro n chi may develop alo n gside of them from the
.

s t em b r on c h u s .

9 . O ccasio n ally
they agree i n their form and type of bra n chi n g .

1 0 They have the same arterial s upply

. .

1 1 In the Mo n otremes their vei n s are correspo n di n gly

.

arra n ged .

O n the basis of these eleve n commo n characters N a rat h de

c lares u n eq u ivocally that A eby s eparterial a n d his apical bro n
’

ch u s is n othi n g else than the fi rst dorsal derivative of the stem

bro n ch u s shifted by migratio n to a n ew site The fallacy of
‘ ’
.

this co n te n tio n is evide n t The characters 1 3 4 7 8 9 listed .

, , , , ,

above agai n o n ly speak for variability i n origi n an d acce n t u ate

that fact that the dorsal l un g segme nts can receive their bro n chial
s upply from any accessible poi n t on the st em b ron chus What .

ever reg ularity of origi n exists for the dorsal bro n chi i n certai n
corrosio n preparatio n s depe n ds upo n the limitatio n s of this
, ,

accessible ran ge N o 2 does n o t hold good i n all cases for the

. .

d orsal bro n chi below the eparterial eve n if the desig n atio n D is 2

s ubstit u ted for A eby s D N o 6 is disproved for the eparterial

’ 1
. .

bro n chial distrib u tio n by i nnu merable i nsta n ces N o 1 0 o nly . .

pertai ns i n the se n se that all portio n s of the l un g receive bra n ches

from the p ulmo n ary artery .
 P ULMONAR Y EVO LUTI O N IN M M A MAL I A 1 47

Of all the characters cited by N arat h the only on es that hold

good are N o 5 the p ulmon ary artery relatio n an d N o 1 1 the
.
, , .
,

relatio n of the mon otreme p u lmo n ary vei n s Y et N arath him .

self is most emphatic i n denyi n g to the vasc ular s upply of the

l un g any morphological si g nifican c e i n the in terpretatio n of the
bro n chi al tree H e qu ite correctly sees i n the arran gemen t of
.

both p ul mo n ary arteries an d vei n s n othin g b ut a close adapta

tion to the i ntrap ulmo n ary architecture the vessels fit t i n g them
,

s elves i n to the i n terbro n chial spaces betwee n the developi n g

b ud s
Comparative an atomical evide n ce i n s upport of the Migration
Theory goes by the board altogether .

EL et u s exami n e the o n togen etic evide n ce with which the last ,

word m ust rest i n an y case .

For this N arath tu rn s to on e so urce where defini t e det erm i n

ation is either extremely di ffic u lt or n ot at all obtai n able V i z ,
.
,

the embryo of a mamm al ian form i n whi ch i n the ad ult the right
eparterial an d fi rst ven tral hyparterial bro n chi arise i n close
p roxi m ity to each other from the right st em b r o n ch u s In ma ny .

forms i n cl uded i n this type the open i n terval betwee n these two
primary bro n chi barely s uffices i n the adul t for the passage of
the mai n trun k of the right pu lmo n ary artery i n its ven tro
dorsal co urse across the lateral s urface of the stem bron ch u s In —
.

these types ( an d N arath s chief examples E chidn a an d L ep u s

’
, ,

belo n g to this group ) the earliest anl agen of the primary bro n chi
of the right l un g appear i n the form of slight swellin gs of the prim
i t i ve l un g t ube gro u ped closely together an d grad u ally shadi n g
-
,

in to the adj ace n t e n toderm of the fu t u re st em b ron ch u s

,
At .

this stage there exists as yet n o clear differe n tiation between the
st emb r o n c h u s an d its primary derivatives It is n ot possible to
.

delimitate acc urately the ce n tral t ube agai n st the fain t swellin gs
de n otin g the fi rst b u ds of the fu t ure bro n chi i n q u estion and of
these anlages agai n st each other The attempt to do so resul ts
.

merely i n an expression of the observers person al j udgme n t an d

aff ords n o proof either for or again st their c onfl uent or discrete
origin becau se the bed from which they arise the f utu re stem
, ,

b ro n ch us has n ot yet declared its defin ite limits

, It is held by .
1 48 G EO . s . HUNTINGT O N

N arath that i n these forms the right eparterial an lage arises

from the cran ial slope of the fi rst ve ntral hyparterial b ud an d is
hen ce to be regarded o n toge n etically as a bra n ch of the same .

E ve n N arat h fi n ds himself i n diffic u lties i n attempti n g to sub

st an t i at e his claim o n material of his o w n selectio n His best .

ill u stratio n s fail to co nvi n ce becau se the co n ditio n they attempt

,

to portray v i z the defini t i on of the bro n chial an lagen agai n st

,
.
,

each other and the fu tu re st em b ro n chus does n o t as yet obtain , .

In the followi n g developme n tal stages the an lage n are said to

‘
move apart A s a matter of fact the st emb ron chu s declares
.
’

itself an d the n the i n divid u al bro n chial b u ds appear as separate

derivatives from the same This movi n g apart o r to speak .
‘
,
’
,

more correctly this clearer defi ni t i on of their relatio n s can n ot

, ,

i n my j u dgme n t be u tilized as evide n ce that the right eparterial

b ud has migrated from the fi rst ve n tral hyp arterial an lage to
‘ ’

a n ew and separate poi n t of origi n on the st em b ron chus It is .

o n the co n trary a demo n stratio n of t w o separate an d i n divid u al

focal poi n ts of e n todermal b u ddi n g which become appreciable

o nly whe n the epithelial model of the f ut u re bron chial system
has become suffi ci ently defi n ed to make the acc u rate locatio n of
the compo n e n ts i n their relation s to each other an d to the
st em b r o n ch u s Vis u ally possible N or do I believe that w e are
.

j ust i fied o n the evide n ce fur nished by the early epithelial an

,

lagen of this type of l un g i n decidi n g that two adj ace n t an d ap

,

p arent ly i n part c o n fl uen t b u d s w h i ch later with the clear d efi

°

, ,

n i t i on of the st em b r o n ch u s appear as t w o separate bra n ches of

,

the same have rehearsed a greatly foreshorte n ed chapter i n

,

their phyloge n y i n which on e of them has migrated or shifted

,
‘ ’ ‘ ’

bodily to a n ew site All that the very earliest stages show i n

.

embryos of this mammalian type is a commo n extremely plastic

e n todermal t ube i n the process of peripheral b u ddi n g This b ud .

di ng follows a ge n etic type which is still see n i n the embryos of

some extan t reptiles ( lacertilia chelo n ia) The i n divid u al cir ,
.

c u m f eren t i all y disposed evagi n atio n s te n d to arra n ge themselves

i n to metameric series .

Hesser (2 2 p , after describi n g the po u ch like evagi n atio n

.
-

of the cran ial pole of the primitive l un g sac i n the mm -

.
1 50 GE O . s . HUNTINGT O N
i g k ei t en
. W e nn di e K n ospen sehr deutlich si chtb ar si n d , d an n h abe n
si e i n d er g r o sse n M ehrz ahl der F alle kei n en Z u samm e nh an g m i t d er
beim K anin chen tiefer sitzen den z ug ehori g en Vent ralb ron chusanlag e .

N u r m an chm al k ann m an u n ter der L u p e o der d em M ikr o sk o pe d ur ch

D rehun g e n d er S chraube ei n e n deutli chen Z us amm enh an g der C ont o ur
li nien erkennen M an m uss auf n o ch j ung ere S tadien z uruck gehen ,
.

w o di e allererste , g an z mi nim al ste Vorwo lb u n g der Epi t h elwan dun g z u

con st at i ren i st u n d d ari n lieg t d i e S chwieri gkeit d er g an zen U nter
su c h un g
. G an z erg eb n i sl os w ar je d och di e A rbeit ni cht Bei auf .

g ehei lte n L un g e n k onn te i ch ei n ig e Male di e allererste A n deutung der

D orsalkn ospen b i ldung erkenn e n D i e A n l ag e der D orsal
.

kn ospe sitzt z u einem guten Theile auf der hinteren und oberen Abda
chun g d er Vent ral kno spen erheb ung .

Hesser (2 2 p 3 0 1 ) com m en ts on this passage i n thes e words

,
.

I n des i st u bem erken dass eine derartig e Erscheinung ausserst

z ,

selte n z u zein schei nt In d en m eiste n F alle n sieht m an kei n e n Z u

.

samm enh an g zwi s che n d er D ors al kn ospe un d der e nt spreche nden v en

t ralen Bro n chusanl ag e ein e Th at sache wovon N arat h durch di e
, ,

Erkl arung abz ukomm en su cht d ass di e un tersu chte n S t adien n i cht
,

fruh g e nu g g ewe sen sin d Ub ri gen s schei nt N arat h selb st sei ner S a ch e
“
.

ni cht s o g ewi ss z u se i n de nn er sag t an ei n er S telle dass er sich Viele

” ,

Muhe g eg eben den U rs prun g der dorsalen B r on chien z u entdecken

,

“ohne einheitliche B efun de z u errei chen .

,

Mammalian pulmon ary developme n t con sists i n progressive

ce n trifu gal epithelial sprou ting from a series of d efini t e focal
poi n ts of the primitive lun g sac An epithelial bud o n ce formed .
, ,

conti nu es to exte n d peripherally It gives rise to lateral daugh .

ter bu ds which carry on the same type of developme n t b ut

-
, ,

n ever leave the pare n t stem n ever divide by splitti n g off an d ‘ ’

,

n ever shift to a n ew locatio n .

N either phyloge n y n or o n to g e ny s u stain the dogma of the

Migratio n Theory They fail to prove that the dorsal bro n chi
.

are primarily bran ches of the correspo n di n g ve n tral bro n chi

which s ubsequ e n tly aban do n their early origi n and become tran s
planted on the st emb ron ch us They fail especially i n provi n g .

that the right eparterial bro n chu s is the cran ial member of the
dorsal series an d that as su ch it is origin ally a bra n ch of the first
ve ntral hyparterial bro n chus N arath himself un co n sciou sly ex
.

presses this when he writes ( 33 p 3 1 3 ) as follows i n conn ectio n , .

,

with the s upposed derivatio n of the cardiac from the first ve ntral
hyp arterial bron chu s
 P ULMONARY EV O LUTI O N IN MAMMAL I A 1 51

D i e U n tersu chung d er lle e sten A nl ag e v orn i nfracardi alen B ron

a r r

chu s i st au sser st schwieri g e s h an d elt si ch j a w i e bei alle n B r on chus

,

anl agen um g an z m inim ale V orb au chung en di e mi t un seren H ilf sm it ,

tel u schwer o der g ar ni cht u e kennen sind W as wi r sehen i st b e

z r .
,

rei t s ei n altere s S t adi um m an m ii sst e n o ch weiter

,
u rii c k g eh en i n der z

Entwi c kl u n g u n d w ii rde d ann auf ei n en be sti mmten Z ellk om pl ex k o m

men au s dem d er B ron chu s herv org eht
,
D er Zell g ru ppe k enn e n w i r .

e s je do ch n i cht an sehe n d ass si e d en B o n ch usk ei m i n si ch trag t

,
r Ich .

d achte ei n e Zeitl an g d ass m an Vielleicht durch ei n e starkere A n h au

,

fung v on Kernt heilung sfi guren einen A nhaltspunkt d afii r g ewinnen

konne um u ent schei den von w o eig entli ch di e Knospenb il du ng
,
z ,

au sgeht D i e Untersu chung en liessen je doch v oll st an di g im S ti ch

. .

The above passage which on accoun t of its si g ni fi can ce I have

,

qu oted i n full really tells the e ntire story an d emphasizes the

,

way i n which a precon ceived idea ten aciou sly retain ed can pro , ,

du ce eve n i n an able observer a men tal attit u de approachi n g

, ,

mysticism For if a morphoge n etic theory dep en ds altogether

.

o n basic di ff ere n tiatio n s ass u med to occ u r i n s u ch early o n to

g e n etic stages that they cann ot be see n or recorded the d edu c ,

tio n s must rest e n tirely on faith an d n ot on sci ent ifi c evide n ce .

This is carryi n g theoretical promorphology beyon d its j usti fiabl e

limits .

N arat h s mo n ograph co n tai n s a mi n e of i n val u able comparative

’

an atomi cal observatio n s with beau tifu l illu stratio n s based on

, ,

preparation s showin g the u tmost tech n ical skill an d the statis ,

tical i n formatio n is prepared an d compiled most carefu lly From .

a sci ent i fi c stan dpoin t the work suff ers from o n ly two draw
backs : the au thor s desire to demolish A eby s res ul ts eve n i n the
’ ’
,

min or details which have n o real bearin g on the problem an d his ,

equ ally stro n g desire to establish his View of the apical bron chi ‘ ’

an d the pri n ciple of migratio n i n the architect ure of the mam

‘ ’

malian bro n chial tree O n e wo uld like to believe that the gross
.

mechani cal con cepts un derlyi n g this theory were employed solely
as fig ures of speech for the p u rpose of Vis u alizin g comparative
,

relatio n s were they n ot un mistakably i n ten ded to be taken lit

,

er ally elaborated with i n fi ni t e an d repeated precisio n of detail

,

i n n early every part of the work an d were it n o t for the attempt ,

to prove their act u ality by o ntogen etic eviden ce .

1 52 GE O . s . HUNTINGT O N

The fallacy of these arg ume n ts is however demo n strated , ,

absol utely as soo n as the i n q uiry is tran sferred to more co n cl usive

mammalian forms than those selected by N arat h If i n place .
,

of the prevale nt mammalia n tree with close approximation of ,

the right eparterial an d fi rst ve n tral hyp arterial b u ds we exami n e ,

the same bro n chial districts i n embryos of a type i n which the

right eparterial compo n e n t is a derivative from the trachea all ,

possible do u b ts as to the correct gen etic i n terpretatio n of the

facts disappear at o n ce Here the eparterial an d the fi rst ve n tral
.

hyparterial bro n chi are separate an d disti n ct anlages from the

very fi rst appearan ce of the epithelial swelli n gs foreshadowi n g
their developme n t the former b u ddi n g from the right side of
,

the trachea the latter from the right st em b ron chus

,
.

There can n ever b e at an y stage an y q u estio n whatsoever of

, ,

a
‘
splitti n g off of the eparterial eleme n t from the fi rst ve n tral
’

a nl age or of its wan deri n g cran iad o n to the trachea The

‘ ’
.
,

facts ren der s u ch a s upposition n othi ng less than abs urd an d speak
c learly for the correct i n terpretatio n N arat h himself recog n izes
.

t h e deplorable weak n ess o f his positio n i n face of the act u al

e mbryological fi n di n g s H e devotes i n his work of 3 8 0 pages
.

o nly 1 5 li n es to this aspect of the s u bj ect H e says (p 3 33 ) . .

Ei n ige Schwierigkeite n ver u rsacht die Erk l arun g des En tstehe n s

der t racheal en Bro n chie n D i e E n twickl un gsgeschichte lehrt
.
,

dass beim Schaf die Kn o sp en v o rwolb un g sehr fril h z ei t i g an des

rechte n Seite der Trachea selber e n tsteht M an ko nn te bi s j etz t .
2

e i n U eb erg eh en der K n ospe vom L u n g en s ac k c h en auf die Tra

chea n icht con st at i ren ” This stateme n t is absol u tely tr u e b ut

.
,

the author might with eq u al assu ra n ce have added that s u ch a

phe n ome n o n n ever W ill be observed H e goes on to say : Es “ .

verliert dieser ei g en t hii mli che E n twickl un gsgan g j edoch sei n e

’

So n derstell ung we nn m an dara n fest h al t dass Trachea un d

, ,

Lunge ei nheitliche Bild un ge n si nd die ursp rii n gli ch oh n e G re n

z en i n ei n a n der ub erg eh en
” ,

N a rat h perhaps u n co n scio u sly

.
, ,

expresses here a far reachi n g tru th b u t evide n tly witho u t grasp

,

i ng its real si g n i fi can ce for the i n terpretatio n of bro n chial ev o l u

tio n i n the mammalia for he proceeds immediately to q u alify
,

I t li m i
a cs ne .
1 54 G EO . s . HUNTINGT O N

respiratory s urface spells a correspo n di n g i n crease i n the n umber

of the termi n al bro n chioles It is a simple mathematical propo .

si t i o n to prove that the n u mber of the ul timate bro n chioles an d

the co n sequ e n t area of the respiratory s u rface will depe n d with ,

the same type and ratio of divisio n of the bro n chial tree u po n ,

their caliber an d u pon the distan ces separati n g the origi n s of

the primary co n d u cti n g t ubes from the st em b ron chus .

Th u s i n the case of the right eparterial bro n ch u s (A ) the exte n t

of its respirat ory area is i n direct ratio to the distan ce X Y sep —

arat i n g it from the n ext adj ace n t derivative ( B) of the st emb ron

Fig . 6 S che ma of b n hi
ro c al an d t ra c h ea l d iv t i n
er a o of r i gh t e p art er a l i
b ron c h u s . A .
,
e p i lb n
a rt e r a ro c hu s ; B
.
,
fi r st v en t l h y p a
ra i
rt e r a l b ro n c h u s .

chu s the first ve n tral hyparterial bran ch In compari n g the

,
.

domi n an t type ( fig 6 ) with that obtai ni n g i n the artiodactyls

A
.

B
(fi g 6 ) the e nlarged respiratory capacity of the latter is meas
.
,

ured by the greater le n gth of the li n e X Y

—
.

The phyletic as well as the o n toge n etic i n terpretation of the

migratory theory wo uld hold that the tracheal eparterial bron
ch us i n the a n cestors of the modern artiodactyls was a bran ch of
the fi rst ve n tral bro n ch us which i n the co urse of evol utio n emi
,
‘

grated from this primitive site craniad fi rst o n to the st emb ron
’
,

chu s and the n still fu rther i n the same directio n beyo n d the bifur
,

cation a variable distan ce up alon g the right side of the trachea .

 P ULMONAR Y EVO LUTIO N IN M AMMAL I A 1 55

In all these wan deri n gs the bron chus retai n ed its own in dividu al
character always represen ti n g the same O dysseu s like phyletic
,
-

eleme n t of the an cestral bro n chial tree N o matter i n to what .

foreign territory the voyage leads the germin al cell group aways ,

carries with it the i nvisible mystic imp ulse to develop i n to a

sp eci fi c bro n ch u s O n the other han d it is evide n t that the act u al
.

facts are adeq u ately and completely met by simply recogn izi n g
that space disposition within the thorax permitted an d i n creased
respiratory requ ireme n ts deman ded i n the evol ution of the ,

P ro un g ul at e type the selecti on of a more cra n ially located poi n t

,

o n the primitive e n todermal p u lmo n ary t u be for the origi n of

the respiratory b u d correspo n di n g to the bro n chial eparterial

area of the domi n an t mammalian type eve n if this origin is thu s ,

made to fall withi n the tracheal segme n t E mbryo n al mu ta .

tio n s probably played a decidi n g r61e i n the evolution ary process

i n volved .

I believe there can be n o qu estio n as to which of these two

"

hypotheses is the more ten able an d co n forms more clearly to the

facts .

Fli n t (2 1 p 3 5) w h o has furn ished u s with a most careful an d

, .

detailed developme n tal history of the l un g i n a form ( pi g) with

ri ght eparterial tracheal bron chu s ( his L I) says : .

If we turn f or a m oment to the considerati on of the ori gin of L 1 .

,

we fi n d the br on chu s is a t ri fle m ore prec o ci ous b u t pr acti c ally si mu l ,

t an eou s with the sec ond lateral br an ch ( ou r V ) i n i t s ori g i n It i s I

.

se parate d fr om L ater al 2 by a c o n si d erable di st an ce If the views of .

W ill ach an d N arat h were corre ct thi s bran ch should n ot appe ar until
,

l ater an d sh oul d be trace able step by ste p from the b u d f orming

, , ,

right L ateral 2 to i t s fin al position on the trache a It s dire cti on i s .

pr acti c ally l at eralwards with a sc ar cely v i sible ten den cy t o po i n t v en

t ralw ards It woul d n ot then from the t o pography of i t s orig in be ar
.
, ,

any an al og y t o a d orsal bron chu s Fr om thi s di sti n c tly l ater al posi

.

ti on of i t s origin I h ave classed it am ong the l ateral group of bron chi

, ,

alth ou g h i n i t s sub se q u e n t g r owth o n e of its br an che s exten ds d own

, ,

int o the dorsal reg ion g iving the bron chus a cert ain superfi ci al resemb
l an ce t o th at series O n the other han d the l ower l ateral elements ,

g r ow v en t r al w ar ds l n the l ater e m bry o n i c st a g e s an d thu s al so l o se

their early stri ctly l at eral ch aracter Thi s m uch i s cert ain ; if L 1
'

. .

ari se s phyl o g eneti c ally fr om the d or sal g ro u p a View f or whi ch there ,

i s n o convin cin g pr oof ab solutely all tr ace of the mi g r ati on i s l ost i n

,
1 56 GE O . s . HUNTINGT O N
the pi g It orig in ates like o ne of the l ate al bron chi and sub se q uently
. r

d evel o p s i t s su perfi ci al re sem bl an ce t o the d o rsal elem e n ts W h at .

ever suppo t f or the rel ationshi p of the bro n chus t o the dorsal series
r ,

c an be dr awn fr om thi s f a c t i s m ulti plie d by the beh a vi or o f a l ater al

,

bran ch of left L 2 whi ch doe s exactly the sam e thin g i n an ad apt ativ e
.
,

pr o c ess o n the p art o f the br o n chu s t o a rel atively u n ob str u cte d

environm ent .

With the artiodactyls an d the ki n dred bro n chial types the

Migratio n Theory defin i t ely collapses an d the fi el d is cleared
phyloge n etically and o n toge n etically of the spec ulation s which
might be fo un ded on a misi n terpretatio n of less clear cu t c on ?
-

d i t i o n s prese n ted by other mammalian types

II I . SELE C TI O N TH E O R Y
The con cl u di n g sectio n of this paper presen ts the viewpoi n t
which has impressed itself u po n me d u rin g these st u dies If I .

dig nify these ded u ctio n s by the formal design ation of a theory
this is do n e solely with the p u rpose of establishi n g more sharply
the co n trast betwee n them an d the V iews co n sidered i n the pre
cedi n g pages A ct u ally they do n o t co n stit u te a theory so mu ch
.

as an attempt to collect coordi n ate an d i n terpret as far as pos

,

sible all the facts accessible to me beari n g u po n the problem of

the phyletic history of the Mammalian L un g At the o utset .

we are co n fro n ted by two q u estio n s u po n the a n swer to which

,

depe n ds the e n tire i n terpretatio n of the mammalian bro n chial

system a n d of the correlated orga n izatio n of the l un g .

D oes a caref ul co n sideratio n of all the established facts n ow i n

o ur possessio n warra n t u s i n co n ti n u i n g to mai n tai n the hypoth

esis of a si n gle mo n ophyletic primitive gro un d plan of mammalia n

bro n chial architect u re an d to derive all exta n t types from the
same by s u ccessive m o di fi c at i o n s ? O r are we o n the co n trary
deali n g with the res ults of adaptatio n of the primitive vertebrate
l un g to co nditio n s which have varied widely i n di ffere n t mam
malian prototypes accordi n g to thoracic str u ct u re and space
dispositio n i n the cra nial portio n of the body cavity activity of ,

respiratory excha n ge mode of locomotio n ge n eral e n viro n me n t

, , ,
.

a n d ma n y other determi ni n g factors which have res u lted i n the

1 58 G EO . s . HUNTINGTO N

the s ubsequ e n t stages i n crease

In in the e x te n t of the respira
tory epithelial area is obtai n ed b y : A . Stru ct ural mo dificat i ons

Fig . 7 S c he ma of p hy l t i v
e c e i
o lu t on of t he v e rt e b ra t e l un g . C on d u c t o ry
p a t h s c ro s s -h a t c h e d ; pi t y
re s ra o r a rea s st i pp l ed .

of the primitive l un g sacs B Stru ct ural m odi fi c at i on s of

-
. . the
primitive extrap ulmo n ary co n d u cti n g can als of the trachea an d

bro n chi .
 P ULMONAR Y E V O LUTIO N IN M MMA AL I A 1 59

A . S tru ctura l modifica ti on s o f the p ri mi ti ve lung—sacs

The fu rther developme n t of the primitive smooth walled l un g

tube dep ends u po n two s ubsequ e n t cha n ges :
.I With the exte n sio n of the l un g sac its circ umfere n ce de -

v el op s poi n ts of i n creased epithelial proliferatio n These areas .

of heighte n ed mitotic activity protru de as hollow epithelial b u ds ,

whose lumin a are i n ope n commun ication with the ce n tral cavity
of the lun g sac The latter thus becomes st u dded with closely
-
.

B
aggregated p ulmon ary vesicles or crypts (fi g 7 ) . .

The lun g of the Anure amphibian ( R an a) fur nishes a good

example of this phyletic stage .

A s the s u rface area of the l un g sac i n creases the previo u s un i

-

form b u ddin g of the p ulmo n ary vesicles is replaced by a smaller

n u mber of localized areas of more i n te n sive prol feratio n which
i ,

develop i nto sac like protru sion s By con ti n u ed ce n trifu gal b u d

—
.

di n g from their walls these repeat i n every detail the stru ctu re
of the primitive vesic ulated l un g sac an d i nitiate the s ubdivisio n
-
,

of the origi n al wide p ulmo n ary cavum i n to a number of cham

bers (fi g 7 ) The walls of the adj ace n t compartmen ts come
.
C
.

in to co n tact by their mesodermal i n vestme n t an d fu se The .

l un g n ow is see n o n section to be divi ded i n to a series of pockets

by septa apparen tly arisin g from the s urface an d directed toward
,

the ce n ter of the origi n al p ulmon ary cavum i n to whose axial ,

remn an t the i n divid u al pockets open by wide mouths The fused .

walls of the origin al ce n trifu gally developed ou tgrowths from the

primitive l un g sac carry the mesodermal blood vessels i n the
-

res ulti n g septa The mu ltiple compo n e n ts of this adva n ci n g

.

lun g have like the a n teceden t primitive l un g sac from which

,
-

they arose the capacity to i n itiate from their e n todermal li ni n g

,

epithelial b u ds each on e of whic h is equ ivale n t to one of the or

,
.
~

i g i n al poi n ts of proliferatio n from the primary l un g sac respo n s -

ible for the produ ctio n of the e n tire gen eratio n of the secon dary
s acs. The l un g n ow co n tai n s a ce n tral tu b u lar space the red uced ,

p ulmo n ary cavu m of the primitive lun g sac which leads by a -

,

n u mber of passages ve n trad a n d dorsad i n to a correspo n di n g

n umber of seco n dary chambers each a replica of the e n tire prim

,
1 60 G EO . s . HUNTINGT O N

i ti ve l un g of the earlier stage The e n toderm is still thro ugho u t

.

respiratory i n character (fi g 7 ) C
. .

A ny of the more simply co n str u cted l u n gs of the L acertili a

an d pal u dal Chelo n ia ill u strates this phyloge n etic stage i n the

evol utio n of the mammalian l un g The same stage is represe n ted .

i n the o n toge ny of the mammal d u e allowa n ce bei n g made f or ,

the smaller n u mber of the seco n dary chambers an d the i n creased

i ntervals betwee n their poi n ts of co nn ectio n with the axial l ume n ,

characters which foreshadow the fu t u re architect u ral plan of the

mammalia n l un g .

Co n ti nu ed exte n sio n of this evol utio n ary process leads i n the

well k n own way to the grad u al prod u ctio n of a racemose orga n ,

i n which each u ltimate compo n e n t is the morphological equ iv

al en t of the si n gle a n cestral primitive l un g sac Types are fur -
.

n i sh ed to the phyletic series by the l u n gs of the soft shelled -

chelo n ian s (A spi d on ect es E retmochelys S phargis )

, ,
.

II With this i n crease i n the complexity of the l un g t h e n ex t

phyletic advan ce is marked by a stru ctu ral modificatio n alon g
d efi n i t e li n es both of the i n trap ulmo n ary epitheli u m an d t h e
related mesoderm repeati n g at a n u mber of poi n ts the earlie r
,

di ffere n tiatio n s of the se tissues throu gh which the primitive

e n todermal lu n g t ube became divided i n to the extra p ulmo n ary
- —

bro n ch u s an d the primary l un g sac First alon g the axial li n e-

.

B
of the ce n tral p ulmo n ary cavu m (fi g 7 ) s ubsequ e n tly i n the .
,

same w ay alo n g the axial li n es of the series of separate chambers

O pe n i n g i n to it ( fi g 7 ) the respiratory epitheli um ass umes the
C
.
,

character of a co n du cti n g epitheli u m while the s ubj ace n t meso

derm di ffere n tiates i n to the s upporti n g str u ctu res of a bro n chu s .

These histoge n etic chan ges begi n at a n umber of separate poi n ts .

The i n divid u al an lagen the n becomes j oi n ed to form a t ub ular

co n d u cti n g system which co nn ects with the extra p ulmo n ary —

bro n ch u s The l un g has thu s become i n vaded by an i n tra

.

p ulmo n ary system of bro n chial tu bes ope n i n g i n to an axial ,

ca n al the stem bronchus which i n t urn is n ow co n ti nu o u s with

, ,

the extra p ulmo n ary bro n ch u s an d meets the correspo n di ng

-

str u ct ure of the opposite site at the bif urcatio n of the trachea
B
(fig s 7.
D
,
7 ) .The l un gs of certai n mari n e turtles ( Cheloni a
1 62 G EO . s . HUNTINGT O N

b uds from any poi n t of i t s s urface an d to develop its i n trin si c

p ulmo n ary organ izatio n i n accordan ce therewith The res ult .

i ng exte n sio n s of the primitive respiratory area are refl ect ed i n the
correspo n di n g modelin g of the con ductin g system and give ris e
to the sp ecifi c form of the bro n chial tree pertai ni n g to the type .

I n cl udi n g th u s withi n the limits of its an lage the commo n basic

materials for the differe n tiation of both of its stru ctu ral compon
e n ts the l un g is i n a position to meet all the evol ution ary
,

demands put u pon it by the en viro nme n t an d the p urpose it

serves i n the org amsm .

3 Compared with the early phyloge n etic stages of the verte

.

brate organ ( fig 7 ) i n which the e n tire available s u rface of t he

A
.
,

primitive l un g sac is respiratory and su pplied by the ex t rapul

-

m on ary bro n ch u s the evol utio n of the mammalian l un g deman ds

, ,

i n accordan ce with its ultimate desig n exte n t and stru ct u re t he

, ,

selecti on of a limited number of poi n ts of i n te n sive epithelial

b uddin g i n place of the precedin g ge n eral use of the e n tire poten

tial s urface This selection is directed an d regu lated by t he
.

exten t an d locatio n of the total peripheral respiratory area which

will ultimately represe n t the c u lmin atio n of the developme n t
derived from each of these poi nts This i n t u rn is govern ed di .

rec t ly by the amo un t an d locatio n of the i n tracoelomic spac e

available for p ulmo n ary ex ten sio n an d its topographical relation s .

The gro upi n g of these selected areas of epithelial prolif eration ,

an d the p a ttern which they prod u ce i n their relatio n to each

other an d to the st em b ron chu s will determin e the typ e of the

,

bron chial tree characteristic for each form Within this form .

the bro n chial type is tran smitted by heredity .

Af ter the primitive l un g sac has been i n vaded by the co n d ue

-

tory system an d the res ulti n g i n trap ulmo n ary bron chu s has
,

replaced the antecede n t ce n tral cavum p ulmo n ale the active ,

epithelial b uddi n g is tran sferred to the li nin g of the secon dary

l un g sacs each of which is now co nn ected by its own short affere n t
—

c an al with the axial st em b ro n chu s ( fi g 7 )

D
The latter is thu s. .

provided with the anlages of its main derivatives the ven tral ,

an d dorsal primary bro n chi These act i n tu rn as st emb ron chi

.

for the i n divid ual seco n dary l un g sacs repeati ng there the g en
-
,
 P ULMONARY EVO LUTI O N IN M MMA AL I A 1 63

etic process which origin ally led to the establishmen t of the main
primary i n trapu lmon ary st emb ron chus (fi g 7 ) L ike the latter
E
. .

they develop the series of secon dary lateral derivatives each ,

termi n ati n g i n a tertiary l un g sac which is the exact morpho

-
,

logical replica of the primary lun g sac on a smaller scale and

-
,

forms the an lage for the con ti nu ation of the process of pul
m on ary u n foldi n g on the origi n al developme n tal plan
, .

The primitive framework of the future bron chial tree has n ow

bee n laid down The actu al details of its organ ization i n any
.

form will depe n d upon the nu mber and position of the seco n dary
l un g sacs b uddin g from the primitive cavu m pulmon ale an d of
—

the prima ry bron chi based on them The further the develop
.

men t of each of these is carried toward the periphery an d the ,

smaller th e n umber of the res u lti n g u ltimate respiratory areas

becomes the lon ger will be the un occ u pied i n tervals of the stem
,

bro n ch u s betwee n the origi n s of the primary bron chi an d t he

g reater th eir i n itial caliber .

In two l un gs of eq u al vol ume an d efficie n cy the st emb ron ch us

of on e may carry a smaller n umber of larger derivatives while the ,

other will show a larger number of primary side bran ches each ,

supplyi n g a smaller peripheral respiratory area Thu s the two .

lun gs shown i n schemata (fig 8 A an d B) belon g to the same

.
,

bron chial type (A) havi n g six ve ntral an d eight dorsal sid e
,

bran ches redu ced i n (B) to three and fou r respectively

,
.

This nu merical ran ge of the bron chial organ ization prevails i n

the mammalian l un g stem proper ( lower lobe ) which i n gen eral ,

is characterized by the marked similarity i n the arran gemen t of

its lateral derivatives from the st emb ron chu s although their ,

nu mber varies co n siderably i n the di ffere n t ge n era an d orde r s

( from three to eight or even n i n e of the ve n tral bran ches ) .

Typical examples of (A) are seen i n the bro n chial trees of the
lun g st em i n t h e Bradi p o di d ae "
Choloep u s] an d i n H g ram of (B) i n ,

that of the carn ivores and lower primates .

Based on the precedin g con dition s the followin g facts may b e

emphasized

1 The e n toderm of the i n trap ulmo n ary bro n ch u s retai n s the

.

i nh ere n t poten cy derived from its phyloge n etic and o n togen e tic
'

,
1 64 G EO . s . HUNTINGTO N

predecessor the primitive en toderm al l un g t ube of developin g

,
-
,

lateral b uds from any poi n t of its circ umfere n ce .

2 In the mammal these poi n ts of epithelial spro u ti n g te n d to

.

arra n ge themselves mai nly i n a ve n tro lateral an d a dorsal -

Fi g . 8 S c h em a sh o w i n g n umer c al i v i ti
ar a on i n t he p i m y d iv t i v
r ar er a es o f

t he st e m b ro n c h u s a n d it s re s u l t s in t h e i g p i ph
r esu l t n er e ra l a re as o f di st ib ut i on
r .

lo n gitudin al row The former are the larger an d expan d further

.

toward the periphery the latter are smaller an d less exte n sively
,

developed They co n stitute the ve n tral an d dorsal primary

.

bran ches of the st emb ron chus an d form with it the fo un dation ,

of the bro n chial tree He n ce the st emb ron chus acquires a dorso
.
1 66 G EO . s . HUN TINGTO N

medial stretch there are fo u r of these poi n ts v i z the ve n tral , .

, ,

ve ntro medial medial an d dorso medial

-
,
-
.

O n e or more of these seco n dary poi n ts may give origi n to

secon dary or accessory bron chi or they may defa u lt .

In the first case depe n di n g u po n the n umber an d locatio n of

,

the poi n ts utilized the st em b ron chu s will be st udded more or less
,

profusely i n the i n tervals between the li n es of its pri n cipal

ve ntro lateral and dorsal bro n chi with smaller seco n dary accessory
-

bronchi In th e seco n d case these portio n s of the st emb ron chu s

.

are n aked Con ditio n s i n this respect vary at diff eren t horizo n tal
.

levels In ge n eral accessory bro n chi become more n um ero u s i n

.

proceedin g cran io cau dad alon g the st emb ron chus Thi s ao
-
.

cords with the more archeal type of bron chial distrib utio n i n the
cau dal reaches of the lun g st em They prepon derate nu merically .

i n the medial as compared with the lateral sector of the circ u m

feren ce of the st emb ron chus the former presen ti n g a greater
,

arc an d he n ce more poi nts for their developme n t than the latter
,
.

i n the proportio n of 4 : 2 The developmen t of lateral an d dors o

.

lateral accessory bro n chi is further restricted by the co urse of

the mai n p ulmo n ary artery which for the greater part of its
exten t lies typically again st the dorso lateral s urface of the stem -

bro n chu s betwee n the rows of the mai n ve n tro lateral and the
,
-

dorsal primary bro n chi The bu ddi n g of the epitheli um lead

.

i ng to the developme n t of the accessory bro n chi is n o t limited to

the mathematical poi nt of the compass i n dicated by their desi g
n atio n . Thu s a member of the dorso medial grou p may develop —

a nyw here betwee n the li n es D and M a ve n tro medial accessory ,

-

bro n ch us an ywhere betwee n M an d V etc If it occ upies the ,

.

latter poi nt it may form either on e of the rarer accessory bro n chi
passi n g directly ven trad or a ve n trally located b ud more com
,

m on ly swi n gs mesad i n attai ni n g its distrib u tio n an d s u pplies a

typical ven tro medial p ulmo n ary district Still further the mai n
-
.

primary dorsal an d ve n tral bro n chi represen t as above stated , ,

the st emb ron chi for the secon dary l un g sacs formi ng their axial —
,

co n du cti n g str uct u re L ike their prototype the mai n stem

.
,

bro n ch u s they retai n the same pote n cy for proliferation from

,

any poi n t of their circ u mfere n ce Thu s a give n ven tro medial .
-
 P ULMONARY EV O LUTI O N IN M AMMAL I A 1 67

d istrict may derive its bron chial s upply from the followi n g
sources
1 Typical ve n tro medial accessory bro n ch u s
.
-
.

2 A ccessory bro n ch u s arisi n g f urther ve n trad or dorsad

. .

3 V e n tral accessory bro n ch u s

. .

4 A ccessory bro n ch u s arisi n g from the an gle betwee n the

.

st em b ron ch u s an d o n e of its primary ve n tro lateral bra n ches -

.

5 From the proximal portio n of the latter itself

. .

This does n o t imply that the accessory bro n chus has migrated ‘

or wan dered from the poi nt of its i n cep tio n the ve n tro lateral
‘ ’
,
-

primary bron chu s fi rst to the an gle betwee n it an d the stem

,

bro n chu s an d the n to the st em b ron chu s itself b ut that the ven ,

tro medial p u lmo n ary area i n question can be s upplied by an

-

accessory bro n chu s derived from any one of several adj ace n t
poin ts on either the st emb ron chu s or on its mai n ven tro lateral -

bran ch This 1 s the very simple ex plan ation of all the observed
.

facts upo n which the e n tir e theory of migration of bro n chial

compo n en ts has bee n erected .

If three corrosio n preparatio n s of the adult l un g ill ustrati n g

three types of bro n chial s upply for a give n ven tro medial pul —

m on ary district the cardiac lobe ) are compared (fig .

101
,
2
they might s u ggest that the ven tro medial accessory
,
-

bro n chu s s u pplyi n g the iden tical peripheral area (X i n all thre e
l un gs had migrated from the i n itial positio n it occ u pied i n
‘ ’

fi g ure 1 0 o n the ve n tral bro n ch u s to the a n gle betwee n it an d

1

the st em b ron chu s (fig an d had reached by co n ti n u ed

. shift ‘

i n g or wan deri n g the ve n tro medial s u rface of the st em b ron

’ ‘ ’
-

chu s itself (fig .

Co n ditio n s ide n tical i n sig ni fican ce with those above o utli n ed

lie at the base of all other cases of s upposed migratio n of what ‘ ’

ever kin d or exte n t In o ur prese n t i n stan ce the soun d i n ter

.

p ret at i on recog n izes that the three bro n chi A B an d C sup

, , , , ,

plyi n g the same peripheral area X i n their termin al di st ri b u,

tio n are fun ctio n al equ ivale n ts

, .

In respect to deri va ti on they are n o t however the ide n tical , ,

morphological elemen t shifted by migration from 1 to 2 or 3 ,

b ut the res u lt of epithelial proliferatio n from three separate an d

1 68 GE O . s . HUNTINGTO N

distin ct poi nts on the respiratory area of the st emb ron chus or ,

of i t s ve n tro lateral derivative or of both and t he selectio n of

-
, ,

o n e of the three ava i lable poin ts for t h e developme n t of each of

the three typ es (fig 1 1 A B C) These three poin ts are fix ed

.
1
, , .

an d do n ot chan ge their positio n d u ri n g the i n divid u al o n to g en y .

Fig u re 11
Fig s . 10 an d 11 V i ti ar a on in or i g in of a v en t ro -m e di l b
a r on c h u s .

Wh en epithelial b uddi n g has started at any o ne of them it will

”
c ont i n u ef an d accordi n gly lead to the correspo n di n g ad ult c on
:
dition of either A B or C , ,
.

It happen s occasion ally that a peripheral l un g segme n t re

c ei v e s its bro n chial s u pply from two so u rces The termi n al .

alveoli of each con trib u ti n g bro n chu s are closed off withi n the
s e g me n t of the dist rict which it s upplies an d do n ot comm un i

c at e w i t h those of its colleag u e Together the two compo ne nts

i‘
.
170 G EO . s . HUNTINGTO N

in a s uffi cie n t n umber of i ndivid uals of each type i n the serie s ,

we might co n ceivably arrive at the followi n g results :

1 In the more primitive types the peripheral p ulmo n ary
.

area X is s upplied i n a large portion of the i n divid uals by bron chus

A i n a m u ch smaller n u mber by bro n ch u s B
,
.

2 The i n termediate gro up of g e n era shows a decided n umer

.

ical prepon deran ce of bron chu s B A few i n dividuals still carry .

bro n ch us A an d a still smaller nu mber have developed bro n ch u s

,

C e i ther alo n e or i n combi n atio n with B

,
.

3 Fi n ally i n a third gro u p comprisi n g the most rece n t a n d

.
,

advan ced forms bron ch u s C has become domi n an t There are

,
.

relatively few B bro n chi and an A bron ch us is o nly occasion ally

,

fo un d .

The n an d o nly then co uld we speak of a p hylogen eti c mi gra

, ,

ti on which has carried bro n ch u s A from its origi n al poi n t of

origi n o n the ve n tro lateral primary bro n ch u s thro u gh the a ng l e
-

site B to its fin al positio n C on the st em b ro n chu s .

In the fi rst of the above hypothetical gro u ps with bro n ch u s A ,

domin an t bro n ch u s B appears as a progressive phyletic varia n t

, .

In gro up 2 with B the prevale n t bro n ch u s A is an archeal rever

, ,

si on al C a progressive phyloge n etic varia n t

,
In gro up 3 both .
,

A an d B are reversio n s the former of the pro g on al the latter of

, ,

the at aval type E ve n the n the term migration is to be in ter

.
‘ ’

p ret ed o n ly fig urat i v ely .

It is true that for the p urpose of rapid examin atio n of a series

of bro n chial trees the con cept an d the term expressin g the same
,

are co nve nie n t and compact an d that they readily i n dicate the ,

cardin al di ffere n ces observed i n the compariso n of diverge n t

types F urther u sed i n the se n se above stated they co n vey
.
, ,

co n cisely certain broad evol utio n ary facts Speaki n g phylo .

ge n etically it might be n ot altogether obj ectio n able to refer to a

bro n ch u s as migrati n g or wan deri ng from its origin al location
‘ ’ ‘ ’

to a n ew site i n compari n g a more archeal type of bro n chial tree

with on e fu rther advan ced i n progressive evol utio n I have so .

u sed the term i n my earlier p ublicatio n But it sho uld be .

employed if at all with a clear defini t i on of its meani n g n ot as

, , , ,

N arat h has do n e as desig n ati n g an act u al shift occ u rri n g d uri n g

,

th e o n toge n y of the i ndivid ual form .

 P ULMONARY EV O LUTI O N IN MAMMAL I A 171

Phyloge n etic migration does n o t imply the bodily t ranspl an

t at i o n of a bro n chial a n lage It mean s the establishme n t of a
.

n ew focal poi n t of bro n chial epithelial proliferatio n whose prod

u c t is a n eomorph rep la ci n g the archeal bro n ch u s a n d from the , ,

vantage poi n t of its n ew relatio n to the bro n chial tree e nl argi ng

-
,

the former s area of respiratory distrib u tio n and thus e nhan cin g
’

its physiological value to the organ ism The older bro n ch us .

thu s replaced loses its earlier si gni fi can ce becomes red u ced and
, ,

either disappears or is co n ti nu ed as a s ubordin ated side bran ch -

.

A bro n ch us per se does n o t travel The phyletic imp ulse to

.

i n au gurate a n ew and more advantageously located bro n chial

derivative i n its place from whatever poi n t of t he uni versally
available e n toderm is best adapted to the p urpose does travel , .

This sums up the esse n tial differe n ce betwee n the migratory ‘ ’

an d
‘
selective theory of bro n chial evol u tio n
’
.

The theoretical example above addu ced co u ld of co urse n o t be

vis u alized l n the lo n g li ne of a sin gle phyletic series with t ypes
tra n smitted by direct i nheritan ce The evol u tio n ary periods
.

req uired for the accomplishme n t of s u ch chan ges exceed by far

the u tmost limit of preservatio n of n on fossilized organ ic tiss ue
-
.

An archeal type has perished ages before we recog nize the res ult
of its evol utio n i n the modern i n heritors But while the un .

broke n phyletic chain of mammalian p ulmo n ary evol utio n i n the

li n e of direct desce n t is n ot actu ally available its reco n stru ctio n ,

is made possible to a large exte n t by the comparative an alysis

of extan t types and by on toge n etic data Thu s for example th e
.
, ,

p ulmo n ary problem receives m u ch el u cidation from the det ailed

st u dy of the l un g i n gro ups like the followin g :
Perissodactyls i n comparison with Tapirus .

Hyrax an d the P rob osci deae .

U n gulates Sire nia some Cetacean s

, ,
.

R elation of Catarrhin e an d Platyrrhi n e Primates .

Terrestrial and A qu atic R ode n ts .

Hystricomorphs .

M u stelidae and Taxidea .

G iraff a an d the Camelidae .

Hippopotamu s i n comparison with the remai ni n g Artioda ctyls .

172 G EO . s . HUNT INGTO N

B S truc tur a l modifica ti ons of

. the pri mi ti ve ex trap u lmon ary
con du ctory chann el s o f the trachea an d bronchi , i n
thei r reacti on on the lung- stem

We have i n the precedi n g pages followed the mai n evolution ary

li nes leadi n g up to the developme n t of the most primitive typ e
of the mammalian l un g co n sisti n g of the archeal l un g stem and
,
-

i t s bro n chial distrib utio n ( fig A dditio nal exte n sio n of the

.

respiratory area of the l ung stem is as s ug g ested above possibl e

-
, ,

to a limited exten t by the i n creased developmen t of its ven tro

me di al accesso ry bro n chi leadin g to the produ ctio n of one or
more cardi ac lobes If th e respiratory deman d is further height
.

en ed t h e cra ni al portio n s of t h e l u n g stem are the fi rst to be called

-

u po n becau se pulmon ary exten sion is o n ly possible for topo

, ,

g raphical reaso n s i n the cran io ve n tral directio n

,
-
.

Th e fi rst ve n tral bro n ch u s is mobilized i n the service of th i s

n ew acqu isitio n of respiratory territo ry It e nl arges and i t s
.

fi rst cran ial side bra n ch ass u mes proportio n s which may bri n g
-

it fin ally to equ al or eve n to exceed the rest of the pare n t stem

V1
. A s the A scen di ng Bran ch (A ) of the fi rst ve n tral bro n chu s
it n ow co n trols an addition al p ulmo n ary district developed from
the l un g stem by a m odi fic at i on of the primitive i n trap ulmo n ary
-

bro n chial plan (fig A s this n ew area e n larges an d exte n ds

.

craniad an d ven trad it may acqu ire a greater i n depe n de n ce by

the developme n t of a cleft i n the mesoderm e n velopi n g the l un g ,

which the n as the a n lage of the mai n i nterlob ular i n cisu re di

, ,

vides the e n tire organ i n to a cran ial an d a cau dal segme n t (fig .

The former is the primitive u pper lobe (fig 1 4 U ) b u ilt u pon the .
, ,

first ve n tral bro n chu s ( V ) an d its asce n di n g bran ch (A ) The

1
.

latter is formed by the balan ce of the origi n al l un g stem sup -

,

plied b V the st emb ro n chu s an d its remai nin g primary ve n tral

derivatives cau dal to V ( V I
all the dorsal bro n chi (D
z l

an d the cardiac bro n ch u s ( C) ( fig 1 4 L s t) The res ult is a mam

.
,
.

malian l un g whose bron chial tree is co n stru cted i n con formity

with A eby s bilateral hyparterial organ izatio n ( type III )
’
.

L et u s follow this fi rst defini t e an d typical mammalian bro n

chial tree i n its further evol utio n ary progress toward the develop
me n t of a still greater peripheral respiratory exten sion .
1 74 G EO . s . HUNTINGT O N

upper lobe are too closely approximated on the bro n chial tree
,

to admit of mu ch additio n al peripheral un foldin g of their ter

mi n al bra n ches
.

Fig u re 14

The right st emb ron chus has n ever lost its archeal i nhere n t
poten cy of i n au g urati n g bro n chial proliferatio n from any part
of its circ umfere n ce Its dorso lateral s urface i n the stretch be
.
-

twee n the origin of V and the tracheal bifurcatio n is an especially

I
 P ULMONARY EV O LUTI O N IN MAMMAL I A 175

favorable site for the exercise of this power owi n g to local c on ,

d i t i on s described i n detail below ( c f i n fra p . A n ew

,
.

bro n chial b ud (fi g 1 4 Ep ) p u shes out i n to the i nterval developed

.
,

0 5( DII

6 ( VIII )

7 ( VIIZ)

Fig 15
u re

b y rotatio n of heart an d oesophag u s bet wee n the foreg ut sup ,

portin g the right vag u s n erve an d the right p u lmo n ary artery
, ,

a nd develop s i n to a bro n ch u s W hich eve n t u ally s upplies the cra n

i al area of the upper lobe an d e n ables it to expan d further This .

176 G EO . s . HUNTINGT O N

n ew eleme n t th u s added to the bro n chial tree of the right l un g ,

is the E parterial Bro n chu s ( fig 1 5 It arises from the

.
,

dorso lateral aspect of the st em b ron chu s cran ial to the origi n
-

of V an d n earer to the tracheal bifu rcatio n It lies behi n d

I
.

rather than a bove the mai n right p ulmo n ary artery which vessel ,

desce n ds c au do laterad over its ve n tral s u rface to dip d orsad

-

betwee n it an d V an d the n co n ti nu e its desce n t dorsal to the

1

remai n i n g ve n tro lateral primary bra n ches of the st emb ron chus
—
.

It might hence be defin ed more correctly as the Kat art eri al

Bro n ch u s b ut all arterial relatio n s are of very seco n dary mor
,

p h o l o g i c a l s i g n ifi can c e i n the a n alysis of bro n chial orga n izatio n .

A eby n amed it the E parterial Bro n ch u s an d this desig n atio n ,

sho u ld stan d for all time on the basis of historical priority n o ,

matter if i n i n divid ual mammalia n forms the bro n chu s starts

behi n d abov e or to the i nn er side of the artery These are
~

,
.
,

secon dary an d un importan t details The cardin al fact remai n s .

that A eby was the fi rst to recogni ze the ex i stence of the bron chu s
an d its morphological si g n ifi c an c e i n the in t r ap u l mo n ary archi

tect u re That fact out balan ces any baptismal slips its g od
.
—

father may have committed Besides t h ere are n umerou s mam

.
'

malian types i n which the design at io n is qu ite perti n ent It i s .

co n cise thoro u ghly i n corporated i n the an atomical literat u re

, ,

an d n o observer wo u ld be misled for a m om en t b y its u se i n cor fi

r ec t l y diag n osi n g any give n bro n chial tree S u rface i n dicatio n s

.

poin ti n g to a n ewly acquired i n depe nde n ce and greater freedom

of the cran ial segme n t of the precedi n g primitive upper lobe ‘ ’

are n ot wan ti n g i n many form s Th e di st ri ct of the n ew ep ar

.

t eri al bro n ch u s becomes separated from that of the fi rst ve n tral

hyparterial bro n chus by a correlated n ew fi ssure the secon dary ,

i nterlobar i n cis ure The former n ow con stit utes the defini t i v e
.

u pper lobe proper the latter forms the middle lobe

‘ ’
confi n ed
‘ ’
, ,

to the ve n tral p ulmon ary regio n (fi g 1 5 M Both are sepa

.
,

rated from the remai n der of t he archeal l un g stem the lower -

,
‘

lobe by the older mai n i n terlobar i n cis ure

,
’
.

O n the left side co n ditio n s remai n m u ch as they w ere before

the i n au gu ratio n of the right eparterial developme n t (fig .

There is ordi n arily n o left eparterial n eomorph bro n ch us The .

1 78 G EO . s . HUNTINGT O N

proximal portio n of the st emb ron chus cra n ial to the a n gle of the
tracheal bif urcatio n or eve n i n part from the right lateral wall
,

of the trachea E very positio n betwee n these two ex tremes is

.

e n co un tered i n the mammalian series These di ffere n ces depe n d .

u po n the poi n t selected for the developme n t of the eparterial

b u d i n the differe n t mammalian forms possessi n g the prevale n t

type of bro n chial organ izatio n I n divid ual differe n ces i n the.

level of origi n of the eparterial bro n chu s are also fo un d within

a si n gle species .

However placed i n this respect the right eparterial bro n ch u s ,

an d its f un ctio n al eq u ivale n t of the left l u n g the asce n di n g ,

bran ch (A ) of the fi rst left ve n tral hyparterial bro n chu s both ,

possess f un damen tal morphological characters of the highest i m

portan ce E ach serves for the n eomorph cran ial p ulmo n ary dis
.

t ri ct i n the same capacity as does the mai n st em b ron c h us for

the l un g stem co n stit u ti n g its exce n tric axial stru ct u re O n to
—
,
.

ge n etically this is the res ult of the ide n tical processes whi c h we
n oted above as respo n sible for the i n trap u lmo n ary orga n izatio n

of the primitive l un g sac ( of p -

. .

The eparterial b u d of the later phyletic stages of evol utio n is

the exact homologu e of a seco n dary l un g sac of the early embry -

o n al period an d develops its con du ctory system on exactly the

,

same morphogen etic li n es We speak for the sake of brevity

.
, ,

o f the a n lage of the eparterial bro n chu s In the strict se n se it

‘ ’
.

is n ot the bro n ch u s which b uds b ut the respiratory en todermal

‘ ’
,

o utgrowth which will later be represe n ted i n the bro n chial tree
by the eparterial bro n chu s ( fig s 1 3 an d 1 4 Ep ) This seco n dary
.
,
.

cran ial st em b ron chus for the upper lobe develops alo n g the same
path as its an cestral prototype the primary st em b ron ch us of ,

the l un g stem an d like it acq uires ve n tral and dorsal branches

-
, ,

a n d spari n gly accessory bro n chi

, ,
Its termi n al distrib utio n sup
.

plies the apex of the l un g j u st as the mai n st emb ron ch us breaks

,

u p i n to a leash of small bra n ches for the c au d o dorsal l un g pole - -

.

In this eval u atio n of Ep an d A as the st em b ro n ch i for the cra

,

n ial exte n sio n from the l un g stem of their respective sides their
-
,

d orsal an d ve n tral derivatives are see n to fall i n to serial li n e with

the correspo n di n g bran ches of the mai n st emb ron chus i n the
 P ULMONARY E V O LUTI O N IN MAMMAL I A 179

l un g stem (fi g s 1 4 an d 1 5 V V D D ) The total n umber of

-
.
,
l— 7
,
l— 7
.

the ven tral an d dorsal bro n chi for the l un g as a whole is thus i n
creased through the s upplemen tal eleme n ts con trib u ted to the
series by Ep an d A The seco n dary acquisitio n of the cran io
.

ve n tral p u lmo n ary segme n ts and their implan tatio n u po n the

primitive l un g stem forms the key to the phyletic i n terpretatio n
-

of the mammalian bro n chial tree an d of the p u lmon ary ev olu

tio n based thereon These grad u ally ac qu ired importan t mod
.

i fi ca t i on s of the primitive architect u ral type an d additio n s to i t , ,

an d the res u lti n g reorga n izatio n s are i n dicated schematically i n

the diagrams 1 2 to 1 5 sho wi n g a lateral profil e view of a right

,

l un g
.

Figu re 1 2 represen ts the archeal co n ditio n The st emb ron ch u s .

gives off fo ur larger ven tro lateral pri mary bro n chi ( V V ) an d
-
I IV —

an eq u al n u mber of shorter dorsal bra n ches (D D ) The

I IV f —
.

p ulmo n ary artery en ters t h e cran ial base of the l un g stem lateral -

to the extrap ulmo n ary bron chu s an d t urn s dorso cau dad over V I -

to descen d on the dorso lateral aspect of the st emb ron chus b e

-

twee n the ven tral an d dorsal primary bro n chi s u pplyi n g a bran ch
to each .

Figu re 1 3 : The cran ially directed derivatives of V ( stippled I

i n diagram ) e nl arge especially the fi rst (A ) foreshadowi n g the

, ,

eve n t u al cran io ven tral exte n sio n of the lun g an d the i n volve
-

men t of V i n the same

I
.

The dotted area (Ep ) on the lateral s urface of the ex t rapul

m on ary st em b ron chu s above an d behi n d the artery i n dicates
, ,

the poi n t of selectio n for the fu tu re developme n t of the bron chial

b ud whi ch wi ll eve n t u all y lead to the establishme n t of the ep ar
t eri al bro n ch u s.

Figure 1 4 : The cranial an d ven tral p ul mo n ary segmen ts have

exten ded greatly The former is b u ilt upo n the first cran ial de
.

ri vat i v e of V whi ch as its A scen di n g Bran ch (A ) n ow passes

I
, , , ,

well above the p ul mo n ary hi l u s alon g the trachea givi n g off ,

ven tral an d dorsal side bron chi an d termi n ati n g i n the di st ri b u

-

tio n to the apex of the lun g .

After s upplyi n g its asce n d in g bran ch (A ) to the crani al de

v el o p m en t
,
1
V co n ti nu es its origi n al co urse ve n trad the fi rst ,
1 80 G EO . s . HUNTINGT O N

ve n tral bran ch of the p ulmo n ary artery keepi n g pace V an d .

I

A together n o w s upply the n ew cra nio ve n tral p ulmo n ary terri -

tory which te n ds to separate by fissurat i on as the primary upper

,
‘

lobe from the remai n der of the archeal l un g stem The stem
’
-
.

bro n chu s havi ng yielded its fi rst ve n tral primary bro n ch u s V

,
I

to the n ewly acqu ired more mobile district s upplies with its ,

remai n in g ve n tral ( V V ) an d with all its dorsal bran ches

H IV —

( D
I
D IV
) h d fi i i l g stem co n stit u ti n g the lower lobe
'

‘ ’
—
t e e n
s
t v e un -
,

of the more highly developed mammalian l un g The left l un g .

of the domin an t mammalian type does n o t pass beyo n d this

evol u tion ary stage and hen ce retai n s a more archeal character
than the organ of the right side i n the great maj ority of the mam
malia In the S pecial gro ups with bilateral eparterial organ iza
.

tio n ( pi nn ipedia cetacean s camelidae ) the left l un g follows more

, , ,

or less closely i n the wake of the right p ulmo n ary exte n sion .

In fi g ure 1 4 the an lage of Ep desti n ed to s upplan t A i n the

,
.

progress of further evol utio n ary advan ce is show n schematically ,

as an epithelial b u d s upplied by a special bran ch of the p ulmo n ary

artery .

The asce n din g bran ch A of V varies i n relative size as compared

I

with the pare n t stem i n di fferen t mammalian types as the n at ,

u ral res u lt of its phyletic developme n t .

1 It may be of smaller caliber than V appeari n g the n merely

I
.
,

as a slightly en larged side bran ch In this case the n ew pul

—
.

m on ary territory is located chi efly i n the ve n tral area the ,

cra n ial exte n sio n remai n i n g small .

.2 The fi rst ve n tral bro n ch u s may divide i n to two equ al

bran ches A an d V The primary upper lobe then exte n ds
,
I
.

equ ally cran iad an d ve n trad .

.3 The fi rst ve n tral bro n ch u s arches almost directly from its

origin cran iad (A ) represe n ti n g the mai n chann el while the con
, ,

t i n u at i o n of V appears as its most cau dal ve n tral derivative

I
.

P ulmo n ary exte n sio n is the n chi efly cran iad .

The three co n ditio n s ill u strate s uccessive stages i n the evo

l ut i on of the cra nial l un g segme n t D iagram 1 4 is based on the
.

seco n d .
1 82 G EO . s . HUNTINGT O N

bro n chu s an d its derivatives with the exceptio n of the fi rst v en ,

tral hyparterial bro n ch u s ceded to the n ew cra nio ve n tral dis -

t ri ct s
. It shows a remarkable reg ularity i n the un iform type of
its co n d uctory system an d i n the bilateral homology of its i n di
,

vid u al compo n e n ts The more rece n tly acquired seco n dary

.

cran io ve n tral n eomorph requires for its organ izatio n the de

-

v el op m en t of a n ew axial co n d u ctory ca n al occ upyi n g withi n ,

the additio n al territory however restricted or exten ded this

,

may b e a pos ition strictly an alogo u s to that of the st em b ron chus

,

of the older caudal p ulmo n ary compo n en t Like this it develops .

its ven tral an d dorsal side bran ches an d its termi n al distrib u tion .

It varies i n the exte n t an d arran gemen t of these eleme n ts with

the varyi n g ran ge an d disposition of the p ulmon ary area un der
its co n trol It likewise varies i n its poi n t of derivation from the
.

archeal i n trap ulmon ary bro n chial system of its predecessor the ,

primitive l un g stem The available ran ge of its origi n exte n ds

-
.

cran iad from the latter s base In the i nitial and lower phases
’
.

of its developme n t it appears as an exaggerated bran ch of the

first ve n tral hyparterial bro n ch u s A s it un folds a n d aims at .

the co n trol of a more exte n ded territory this origi n al poi n t of its
origi n proves i n adequ ate i n accordan ce with the law governi n g
,

the relation betwee n the peripheral expan sion of a bro n ch u s an d

the distan ce separati n g its origi n from the adj ace n t bran ches of
equ al rank The n eighborhood has become too crowded so it
.
,

moves But it does n ot carry its old hou se with i t It selects a

. .

site i n territory less thickly settled and begin s to b uil d a n ew on e ,

.

It does n o t migrate it emigrates It selects a poi n t on the

‘
,
’ ‘
.
’

still un occ upied stretch of the st em b ro n chu s If the lo n g estab .

l i sh ed path of the mai n p u lmo n ary artery s upplyi n g the archeal

system lies i n the way it s urmo un ts it an d fix es on a S pot b e
,

yo n d confi dent i n the k n owledge that the old stream c an be

,

tapped to furn ish the n ecessary s upply for the n ew habitatio n .

In doi n g this the bro n ch u s has n o t o n ly aban do n ed its a n cestral

birthplace it also has chan ged its citize n ship It leaves the
,
.

archeal hyparterial comm un ity of its fello ws an d appears as the

pio n eer fo un der of the n ew eparterial colo n y The on ly path for .

still further advan ce lies ope n cran iad D epen din g u pon the .
 P ULMONARY EV O LUTI O N IN MAMMAL I A 1 83

dema n ds for i n creased p u lmo n ary area the n ew arrival appro

p ri at es a still more proximal portion of the st em b ro n c hu s The .

tracheal bifurcatio n the trachea itself to a varyi n g height fur

, ,

n ish the epithelial fu n dame n t for i n creased respiratory exte n sio n

i n n ew poi n ts of eparterial bro n chial derivatio n from the origi n al

stem li n e
-
.

In all cas es the available i n trathoracic ran ge of eparterial

exte n sio n is more or less restricted so that the opportun ity or
,

n ecessity for the developme n t of more tha n a si n gle eparterial

bron chu s does n ot arise The anl age for it has o nly to b u d far
.

en o ugh away from its n earest hyparterial n eighbor to obtai n all

the space requ isite or available for its developmen t Still th e .

possibility for the un foldin g of more than on e eparterial b u d is

i nhere n t i n the organi zatio n as shown by i n stan ces of do u ble
,

eparterial bro n chi i n a few forms (G iraffa Camelu s Au che nia , , ,

D elphi nu s Myrmecophaga) i n which a large tracheal eparterial

, ,

bro n chu s is accompan ied by a smaller eparterial con trib u tion .

This is u s ually derived from the right stem bro n chu s b ut i n ,

G ira ffa arises from the trachea cau dal to the mai n eparterial
trun k .

F actors resp on si ble f or the p reva len t a symmetry of the mammali an

lun g

In the domi n an t type of mammalian i n trap u lmo n ary archi

tect u re eparterial developmen t is c on fin ed to the right l un g with ,

the eparterial bro n chu s derived from the right st em b ron chu s .

The res ulti n g p u lmon ary asym metry is e n co un tere d i n at least

95 per ce n t of the mammalia n ge n era a n d species whose bro n
chial trees are recorded an d he n ce calls for detailed co n sidera
,

tio n i n any critical an alysi s of mammalian p ulmon ary evolutio n .

The qu estio n has lo n g attracted atte n tio n and the etiological

factors held respon sible for the prevale n t co n ditio n have bee n
disc u ssed i n a ge n eral way .

The e n croachme n t of the heart on the left half of the thoracic

space is evide n tly the cau se of the relative redu ctio n an d loss of
volume of the left as compared with the right l un g This is .
1 84 G EO . s . HUNTINGT O N

born e out by the fact that i n the aq u atic mammals ( pi nn ipedia ,

cetacea sire n ia) with bilateral p ulmo n ary sym metry the heart
, ,

occ upies frequ e n tly a striki n g median positio n exte n di n g n early ,

equally i n to both sides of the thorax It is of co urse impossible .

to decide whether this is the origi n al co n ditio n favori n g sym

metrical p ulmo n ary developme n t or whether the i n itial devi ,

atio n of the heart to the left has been corrected seco n darily by
an i n crease i n the vol ume of the left l un g an d possibly by the

operation of hydrostatic factors .

Fli n t (2 1 ) regards the u n paired right eparterial bro n ch u s as

the n ormal co n ditio n of mammalia d ue to a phyloge n etic pro ,

visio n for the desce n t of the heart an d great vessels thro ugh the
s uppression of the eleme n t of the left side .

Fli n t explai n s the mechan ical factor which he co n siders opera

tive i n preve n ti n g the developme n t of a left eparterial bro n ch u s
as follows (1 c p . .
,
.

In the de scent of the aorti c a ch an d the D u ctus arteri osus duri ng

r

embryon i c life from a point ab o ve the ori gin of L ateral 1 t o a point

bel ow we h a ve an expl an ati on fo the supp essi on of thi s elem ent on
,
r r

t h e left si de f o r if thi s br o n chu s were f orm e d b oth aort a an d the

, ,

Bo t al li an du ct wou l d be c au ght u p on it a n d t h ei r desc en t p ev e nte d

°

r .

L ikewi se the V e n a pu lm on ali s a pp e ar s i n the m i dlin e an d i s c arrie d t o

the left un til it fin ally re sts o n the portion of the stem where a left
Ve n tr al 2 shoul d d evel o p The u su al su ppression of the se t w o elem en ts
.
,

therefo e m ust be l ooke d u pon as a phyl og en eti c pro vi sion t o allow

r ,

f o r the d esce n t of the g e at ve ssel s o n the on e h an d an d the shifti n g

r

of the V e n a pul m o n ali s o n the o ther It i s n oteworthy th at i n those

.

an im als where the se br o n chi a e f o rm e d o n b o th si de s

r they a e so ,
r

sit uate d as t o o ffer n o re si st an c e t o either of the se fe at ure s of the d evel

o pmen t o f the g re at ve ssel s .

It is di ffi c ult to see j u st what Fli n t mean s i n ass umi n g archi

tecto n ic di ffere n ces i n the developme n tal gro un dpla n of mam
malia possessi n g bilateral eparterial bro n chi which favor sym
metrical bro n chial un foldi ng i n them whereas it is preve n ted on ,

the left side i n the domi n an t mammalia n type It is also di ffi .

c ult to bri n g his explan atio n of p ulmo n ary asymmetry i n to li n e

w ith the h uma n i n sta n ces of dextro thoracic aortae witho u t -

situ s i n vers us i n which the aorta u ses the right fo urth arch an d
,
186 G EO . s . HUNTINGT O N

also exists (pi nnipedia perissodactyls camelidae aq u atic ro

, , ,

de n ts arboreal primates Hippopotam us ) The early o n to

, , .

gen etic stages mu st therefore offer opport un ities which corre

sp o n d to the above fi n di n g s i n the ad ult The ve n tral wall of .

the foreg ut with the vagi forms i n rabbit embryos of 1 0 mm .

to 1 1 mm the backgro un d s upporti n g the tracheal bifurcatio n

.

an d the st em b ro n chi with their primary b u ds The p ulmo n ary .

arteries accede to the ve n tral lateral circ umfere n ce of the trachea

-

alo n g which they at fi rst desce n d i n a fairly symmetrical co urse .

In approachi n g the tracheal bif urcatio n the right artery grad u ally
in clin es ve n trad while the correspon din g vessel of the left side
,

t u rn s dorso cau dad This is the res ult of the rotatio n of the
—
.

heart an d its arterial pedicle to the left At the same time the .

foreg u t carryi n g the two vagus tr un ks un dergoes the axial rota

tio n to the right thro ugh which the left side of the gastric
en largeme n t attai n s its directio n ve n trad .

A s the res ult of these rotatio n s of heart an d foreg ut i n opposite

directio n s the topographical co n dition s are chan ged on the two
sides The left vag us an d left p u lmo n ary artery grad u ally
.

approach each other an d are at the level of the tracheal bifu rca
,

tio n an d origi n of the st emb ron chi i n close co n tact The reverse
,
.

obtai n s on the right side The right p ulmo n ary artery t u r n s

.

more an d more ve n trad i n desce n din g while the right vagus , ,

followi n g the rotatio n of the foreg ut moves dorsad An arterio ,

.

n e u ral i n terval is th u s ope n ed up on the right side toward which

the latero dorsal circ umfere n ce of the right st em b ron chu s faces
-

directly an d i n to which it se n ds the right eparterial bro n chial

b ud . The latter thus comes to lie i n fro n t of the right vag u s
an d right side of the oesophag u s an d behi n d the right pl um on ary
,

artery O n the left side this arterio n e u ral portal for eparterial
.
-

developme n t has bee n blocked by the approximatio n of left

vagu s and left p u lmo n ary artery or red u ced to s u ch an exte n t
,

that it n o lo n ger affords a favorable path for the protr usio n of

an eparterial b u d from the left st em b r o n ch u s The fi rst pri .

mary derivative of the latter is he n ce forced to pass ve n tral to

the artery and th u s becomes the fi rst ve n tral hyparterial bro n
chu s of the left side This s upplies thro ugh its large asce n di n g
.
,
 P ULMONARY EV O LUTI O N IN M AMMAL I A

bran ch the cran ial pole of the left l un g correspo n di n g to the ,

area which on the right side receives the eparterial bro n ch u s .

The n arrow i n terval betwee n left vag u s an d left p ulmo n ary

artery may at times s uffice for the passage of a b ud derived i n
this sit uatio n from the left st emb ron chus In s u ch i n divid u als .

the varian ts me n tion ed below ( cf p 1 9 3 ) wo uld occ ur The

. .

n ormal asce n di n g bra n ch from the fi r st left ve n tral hyparterial

bro n chu s u s ually s upplyi n g this segme n t would the n be corre

, ,

sp o n di n g ly red u ced or defau lt altogether .

It is si g ni fi cant to n ote i n this co nn ection that i n the rabbit ,

i n which form both the adu lt variatio n s above me n tio n ed an d

their embryo ni c anlagen have bee n recorded ( cf p the . .

cra ni o ve n tral p ulmo n ary exten sio n formi n g the upp er pole of
—
,

the left l un g is very m u ch red u ced compared with the right side
,
.

This is especially marked i n some wild species as i n L ep us cam ,

p estms The imp ulse to sen d an eparterial b ud from the left

’

st em b ron ch u s thro u gh the n arrow vago arterial i n terval i n -

,

S pite of the red u ced space available wo uld be fostered un der

,

s u ch co n dition s by the smaller p ulmon ary area which it wo uld

be called upo n to serve .

I believe that the co n ditio n s j u st ou tli n ed f urn ish the adequ ate
o n toge n etic explan atio n of the prevalen t asymmetry of the mam
malian bro n chial tree an d that this is based primarily on the
,

differe n ce i n the opportun ity for cran io ve n tral p u lmo n ary -

exte n sio n afforded n ormally to the right an d left l un g resp ec

t i v el y i n co n seq u e n ce of the cardiac an d oesophageal rotatio n i n
,

opposite direction s The si nistral t u rn of the heart an d its

.
,

effect on the i n itial directio n imparted to the p ulmo n ary arteries ,

depe n ds upo n the typical developme n t of the left sided mam -

malian aorta an d the rete n tio n of the dorsal segme n t of the left
sixth aortic arch as the D u ct u s Bot all i This stru ct ure fix es the
.

cardiac twist an d t ur n s the left p ulmo n ary artery dorsad while ,

i n co n trast the right p ulmo n ary artery freed by the obliteratio n

,

of the right sixth aortic arch i n its dorsal portio n t urn s ve n trad ,

i n respo n se to the rotatio n The e n ormo u s mechan ical force

.

exerted by the Bo t alli an D u ct o n the positio n of the heart an d

o n the ad ult arterial patter n is striki n gly ill u strated by compar
188 G EO . s . HUNTINGT O N

i ng the freq u e ntly observed cardi n al variatio n of a left s ubclavia n

artery arisi ng from the n ormal thoracic aorta with the rare case ,

i n which the same variatio n of the primary arterial bra n ches is

combi n ed with a ri ght sided thoracic aorta an d the rete n tio n of
-

a left Bot alli an D uct .

The facts observed d uri n g the o n toge n y of the domi n a n t p ul

m o n ary type i n the mammalia acco un t for the co n ditio n s fo un d
i n the ad ult An y exte n sive series of typical mammalia n p u l
.

m c n ary corros i o n s will show that i n the great m aj ority the car
diac rotatio n has carried the origi n of the mai n p ulmo n ary tr un k
so far ve n trad an d to the left that the primary division of the
vessel takes place to the left of the trachea or of its bifu rcatio n .

The lon ger right p ulmon ary artery he n ce crosses the ve n tral sur
face of the lower en d of the trachea or its bifurcatio n from left to
right In desce n di n g to the poi n t of its i n tersectio n with the
.

dorso lateral s urface of the st em b ron chu s i n the an gle formed by

-

the fi rst ve n tral bro n chu s with the latter the artery lies chiefly
,

ve n tral to the proximal portio n of the right st em b ron chus b e ,

twee n the origi n of its fi rst ve n tral bran ch an d the tracheal

bifu rcatio n The cran io lateral s u rface of the st emb ron chu s th us
.
-

left free is occ upied by the origi n of the right eparterial bro n ch u s .

The shorter left p ulmo n ary trun k t urn s almost sagittally dorso
cau dad coveri n g directly t he cran io lateral circ umfere n ce of the
,
-

left st emb ron ch us i n the i n terval betwee n the tracheal bifurca

tio n an d the origi n of the first ve n tral hyparterial bro n ch u s .

The artery th u s occ upies the larger part of the segme n t which
wo uld be otherwise available for left eparterial developme n t .

If the ge n eral premises of the selectio n theory are well take n

a n d if he n ce the eparterial compo n e n ts of the bro n chial tree

are s ec o n da ri l y e n grafted d u ri n g evol utio n o n the archeal l un g

'

stem the m ut ual relatio n s of the latter were already established

,

at a time whe n additio n al cra nio ve n tral exte n sio n of the l un g

—

w as i n a u g urated The eparterial acquisitio n m u st fit i n to an d

.

become adapted to already existi n g co n ditio n s which are more ,

favorable o n the right as compared with the left side .

I co n sider these facts as chiefly respo n sible for the prepo n der
an t developme n t of the eparterial bro n ch u s o n ly o n the right side ,
1 90 GEO . s . HUNTINGT O N

S elec ti on theory

S ummary

The theory of selectio n is based on the morphological an d

physiological co n ditio n s which have combi n ed i n the evol utio n
of the mammalian p u lmo n ary organ izatio n .

I M orp hogen ett c p rm ctp les The phyletic i n terpretatio n s o f

'

the extan t types of bro n chial architect ure i n the mammalia and ,

the h om ol o g i zat i on of the right and left bro n chial compo n e n ts

i n the asymmetrical forms does n o t depe n d u po n the rearran ge
,

men t of afix ed number of primary bro n chi derived from a hypo

thetical archeal bro n chial tree and shifted withi n this frame i n the
vario us types m u ch as chessme n are moved aro un d on the
,

board It rests on the fact that the e n tire e n todermal respira

.

tory an lage of the mammal has evolved o n the basic p ulmo n ary
organ izatio n tra n smitted from its b at rach i o reptilian a n cestry -
,

shari n g i n all the developme n tal pote n cies of the latter It is .

1

n o t o n ly unn ecessary to s u ppose that exta n t reptilia n types if ,

determi n ed adeq u ately wo uld yield an un broke n and closely

,

graded series leadi n g directly to the mammalian p ulmo n ary or

g ani z at i on b u t all the available phyloge n etic o n toge n etic an d
, , ,

comparative evide n ce n egatives s uch an ass umptio n .

It is probable as o utlin ed above that the promammalia emer g ed

, ,

from a reptilian stem with l un gs correspo n din g approximately

to the organ s of the simpler modern lacertilia or at most ad ,

van c ed to the stage of p u lmo n ary evol u tio n occ u pied to day by —

the more primitive pal u dal and littoral chelo nia S u ch a l un g .

prese n ted the p ulmo n ary cav um still co n tinu o usly lin ed by
respiratory en toderm The more complex and highly organ ized
.

l un gs of the mari n e chelo nia and of the crocodilia developed the

di ffere n tiation of the i n trap ulmo n ary bro n chial system seco n d ,

ari ly e n grafted o n the primitive str u ct ure ( c f p an d rep . .

rese n t specialized adaptatio n s co n ti nuing the li n e of reptilian

developme n t beyo n d the poi n t at which the mammalian deriva
tio n left the commo n a n cestral stock In this they follow the .

same un derlyi n g ge n etic laws which determi n e the further course

of the mammalia n evol utio n b ut they lead to higher stages of
,
 P ULMONARY E V O LUTI O N IN MAM MAL I A 1 91

p ul mo n ary developme n t distin ctly reptilian i n the character of

its details an d diverge nt from those attain ed by the mammalia .

The archeal promammalian l un g t u be possessed like its lacer

—
,

t ilia n prototype the pote n cy of differe n tiati n g co n d u ctory li n es

,

an d peripheral respiratory areas by a selective developme n t of

bro n chial b u ds from any poi n t on the e n todermal s urface fol ,

lowi n g t he m orp h og en et i c li n es detailed above for the reptile

‘
.

The n umber of these primary foci of heighte n ed mitotic activity

leadi n g to epithelial proliferatio n an d their positio n relative to
,

each other an d to the axial p ulmo n ary can al determi n ed the

,

bron chi a l typ e of distrib u tio n .They m u st have from their i n

c ep t i o n represe n ted the sum of the reactio n s of the m i lie u o n a

p ulmo n ary organ ization which at its emerge n ce from the rep
,

tilian stem w as i n a highly plastic co n ditio n an d respo n sive to

,

n ew e n viro n me n tal an d metabolic factors It is he n ce qu ite

possible that the variou s types of the modern m
.

ammalian l un g
o w e their diversity i n part to pol yphyletic derivatio n The .

bro n chial organ ization best adapted to respiratory co n ditio n s

obtaini n g at any one evol utio n ary phase for each mamm alia n
form became fix ed for that form by heredity as lon g as the
respiratory factor of the e n viro n me n t remai n ed un chan ged .

The extan t ordi n al ge n eric an d sp ecifi c types are the resu lt of

,

the tran smissio n of selective pattern s to the desce n dan ts .

A ll the available evide n ce goes to show that i n certai n mam

malian groups the more primitive bro n chial types have d u ri n g ,

t h e progress of this evol u tio n un dergo n e m o di fi c at i o n s some

, ,

times of wide import i n respo n se to altered e n viro nm en tal an d

,

fun ction al deman ds The opport un ity for the con ti nued devel
.

o p m en t of s u ch seco n darily acq u ired adaptatio n s rests i n the per

sisten t morphoge n etic plasticity of the bro n chial system an d the

rete n tion by it of the poten cy derived from the primitive an ces
,

tral e n todermal l un g t ube of i n au g urati n g additio n al or atypical

—
,

poi n ts of epithelial activity u n der the appropriate stimuli leadi n g,

to the produ ctio n of n eomorph bro n chial compo n e n ts .

I base these premises on three sets of facts :

I The above o utli n ed phyletic developme n t of the bro n chial
.

s ystem i n the vertebrate series .

1 92 G EO . s . HUNTINGT O N

II The m u t u ally i n tercha n geable character of co n d u ctory

.

an d respiratory a n lages i n the most primitive mammalia .

III V arian t fl uct uatio n s an d m utatio n s of the mammalia n

.

bro n chial system .

I Cf s u pra p 1 57
. .
,
. .

II The l un gs of A place ntalia develop o n li n es ide n tical with

.

those determi n i n g the place n tal p u lmo n ary u n foldi n g u n til the
mai n feat ures of the fut u re bro n chial patter n characteristic of
the ad ult are cl early mapped out i n the dispositio n of the pul
‘

m on ary e n toderm ( 5 3 2 A t birth u n der the stress of

‘ ’
, , ,

altered physiological co n ditio n s they revert for a period to an ,

earlier phyloge n etic phase The bro n chial patter n is wiped ou t .

by the s u dde n diste n sio n of the l un g an d the en tire f ut u re c on

d u c t o ry apparat u s is employed for respiratory excha n ge We .

are probably n ot far from right if we pict ure the early p rom am
malian l un g as prese n ti n g abo u t the co n ditio n s see n i n the third
o ntoge n etic stage of the mo n otreme an d mars u pial organ i n ,

which the immat ure embryo n ic l un g at the mome n t of the

‘
birth is blow n up by the fi rst i ntake of air an d reverts tempo
’ ‘ ’

rari ly to its reptilia n prototype The apl acen t ali an l un g of this .

stage s uggests eve n ts which might have occ urred if a reptilian

l un g began to experime n t with adaptatio n s to mammalian o r
g ani z at i o n an d the experime n t w a s i n terr upted
,
The grad u al .

ret urn of the aplace n tal l un g d uri n g the fo u rth developme n tal ,

period to the temporarily aba n do n ed type of place n tal bro n chial

,

organ izatio n poi n ts the evol utio n ary path alo n g which the rep
tilian l un g attai n ed the mammalian form The aplace n tal o n to .

ge n etic p ulmo n ary cycle demo n strates the developme n tal pote n
t i ali ty of the primitive mammalia n p u lmo n ary a nlage It .

proves that the early st emb ron chu s an d its primary derivatives ,

eve n after they are gro uped i n to the mai n li n es of the fu t u re

bro n chial pattern are n ot yet bro n chial str u ct ures i n the se n se
,

i n which w e employ the term i n ad ult a n atomy altho u gh they ,

o utli n e clearly the path alo n g which the s ubseq ue n t differe n tia
tio n of the co n d uctory passages an d the respiratory alveoli
occ urs The early bro n chial system of the place n tal embryo
.

still retai n s the archeal character of a pote n tial respiratory

1 94 G EO . s . HUNTINGT O N

In a series of examples of Eri n aceus europ aeus recorded by

29
N ara t h p 1 3 1 ) 2.0 specime n s carried o n ly the u s u al right
sided eparterial bro n ch u s b ut i n 9 i ndividu als there was a
,

bilateral eparterial developme n t .

These are examples of exceedin gly variable species with un ,

stable bro n chial organ izatio n b ut the same co n ditio n s obtain to a

,

greater or lesser degree thro ugho u t the mammalian class They .

are i n stan ces of the selectio n by the primitive e n todermal anlage

of developme n tal li n es of bro n chial b u ddin g which are atypical
for the species co n cern ed altho u gh typ ical for some other forms
,
.

Their appeara n ce as mu tatio n s is re n dered possible thro u gh the

u n iversal plasticity of the embryo n al a n lage Thu s the primitive .

l un g t ube with the typical plan of its unfoldin g determi n ed by

-
,

heredity for each species yet retain s the archeal pote n cy of devel
,

O pi n g bro n chial a n lage n from any poi n t of its e n todermal li n i n g ,

an d is he n ce able to prod u ce i n the i n divid u al both m i n or depar

tu res from the reg ular type an d cardin al varian ts whi ch alter
,

the morphological character of the bro n chial tree as i n the i n ,

stan ces cited .

I I P hysi ologi ca l fac tors op era ti ve i n p u lmon ary evolu ti on The .

o u tstan din g fact i n all vertebrate p ulmo n ary evol u tion is the
i n crease i n stru ct u ral complexity an d the resultan t heighte n ed
fun ctio n al activity of the cran io ve n tral as compared with the -

more archeal cau dal l un g segme n ts This distin ctio n begi n s to .

appear far back i n the reptilian stem It is well defined i n the .

higher lacertilian l un g becomes domi n an t i n the pal u dal and

,

mari n e chelon ia an d reaches its highest developme n t i n the

crocodilia It is a pro n o un ced factor i n the organ izatio n of the
.

avian l un g modified by the pec uliar morphoge n esis of the organ

,

i n this class an d forms the g uidi n g li n e as above detailed i n

, , ,

the i n terpretatio n of the mammalian p ulmo n ary evol u tio n .

Selectio n bases the morphoge nesis of the organ on the u n iver

sal adaptability of the primitive promammalian e n todermal
l ung sac to respo n d to the fun ctio n al deman ds made on i t This
-
.

i nhere n t pote n cy derived from the mammalian an cestry to de

v el o p bro n chial b u ds from a ny poi n ts of its epithelial s u rface ,

a n d to assemble them i n to a patter n adapted to the respiratory

 P ULMONARY EV O LUTI O N IN M AM MAL I A 1 95

req ui reme n ts of the organ ism lies at the root of the bro n chial
,

architect ure of the extan t mammalia In place of mechan ical

.

modificatio n of a preformed str u ct ure we see everywhere i n the

mamm alian bro n chial tree the selection of developme n tal paths
from a fi el d o n ly limited by the exte n t of the p ulmo n ary e n toderm
itself Selectio n of the n umber locatio n an d directio n of the
.
,

poi n ts at which epithelial prol i feration is foc u sed i n bro n chial

developmen t furn ishes the archeal types from which the extan t
pattern s have passed to the modern desce n dan ts For any .

give n mammalian grou p the force of i nh eritan ce will ordin arily

assemble the e n todermal b u ds progressively as they develop i n
the embryo accordin g to a pattern which o u tli n es the type of
,

bro n chial tree characteristic for the form i n q u estio n i n the n or

mal ad ul t i n dividu al.

Comparative an atomy i n dicates clearly that the forces which

determi n ed the selectio n of this pattern i n the a n cestor were en
v i r o n m en t al an d metabolic i n character A primeval an cestral
.

bron chial type tran smitted to the descen dants can be altered an d
modified by the same morphoge n etic processes determin i ng the
selectio n of n ew bro n chial lin es as soo n as the desce n dants
,

themselves or a large proportion of them have begu n un der

, , ,

chan ged co n dition s of en viro n me n t an d adaptatio n to co n stitute ,

n ew mammalian types req u iri n g s u ch respiratory m o di fi c at i on s .

A n example of this is fu rn ished i n the rode n t s u bfamily of the

Hydromyi n ae : Xeromys a p urely terrestrial form exh ibits the
, ,

domin an t asymmetrical bron chial type common to most mem ,

bers of the order with the eparterial developmen t con fin ed to

,

the right side Its close relative H ydromys is strictly aqu atic
.
, ,

i n habit an d develops a b i lateral symm etrical eparterial district

,
.

The same co n ditio n is e n co untered i n M yop otamus i n which a ,

very exte n sive left eparterial bron ch u s appears i n association

with the equ ally well developed left cardiac bro n ch u s .

These i n stan ces of left eparterial bro n chial exte n sion i n some
of the aqu atic roden ts place themselves i n lin e with the corre
sp o n di n g p u lmo n ary orga n izatio n fo u n d i n its f u ll developme n t

i n the pi n ni pedia an d some cetacea n s Li ke these they are

.

examples of p ulmon ary adaptatio n s to sp eci fi c e n viro nme n tal

an d f un ctio n al co n ditio n s.
1 96 G EO . s . HUNTINGT O N

Perhaps the most si g n i fican t example of the respo n se of pul

m on ary architect u ral orga n izatio n to aq u atic adaptatio n is
a fforded by the l un g of H i pp op otamus ltberi ensi s which i n c on ,

trast to all the other typical artiodactyls with right eparterial

bro n ch u s derived from the trachea possesses a bilaterally sym ,

metrical tree each st emb ron chu s furn ishi n g an eparterial com
,

po n e nt .

We have previo usly referred (p 1 2 8 ) to l un gs i n which i n the.

phyloge n etic evol utio n this selective m o difi cat i on of bro n chial
stru ct ure has n ot bee n active becau se the desce n dan ts of a more
,

primitive l un g type i n their adaptatio n to an altered e n viro n

-
,

me n t chan ge o n ly those parts of their organ izatio n which are

,

directly affected by the n ew e nviro n me n tal co n ditio n s If the .

l un g is n ot o n e of the parts thu s acted u po n the respiratory ,

apparat u s is n ot called upo n to modify its stru ct ure The pul .

m on ary organ izatio n both i n the archeal evol u tio n ary stage an d
,

i n the later adaptatio n s to n ew co n ditio n s was an d remai n s , , ,

adeq u ate for the req uired respiratory exchan ge We fo un d .

(p 1 3 0) i n this balan ce betwee n l un g stru ct ure an d specific en

.
—

v i ronm en t al factors the expla n atio n of the appare n t discrepa n cy

of the bro n chial tree of Tax i dea compared with that of the
remaini n g M u stelidae of the Hystricomorphs as agai n st the rest
,

of the rode n t order an d of the atypical forms amon g the cetacea

, ,

if s u ch really exist .

The factors which on the physiological i n terpretatio n of the

,

comparative an atomical evide n ce may be regarded as determi ni n g,

sp ecifi c all y p u lmo n ary evol u tio n ran ge themselves un der the ,

followi n g headi n gs :
1 In its broadest li n es the vertebrate l un g emphasizes i n its
.

str u ct ure the effects of the e n ormo u s metabolic chan ge which

the i n crease of oxyge n ation has e n tailed i n passi n g from the poi
kil o t h erm al amphibia n s an d reptiles to the homoeothermal birds
an d mammals This fi nd s its morphological expressio n i n the
.

n eomorph developme n t of the cra n io ve n tral p u lmo n ary districts -

of the two higher classes partic ularly i n the acqu isitio n of the
,

eparterial system O n the other han d the higher l ung type of the
.
-

more adva n ced reptilia n forms depe nds largely upo n the i n creased
 1 98 G EO . s . HUNTINGT O N

here a seco n d factor presen tly to be con sidered is chi efly

respo n sible .

The greatly i n creased eparterial developmen t of the typical

avian l un g depe n ds on the high rate of tiss u e comb u stio n as -
,

eviden ced by the high bodily temperat ure an d on the e n or ,

mo u s relative developme n t of the specialized pectoral muscul a

t ure with its great and lon g con tinued activity d uri n g fli ght In .

S pite of the fact that the weight an d b u lk of the whole body has

been redu ced as far as possible by special adaptatio n s to this

p urpose ce n tred chi efly i n the appen dages of the respiratory
,

tract the remaini n g factor is correlated with a great cephalo

,

ven tral exte n sio n of the l un g an d the establishmen t of the type

of p ulmon ary organ izatio n characteristic for the class .

.3 Chan ges i n the ge n eral e n viro nm e n t which are associated

,
:

with chan ges i n the stru ctu re of the lun g may lead to the replace
,

men t of a slow regular respiratory rhythm by a type in which

periods of exceedi n gly active respiratio n altern ate with more or
less prolon ged in termissio n s d urin g which respiratory exchan ge is
at a stan d still This co n dition is the result of the adaptatio n
.

of the mammali an organ ism i n a greater or lesser degree to the

aq u atic habitat Shorter or lo n ger periods of s ubmerge n ce
.
,

durin g which respiratio n is s u spen ded are su cceeded by i n ter

,

vals of active su rface breathi n g The reactio n of thi s e n viron

.

men tal factor on in trap ulmon ary organization is see n i n the

hi gh degree of bilateral eparterial developmen t i n the pinn ipede
carn ivores sire ni a an d typical cetacea
,
.

The qu estion of the total expe n dit ure of respiratory e n ergy

withi n a give n period of time becomes importan t i n its effects
o n p u lmo n ary str u ct u ral m o difi cat i on s of evol u tio n ary character .

In the mammal the i n terdepe n de n ce of the biochemical factors

of r espiratio n an d l un g str u ct ure requ ire an alysis prim arily i n re
-

spect to the rapi di ty with which the gas exchan ge is e ffected -

.

The more exte n sively organ ized mammalian l un gs i n dicate either

a high degree of tiss ue comb ustio n at a fairly constan t rate ( Un g u
—

late type ) or a very rapid i n termi tten t metabolism called for at

,

definite periods with i n tervals d uri n g which respiratio n is sus

pen ded ( aqu atic adaptatio n s) .
 P ULMONARY EV O LUTI O N IN MAMMAL I A 1 99

In the latter type all the accessory m odi fi cati on s are i n the
directio n of stori n g 0 0 un til the same can be exchan ged i n large
2

qu an tities an d rapidly duri n g the resumptio n of respiratio n

( Caval s i nu ses of P hoca an d M an atus abdomi n al ve n o u s plex
,

u ses of M acr orht nu s) The l un g therefore mu st be so organized

’

as to meet this i n termitte n t deman d for high efficien cy altho u gh ,

n ot called u po n i n the i n tervals of respiratory s u spe n sio n The .

l un g acqui res the morphological complexity correspo n di n g to

the highest degree of effi ci en cy which c an be deman ded i n any
give n form un der the n ormal e n viron men tal co n ditio n s .

C O N C LUSI O N

The search for the el u sive hypothetical promammalian

groun d plan of bron chial architect ur e has termi n ated somewhat
-

like the hun t for the Philosopher s Sto n e ’

.

The mechan istic con cept of a defin i t e an d crystallized archeal

bro n chial tree from which all extan t mammalian types are de
,

rived by m odifi c at i on s of the pattern throu gh migratio n doe s ,

n ot exist i n the commo n ly predicated form of a co n crete an d

fi x ed morphological e n tity It had n o reality i n any definite

.

an cestral stru ct u re save i n the sen se that the primitive reptilian

,

l un g an d its phyletic derivative the mammalian e n todermal pul

,

m on ary anl age both retai n the pote n cy of developme n t by selec

,

tion in to the type deman ded by the en viron me n t In place of .

the myt hical commo n an cestral bro n chial tree appears a livi n g
plastic organ ization respo n sive to the chan gin g deman ds of
,

biological evolu tion an d replete with an swers to the m odern

probl em s of morphoge n etic in qu iry .
2 00 GE O . S . HUNTINGT O N
BIBLI O G RAPH Y
1 A E BY ,
CH R . 1878 D ie G t lt d B
es a hi lb es r o nc a au m e s un d d ie H om o lo gi e

d er L g un en la pp en b e i m M en sc h en . Cen t r a l b l d m ed W i s sen sc h . . . .
,

Nr 16 . .

1 88 0 B D er ro n c h a l i b au m d e r S a u g e t h i e re un d des M e n sc h e n ne b st

Bm k g e er un en u b er d en B r on c h a l i b au m der Vog el un d R p t i li e en .

L i pz i g
e .

BL I S N I A N S K A J A , G R U NI A
1 90 4 Z u r n t w c l un s esc h c h t e d e r m en s c h E i k gg i
l c h en iu n en : L g
r o n c h a l a u m , L un g en f o r m ss Zur c h B i b . Di . i .

BO E CK H , R UD O L F 1 9 1 4 D i e n t w c l un d e r S au g e r lu n g e In A l e sc h E i k g . . F i
m ann , M o rp h o g enet i sc h e t u en I M amm a l a G e g en b au r s M o r h
’
S di . . i . p .

J ah r b .
,
48 . Bd .

BR E ME R JOH N ,
L . 1 904 O n t he l u n g of t he o p o ssu m . Am . J ou r An at . .
,

vo l iii .
, pp
67 7 3 .
—
.

C HI A R I ,
H . 1 889 e er Ub i
e n e n n e u en T yp u s von M i ssbi l d un g an d e r T ra c hea

d es Men sc hen . Z i e g ler ’

s B it g
e p ra e z . at h . A n at . u . a ll g e m . P at h o l .

5 Bd. .

1 890 Ub e e r e i n c o n g en i t a l e s D iv ti k er el d es re c h t en St a mm b ro n c h u s .

P g ra m e d W o c h en sc h r , N r 46
er . . . .

1 890 e b e r d as U
o r o mm en e n e s o V k i d pp e l t en S e i t en b r o n c h u s an d em
r e c h t en t amm r on c h u s d e s Men sc h enS b . Z e i t sc h r . f . H ilk d e un e, Bd .

10 , s . 47 04 7 8 .

1 89 1 Ub e er e n e n eu e i F o rm v o n
‘
D re i t h e i l u n g d e r T ra c h ea
’
b ei i
e n em

16 T g a en a l t en Kn b a en mit so n st ig en Bi l dun g s - A n o m a l en , i d a ru n t e r

auc h M an g el d er M i l z un d V er la g e ru n g d es Lig am en t u m h ep at o
d uo d en a l e P g md
. ra er e . W o c hen sc h r .
,
Nr 8 . .

D ALL A R OS A 1 88 9 B i t g e ra e z ur K u i t i k un d M p h l g i
as s or o o e d e r V a r i et at en
d es m en s c hl i h B c en ro n ch i l b um a Wi n k l in
a es . e er . W o c h ensc hr , .

J ah r g 2 , N r s 22 , 2 3 , 2 4 . . .

D H A R D I V I L L E R L a r am i fi c a t i o n
’
b ron c h i qu e c he z le l a pi n . Bi b l i o g r . An at .

S pt e .
—
O ct ob r . 1 8 96, J an v .
—
Fév r 1 8 9 7 . .

1 896 Dév l pp m e o e en t de l a ra m i fi c at i o n b ron c h i qu e et b ron c h es ép ar

t é i ll h z l
r e M es c e es a m m i f ere s . So c d . . la Bi o lo gi c, 19 Dé mb ce re .

1 89 7 H m l g ti o o o a on d es b r o n c he s c he l e l a z pi n . Bi b l i o g A t r . na . J an v .

—
F év r .

1 8 97 Les b r on c h e s ép a r t ér i e ll es c he z le s M am m i f ere s e t spe c i a l e men t

c he z l h o mm e
’
. Ex t r . d . Co mpt es r en du s d . l A c ad
’
. de s Se , 2 o ut
. A .

1 897 b i du m u t n E t
Or i g in p e de s b r o n c he s lo a re s o o . x r . d C om t e s
.

r en d d é n d l S o c 2 0 N ov
. . s a c es e a .
,
.

1 8 97 Dév l o pp m en t d b n h p in i p l h z l m u t on E t
e e es ro c es r c a es c e e o . x r .

d C mpt . n du d é n d l S o c d B i l 4 Dé m b
o e s re s . s a ces e a . . o .
,
ce re .

1 8 9 7 Dév l pp m n t d b on h h z l m u t n E t d C m p t
e o e e es r c es c e e o o . x r . . o es

re n du d é n e d l S o c d B i lo g i 1 1 Dé mb
s es s a c s . a . e o e, ce re .

1 897 Dév l pp m n t t h m l g at i n d b n h p in i p le c he z
e o e e e o o o o es ro c es r c a s

l s M mm i f e
e ( L p in ) The N n y
a re s a . se , a c .
R esumen por los autores Baumgartn er M F N elsen y W ,
E A
. .
,
. . .

D ock
Hospital Halstead Halstead Kan sas y Sai n t L ou 1 s Missou ri
, , , , ,

El desarrollo de las g l an dul as u teri n as en la especie human a .

En feto human o de seis a siete meses de edad 1a cavidad

el

uteri n a presen ta bordes irregularme n te a serrado s a expe n sas de

los c uales se desarrollan las glan dul as uterin as Ad em as existen .

proyeccio n es tran sversas de forma semil un ar en las sup erfi ci es

an terior y posterior del utero estan do formadas estas proyec ,

ci on e s por d i v ert i cul o s de la cavidad uteri n a E stas proyeccio n es .

de la cavidad u terin a llevan tambié n glan dul as L as g lan dul as .

mas tempran as aparece n en forma de crecimien tos redon deados ,

q u e existe n ya en este periodo T o d o d los ej emplares de edad '

mas avan zada examin ados por los au tores presen taban glandul as
mas o men os desarrolladas L as gl andulas son primeramen te .

redon deadas dividién dose después en ramas que p ueden exten

,

derse paralelamen te a la su perficie en cierto trayecto para divi d

irse n uevamen te al mismo tiempo que se hun den en la pared
u teri n a estan do f rec uen t am en t e colocadas estas n uevas ramas
,

en un plan o perpe n dic ular al plan o de las divisio n es precede n tes .

Tales subdivision es p uede n te n er l u gar varias veces hasta que

se forma un a gl an dula completame nte ramificada En los ad ul .

tos los extremos distales de las gl an dul as se extien den p aral el

amen te a la superfi ci e del epitelio proyect an do se gen eralm en te ,

dichas gl an dul a s en la misma di rec ci on S on frec ue n tes las an as .

tomosis e n tre diferen tes gl an dulas .

T r a ns l a t i
by J on o se F N o m d ez
C ar ne gi I
e t i t ut i
ns o n of W hi
as ng t o n
 BY T H E B BI LI O GR A H I C P S ER V ICE AP RI L
, 19

D E V E L O PM EN T O F TH E U T E R I N E G L AN D S IN MAN
E . A . B A UM G A R T N E R M ,
. T . N EL S O N , AND WM . D O CK
H a l s tea d , Kan s a s , a nd S t L ou i s ,
. Mi ssou ri

S E V EN FI G UR ES

Inhis book A L aboratory Textbook of E mbryology

,

Min ot i n speaki n g of a model of the hum an u teri n e g lan ds

“
,
,

said : The model demo n strates that the c on cep

tion of the character of the uterin e glan ds which has hitherto ,

prevailed is very i n adequ ate

,
D escriptio n s of the morpholo gy .

of the u teri n e glan ds i n the vario u s histological textbooks vary

slightly Piersol . describes them as tu b u lar or slightly
bifu rcated wavy i n vagin atio n s with tortu o u s bli n d e n ds dis
, ,

trib u ted at fairly regular i n tervals L ewis descriptio n based .

’
,

o n H edbl om s model states that they are bran ched tortu o u s

’
,

glan ds which occasion ally a n astomose and which have i n their ,

deeper portio n s lo n g horizo n tal bran ches at right an gles to the

mai n tube .

In the older texts ( Stricker 7 3 ) the gla n ds are described as

’
,

simple althou gh occasio n ally they give off bra n ched tubes from
,

the ce n ter or j ust below the ce n ter They are twisted or cork .

screw like and i n the fun dus may run horizo n tally to the s urface
-
, .

The o n ly models of these gla n ds i n any stage k n ow n to u s , ,

are those of H ebl om s on e of which is fi g ured i n Mi n ot s L ab o

’
,
’

ra t o ry Textbook of E mbryology The model fi g ured is mu ch .

more complex than the descriptio n s of the tex t books wo u ld

,
-

in dicate From sectio n s H i t schm an an d A dler

. have
described the character of the glan ds d u ri n g the vario u s stages
of the men stru al cycle b u t they give very little i n formatio n ,

other than is fo un d i n the u s u al texts an d n o addition al i nf or

matio n l n regard to the glan ds d uri n g the i n terval stages Wax .

models of the gla n ds i n the vario u s stages of the cycle sho uld
2 03
2 04 E . A . B AUMGART N E R ,
M . T . NE L SON ,
A ND WM . D O CK

show some very i nteresti n g res u lts an d the pla n was to i n cor ,

p o rat e these res u lts i n this paper However lack of material .

,

preve n ted this at this time .

There is an eve n greater lack of descriptio n of the characteristics

of the glan ds du ri n g developme n t L ewis states that the glan ds .

develop at the bases of folds In Kei b el and Mall s E mbryology

.
’

very little atte n tio n is give n to their developme n t an d o n ly

o n e i n vestigator (W yder 7 8 ) is q u oted This i nvestigator
’
.
,

states that t he glan ds develop i n depe n de n tly of the age of the

i n divid u al .

It seems therefore from this brief review of the literat u re

, ,

that very little is k n ow n of the developm ent of the u teri n e

glan ds or of their ultimate form For that reaso n the se n ior .
,

author su ggested that this problem sho u ld yield resu lts worthy of
p u blicatio n The models were all co n stru cted by the j un ior
.

i nvestigators .

DEVEL O PM E N T O F TH E G L A ND S

The material u p on whi ch this stu dy is based co n sists of serial

sectio n s of portio n s of the corpora of u teri of fet u ses and of
adu lts The yo un gest u teru s st u died was of a six to seven
.

mo n th fet u s the oldest a u teru s of a twe n ty fiv e year old Virgin

,
- — -
.

The e n tire fun d u s of the six to seve n mo n th fet u s was sectio n ed —

a n d a model made of the l u me n of the orga n i n cl udi n g the ,

epithelial li n i n g and any ou tp ou chi ng s or gla nds This model is .

show n i n fi g u re 1 A n i nteresti n g partic u lar of this stage is

.

the pec uliar S shaped l ume n with irreg ular sides There are
-
.

folds of the uteri n e mu cosa from which small o ut p ou chi n g s

protru de .

The irregular folds of the epitheliu m on the sides of the lu me n

give the effect of a scalloped edge R u dime n ts of glan ds are .

prese n t The most promi n e n t epithelial folds are fo un d on the

.

an terior an d posterior s urfaces an d are directed toward the

cervix O n ly t wo are shown i n the model These tw o run
. .

almost tra n sversely across the s urface of the l ume n and have
the appearan ce of cresce n t shaped hood like evagi n atio n s -
,
-
2 06 E . A . B AUMGAR T N E R , M . T . NE L SON ,
AND WM . D O CK

t u b u lar en d pieces exte n di n g obliqu ely i nto the m u cosa ( fig .

O n e glan d shows the two t u b u lar bra n ches r unn i n g parallel to

the su rface of the l ume n as described above with on e en d piece
, ,

t urn i n g toward the mu sc ular layer almost at right an gles an d ,

se n di n g o ut two bran ches on e from each side the n dividi n g , ,

i n to two parallel t u b ular end pieces .

A n other glan d is very m u ch more complex A fter the for .

matio n of the T shape on e of the arms of the T t u rn s sharply

-
,

and divides formi n g a Y shape O n e of the arms of the Y

,
-
.

divides agai n whereas the other arm which is larger gives off
, , ,

three pairs of bra n ches and termi n ates i n a cl u ster of fiv e bran ches .

The bran chi n g is dec u ssate O i the termin al cl u ster o n e short .

ce n tral termi nu s appears to be the co n ti nu atio n of the main

stalk The termin atio n s u s u ally poi n t toward the mu scle layer
.

an d show e n largeme n ts .

O wi n g to a lack of material we have bee n u n able to model ,

or describe the stages betwee n a fo urtee n year old before p u berty — -

an d the twe n ty fi v e year old n u llipara of the i n terval stage

— - -
.

O f the latter an area of attachme n t of abo u t twe n ty fi v e gla n ds

,
-
,

abo u t 1 mm by mm was modeled whi ch area i n clu des o n e

. .
, ,

hun dred and fi fty sectio n s 1 0 i n thick n ess Some of the glan ds u
,
.

are gro uped fo ur glan ds comi n g from on e slight furrow i n the

,

mu cosa ( fig The others are all more or less isolated bu t i n

.
,

qu ite d efini t e rows In st u dyi n g the portio n s of the glands

.

which lie n ext to the m u sc ulat ure on e is impressed with the ,

fact that the gla n ds give the appearan ce of r unn i n g parallel to

the s u rface of the l ume n and all i n the same directio n an d that
, ,

the great prepo ndera n ce of glan dular tiss u e is i n the o u ter third
or half of the e n dometri um The co n stricted n ecks of the glan ds .

are short the stalks grad u ally e n largi n g as they exte n d obliqu ely
,

toward the mu sc u lature Some glan ds show co n stricted areas

.

at i ntervals others prese n t a slightly spiral appeara n ce but

, ,

o n ly few show any bran ches u n til they are deep i nto the m u cosa ,

whe n they all t u rn more or less grad u ally i n to a directio n at

,

right a n gles to the o n e which they have j u st bee n followi n g

a co u rse n early parallel with the s u rface of the l ume n .
 D EVELO PM E NT OF UTE R IN E GLAND S 2 07

The mode of termi n atio n varies a great deal O n e gla n d .

forms the familiar T shape the cross bar bei n g t ub u lar a n d

-
,
-

short A n other sen ds off a lateral bra n ch which immediately

.

divides di chotomo u sly each bran ch agai n dividi n g on e of the

, ,

l atter bra n ches exte n di n g back two thirds of the distan ce toward -

the epithelial s u rface termi n ati n g i n an e n larged en d A third

,
.

gla nd aft er t u rn i n g at right a n gles to the s urface breaks up

, ,

i n to two parallel t ubes on e s u bdividi n g the other an astomosin g

, ,

with a parallel en d piece of an other gla n d .

Some are eve n more complex i n their bran chin gs an d divisio n s

( g )
fi . 7 approachi n g the complex type described for the fo u rtee n

year old except that the bran ches are fo un d more deeply em
-
,

bedded i n the mu cosa and i n ge n eral run parallel to the s u rface

at their termin ation s O n ly o n e si n gle tu b u lar glan d is prese n t
.

a n d it r u n s parallel to the s u rface at its distal en d .

The models show that a n astomosin g be twee n glan ds an d

between bra n ches of glan ds is n ot un comm o n Two simple .

gla n ds an astomose the n divide i n to several bran ches (fi g .

A gai n two rather widely separated glan ds a n astomose then

, ,

divide i n to a complex set of bran ches which exte n d for some

dista n ce alo n g the m u scle layer .

DIS C USSI O N
G u yo n de Si n éty
an d fo un d n o gla n ds properly ,

s peaki n g i n the corp u s of the fetal u ter u s altho u gh they fo u n d

, ,

well developed o n es i n the cervix D e Si nét y states that gla n ds

-
.

are n ot developed un til the sixth or seve n th year W yd er .

who is q u oted i n Kei b el an d Mall s E mbryology fo un d ’

,

n o gla n ds i n the u teri of t en year old girls an d co n te n ds that

— -
,

the stages of developme n t bear n o relatio n to age T ou rn eux .

a n d L egay fo un d glan ds i n the cervix of a late fetu s b ut ,

n o n e i n the corp u s eve n at birth M 6ri cki

,
described .

glan ds i n the cervix of fetal u teri In a premat u re of n i n e .

mo n ths he fo un d well developed glan ds W ith bra n ches i n the

,
-
, ,

corp u s an d i n a n ew born he fo un d many gla n ds i n the f un d u s

- .

The glan ds varied i n form b u t were u s u ally t ub u lar with relative

,
2 08 E . A . B AUMGART N E R ,
M . T . NE L SON ,
A ND WM . D O CK

large l umi n a In ad ults the glands forked n ear their e n ds

. .

N agel fo un d that the gla n ds of the corp u s developed mu ch

later tha n those of the cervix altho u gh he fi g u red gla n ds i n the ,

corp u s of a 1 7 c m fetu s W yder appears to have bee n abo u t

-
. .

the o nly on e w h o did n o t fi n d gla n ds i n the corp u s at birth an d

is the o n ly o ne w h o has stated that the developme n t of the gla n d s
is e n tirely i n depe n de n t of age .

The pec u liarly shaped lu me n of the u teru s of the fetu ses and
yo un g ad ults has bee n n oted by other i n vestigators an d is
appare n tly co n sta n tly presen t N o explan atio n of this pec uli .

ari t y has bee n o ffered The theory that it may be du e to the

.

u n io n of the early M ii ll eri a n d u cts is worthy of co n sideratio n .

In all of the specime n s u p o n which this st u dy is based glan ds ,

or r u dime n ts of gla n ds are fo u n d from the six mo n th fetu s ,

—

o nward In the six mo n th fetu s they are prese n t as smal l

.
—

o u t p o u chi n g s from the folds of epitheli u m Co n trary to t h e .

fi nd i n g s of M 6ri ck e o u r specime n of n ew bor n shows gla n ds

,
—
,

ofte n closely crowded witho u t bran ches an d n ot as yet t ubu lar

, ,

b u t rather havi n g the appearan ce of simple o u tgrowths with

e nlarged fl at t en ed en d pieces and slightly co n stricted stalks .

This shape is typical of early stages of u teri n e glan ds as well as

of the early stages i n the developme n t of man y other glan ds .

The t ub ular shape and T shaped bran ches appear i n the first -

year however
, .

The glan ds i n crease i n le n gth and i n n u mber of bra n che s

d uri n g the early years of life althou gh the growth is n o t extreme , ,

an d the stages from o n e to seve n years show n o marked di ffer

en t i at i o n The T shaped bran ches an d the growth from t h e

.
-

e nds of the folds of the mu cosa are co n stan t characteristics .

J u st before puberty gla n d ular growth seems markedly

,

haste n ed altho ugh the muc osa may n ot be m u ch thicker O i

,
.

the glan ds modeled man y have T sh ap ed bran chi n gs Further — -

.

bra n ch in gs an d s ubdivisio n s beyo n d the T shape are fi rst ,

-
,

observed at this time Some of the glan ds are closely crowded

.

as i n yo un ger stages The t ub ular shape is co n sta n t altho u gh

.
,

e n larged e n ds an d irreg ular e n largeme nts of the stalk are fo un d .

The short co n stricted n ecks are here prese nt

,
.
210 E . A . B AUMGART N E R ,
M . T . NE L SON ,
AND WM . D O CK

of an eight mo n th child a t en year child an eightee n year

-
,
-
,
-

n u llipara an d of an eighty two year old m u ltiparo u s woma n

,
— — -
.

The yo u n gest shows irreg u lar folds with small o ut p o ck et i n g s .

The folds w e believe are mu cosal folds s u ch as we have fo u n d

,

i n o ur specime n s The glan d u lar r u dime n ts are very small i n

.

compariso n with those of o ur specime n of a day old an d really ,

appear i n shape mu ch more like the gla n ds of o ur six mo nth -

fet u s The t en year old specime n shows small irregu lar short
.
— -
, ,

tu b u lar o u tgrowths with some an astomoses an d irreg ular folds .

Here the type does n o t correspo n d at all to any of o ur specimen s .

The bran chi n gs are few an d dissimilar to those foun d by u s .

The short sle n der t u bu lar glan ds here also appear more like
small o u tgrowths fo un d i n ou r six mo n th fet u s -
.

The eightee n year old specime n fi t s well i n to o ur series betwee n

— -

the fo urteen an d the twe n ty five year old stages The short - — —
.

n arrow stalk an d T like divisio n s which w e describe are n ot —

fou n d b ut i n stead a Y like bran chi ng from a lo n ger stalk O n e

,
-
.

arm of the Y has o n e or two Y like s ubdivisio n s whereas the -

,

other arm se n ds off ma ny irreg ular si n gle bran ches which fu rther
s ubdivide Several bran ches exte n d for some distan ce toward
.

the mu sc ular coat and o n e gives o ff a bran ch which retu rn s

,

o n e third
-
of the dista n ce to the u teri n e l ume n the n tur n s ,

sharply and ret u r n s i n its origi n al directio n Ma ny termi n al .

bran ches have e n larged e nds similar to those w e have fo un d .

An astomoses wi th other gla n ds are prese n t .

The eighty two year old specime n is a qu ite remarkable o n e

— — -
.

Ma n y cystic t ub ules are fo un d at the e n ds of short n arrow ,

t u b ules A na stomoses betwee n these cysts by mea n s of small

.

t u b u les are fo u n d The cystic e nlargeme n ts are elo n gated oval

.

str uct ures and are fo un d close ly crowded an d n ear the s urface
epithelium A s is well k n ow n the mu cosa is thi n an d the e n tire
.
,

depth of the gla n d is o n ly abo u t on e third that of o u r adu lt —

specime n an d eve n less than that of the eightee n year old model — -

of H edb l om .

The work of Scammo n rep orted at the meeti n g of the A merican

A ssociatio n of A n atomists shows that some i nterestin g
phases may have b ee n overloo ke d It is extremely imp orta nt .
 D EV E LO P M E NT OF U T E R IN E GLAN D S 211

an d wou ld be val u able to have models of the vario u s stages

from birth to three mo n ths of age i n stead of o nly at birth , .

Th en correspo n di n g to the periods of growth as i n dicated by

, ,

le ng ths of the u ter u s models of the sixth year an d agai n abo u t

,

the eleve n th wo u ld be desirable The models of seve n mo n ths .

a n d of fo u r years as i n dicated are q u ite similar A specimen .

from a seve n year old child shows rather p oorly developed

— -

gla nds certai nly n o t m u ch beyo n d the fo u r year stage s wh i l e

— - —

a model of a twe n ty on e mo n th child is very similar to the o n e

- -

of seve n mo n ths .

It can n ot be den ied therefore that it wo uld be i nteresti n g to

, ,

reco n stru ct more of the stages of the glan ds betwee n fo u r and

fo urtee n years as well as d uri n g the me n stru al period the me n o ,

pau se an d of a paro u s woman and that these stages sho uld be

, ,

i n cl uded i n this paper Un fort un ately we have n o t fo u n d the

.
,

material available .

C O N C LUSI O NS

1 . G lan ds
are fo u n d i n the corp u s of the u teru s i n a six to
seve n mo nth fet u s an d i n all material st u died beyo n d this age
-
.

2 The earliest gla n ds are small irreg u lar o u t p o u chi n g s fro m

.

semilun ar mu cosal folds later developi n g co n stricted n ecks an d

,

e n larged en d pieces similar to ma n y other gla n d ru dime n ts .

3 The n ecks persist eve n to the ad u lt stage the e n larged

.
,

e n ds becomin g t ub ular an d dividi n g T like with sometimes a -

seco n d lo n gitu din al divisio n of the en d bra n ches .

4 The stalks follow an obliq u e co u rse i n the ad u lt sometimes

.
,

almost a S piral o n e .

5 N ear the m u scle layer the bra n ches r u n parallel to the

.

s u rface an d all i n o n e directio n sometimes formi n g a n etwork by

,

an astomoses of di ffere n t bra n ches .

6 The greater part of the gla n du lar tiss u e lies i n the lower o n e
.

third or fo u rth of the e n dometri u m .

7 A d u lt u teri n e gla n ds are compo un d a n astomosi n g t u b u lar

.
, ,

g la n ds .
212 E . A . B AUMGART N E R ,
M . T . NEL S ON ,
A ND WM . D O CK '

LITERATURE C ITED
C A DI AT , O . 1 88 4 M em o i re
’

su r l
’
u t e ru s e t l e s t ro m p es . Jo u r d e .
’
1 An a t . et de
la Ph y s ,
T 37 . .

GU YO N, H . 1 8 58 Et u d e su r l es vité d l
ca s e
’
u t erus al e ta t de v a cu i té . P i ar s

(Q uo t ed f ro m T o u r n eu x t L g y)
e e a .

H I TS C H M A N ,
F .
,
UND ADLER ,
L . 1 90 7 D i L h'

e e r e vo n d er E d
n o m et r iti s . Zei t sch .

f . G e b u r t sh . u . G yn .
,
Bd 60 . .

L WI
E S ,
F T . .
,
A ND S T OH R ,
PH . 1914 A t xtb k f h i t l g y B l k i t Ph i l
e oo o s o o . a s on , a .

MI NO T C S ,
. . 1 9 10 Al b a o ra t o r y t xt b k f m b y l g y P Bl k t
e oo S o e r o o . . ac s on s
’
on

Ph i l a .

M O R I CK I R ,
. 1 88 2 D i e U t e ru s s c h l e i m h a u t i n d en e r sc h e en en l t e r s e r v i d A p i d o en

un d z u r i
Z e t d e r M en s t r u a t o n r c h 1 G e b u r t sh u n d G yn i . A . . . . .

N A GE L W ,
. 1 89 1 Ub e er di e E nt w c i k lun g d es Ut e ru s un d d er V gi a na b ime

M en c h en . A r ch . f m i kr
. . A n at .
,
Bd 3 7 . .

SC A MM O N ,
R . E . 1 919 S t di u es i n t he g ro w t h o f t he u t eru s . A nat . R ec .

DE S I N EJT Y
’

1 879 T i t é d gy
ra e n éc a l o g i e .

TO U R N E U X F .
,
ET LE G A Y ,
OH . 1 88 4 M é mo i re su r le d e v e l O pp em en t de
1 u t e ru s e t du v gi a n . Jou r . l An a t
’
. et d e la Ph y s .
,
T 37. .

W Y D E R , A T 1 878 .e r e z u r n o rm a l en
. B i t ag un d p th l gi
a o o s c h en Hi st ol o gi e d er
men s c h l i c h en U t e r u ss c h l e i m h au t . A r ch . 1 G yn a k
. .
,
Bd 1 3 . .
 R es ume n por los au tores ,
Eliot R . Clark y Elea n or L i n to n Clark .

La r ea c c de las cél ulas de la cola d el re n ac u aj o hacia la

mn
i nye c c i on de aceite de croto n en el a n i mal Vivo , .

L os au tores i n yectado g l ob ul o s m i c ro sc é p i co s de aceite

h an
de croto n dil u ido en aceite de p a rafin a en la ex p an si én caudal de
“
,

larvas de Hyla observan do en el an imal vivo la i n flam a ci on

” ,

aséptica res ultan te El p un to est u diado especialme n te es la .

c u e st i é n de s i d u ra n te la i n fl am a c i on las cél ulas de tipo d efi n i d o

'

, ,

perman ece n como eleme n tos esp eci fi c o s o si se tran sforman en

otros tipos cel ulares Primeramen te se obt uvo u n a idea ge n eral .

de los procesos qu e tien en l u gar est u di an d o se desp ués c uidado ,

sam en t e los difere n tes tipos de cél u las Se h a observado qu e .

los leu c o ci t o s y cél ulas emigran tes del tej ido cami n an hacia cl
aceite de croto n pero n o llegan a po n erse en co n tacto c on él
, ,

perman ecien do a un a corta distan cia y hacié n dose estacio n arios

al mismo tiempo q u e prod u ce n en s u su p erfi c i e apé n dices p un tia

g udos q ue les asemej an a las cél ulas pequ e nas del tej ido c on j un
ti y o Si n embargo n o se tran sforman en cél ulas co n j un ti vas
.
, .

Por el co n trario mas tarde se hacen esféricos y desp u és de varias

,

horas o aun dias v uelven prese n tar su actividad am i b oi de y

,

emigran habien do sido eli mi n ado el aceite d uran te este tiempo

, .

L as cél ulas co n j un tivas cercan as a las gotas de aceite se t ran sfo r

m an en eleme n tos gran u lares se hi n chan y lle n an de v a cu ol a s y ,

algun as de ellas mu eren L a mayor parte de ellas san an des .

p u és de la elim i n a ci é n del aceite y vu elve n a adqu irir la fac ultad

de moverse L os capilares san g u in eo s prese n tan v a cuoli z aci é n
.

en d o t eli al y r et ra c ci é n a las qu e sigu e nu evo crecimie n to El,

.

e n dotelio li n fatico se comporta de un modo semej an te Todos

'

los tipos cel ulares perman ece n esp ec i fi c o s L as c éullas emi .

g ran t e s n o se forman a expe n sas sel e n dotelio o de las cél ulas

co n j un tivas n i tampoco origin an n i n gun o de estos tej idos L as
,
.

cél ulas co n j un tivas y los en d ot el i o s san gu i n eo y li n fatico tambié n

se co n servan esp ec i fi c o s y n i se co n vierte n en otros tipos cel ulares
,

n i se deriva n de ellos .

T r a n sl a t i on by J é Fos N o m dez
C ar n e gi e I ns t 1 t u t 1 o n of h
Was i n g t o n
2 22 E LI O T R . CLARK AND E L E ANOR LINT O N CLARK

co nn ective tiss u e cells are fewer i n n umber than i n the larvae of

-

frogs an d toads They are therefore wo n derfully tran spare n t

.

a n d the i n divid u al cells may be followed readily A s was .

previo u sly me n tio n ed (E R Clark this form is m u ch easier

. .
,

to a n aesthetize tha n the larger larvae of the Anu ra it respo n ds ,

more q u ickly to chl oret on e an d remain s motio n less for lo n ger

periods witho u t deleterio u s e ffects o n the circ ulatio n .

The irritati n g s ubsta n ce selected for these experime n ts w as

croto n oil The un dil uted oil was i n j ected i n a few larvae b u t
.
, ,

i n the maj ority of cases 1 0 per ce n t croto n oil ( made of o n e part

,

croto n oil to n i n e parts paraffi n oil ) w as employed The mixtu re .

was sterilized before i n j ectio n .

The tadpoles were an aesthetized i n chl oret on e 1 to 3 000 ,

made u p i n po n d water Un der the bi n oc u lar microscope . .

,

small glob u les of croto n oil were i nj ected i nto the fin by mean s
of small glass ca n nu lae The larvae were the n tran sferred .

immediately to the observatio n chamber previo u sly de scribed ,

( E R Clark 1 2 ) an d exami n ed un der the compo un d microscope

’
. .
,

i n 1 to 40 00 chl o ret o n e With the microscope t ube horizo n tal

.

a n d a drawi n g board placed be n eath at an a n gle of 4 5 degrees

-
, ,

a n d the u s e of a L eitz drawi n g eye piece ( no it was possible -

.

to make co n sec u tive sketches of i n divid u al cells and to record

the chan ges which occ urred In some cases the regio n w as .
,

watched and freq ue n t records made over periods of six to twelve

ho u rs after which the larvae were ret u rn ed to fresh water over
,

n ight an d the regio n agai n observed o n the followi n g day

,
In .

other cases especially where it was desirable to follow i n dividu al

,

rapidly movi n g and chan gin g cells the regio n was kept un der ,

co n ti nu o u s observatio n for twe n ty fo u r ho u rs or more -

.

The cells an d tissu es prese n t i n the tran sparen t fi n before ,

he prod u ctio n of an i n j ury are the followi n g : ,

1 Stellate co nn ective tiss u e cells

.
—
.

2 Wan derin g cells : a) pigme n ted ; b) n on pigme n ted

.
—
.

3 Blood capillaries li n ed with a si n gle layer of e n dotheli um

. .

4 L ymphatic capillaries also li n ed with e n dotheli u m

.
,
.

5 Blood cells red blood cells an d le u cocytes i n side the

.
- — -

vessels .
 REA C TI O N OF C ELLS TO C RO T O N O IL 22 3

6. Station ary pigme n t cells .

. 7 N erve cells .

. 8 E pidermal cells coveri n g the s u rface of the fi n

,
.

O f these ce lls the co nn ective tissu e cells the wa n deri n g cells

,
-
,

( i n clu di n g migrated le u cocytes ) an d the e n dothelial cells li n i n g

,

the blood vessels an d lymphatics were the o n es observed p art i c

-

ul arly . These are the cells which had formerly bee n studied i n
the n ormal larvae an d i n the experimen ts i n which vario u s
fore i gn su bstan ces had bee n i n j ected i n to the fin The character .
,

behavior an d growth of the blood vessels an d lym phatics i n the

,
-

n ormal tail have bee n described i n detail i n earlier papers The .

n ormal growth of co nn ective tiss u e cells has also bee n st u died

-

in te n sively (E R Clark . . D u ri n g the co u rse of the prese n t

,

stu dy records were also made of the wan deri n g cells of the tissu es
i n n ormal un i n j ured larvae Co n sec u tive records of these cells
.

showed that they are co n stan tly movi n g thro ugh the tissues ,

wa n deri n g aro un d between the co nn ective tiss u e cells an d comi n g —

back ag ai n to their starti n g poin t E ach cell seems to have a

-
.

more or less d efini t e ro u te of its own .

The blood cells of Amphibia have been described by M ax im ow

-

an d by F ri ed so h n The latter divides the le u cocytes

i n to the followin g classes :
.a L eu cocytes witho u t granu les .

1 Small le u cocytes
. .

2 L arge leu cocytes

. .

3 Polymorpho nu clear le u cocytes

. .

4 Small oval cells with oval nu clei

. .

( S pin dle cells ) .

.b L e u cocytes with gra nu les .

1 E osi n ophile le u cocytes

. .

2 Mast le u cocytes ( basophile )

. .

3 Pigme n t le u cocytes (n o t prese n t i n mammals )

. .
22 4 ELI O T R . CLARK AND E L E ANOR LINT O N CLARK
GE N ER A L PI C TURE O F TH E REA C TI O N TO C RO T O N O IL

Immediately after the i nj ectio n of a small glob ule of croto n

oil a decided chan ge was n oticeable i n the fi n n ear the site of
,

i n j u ry The whole regio n became more opaqu e than other por

.

tio n s of the tail This opacity is du e to a granular appearan ce of

.

the epidermal cells and co nn ective tissu e cells of the n eighborhood

-
.

Higher m ag n i fi cat i on shows that their gra nu lar appearan ce is ‘ ’

cau sed by the formation of vac u oles i n the cells An opaqu e .

‘
granu lar zo n e exten ds for a distan ce aro un d the glob ule This
’
.

zo n e is wider proxim ally toward the mu scle than it is i n the

, ,

regio n di stally betwee n the glob ule an d the fi n margi n evide n tly ,

correspo n di n g to the directio n of fl ow of the c u rre n ts of fl uid i n

the tissue spaces The processes of the co nn ective tissue cells
.
-

i n this zo n e are draw n i n the cell bodies become swolle n an d

,

vac u olated an d the n u clei become Visible The e n dothelial

,
.

cells of the lymphatic capillaries withi n this zo n e also become

swolle n an d vac u olated and their nu clei show up plai n ly .

Fiftee n or twe n ty minutes after the i n j ectio n a distortio n of

t h e fin at the i n j u red regio n may be observed Most frequ en tly .

this takes the form of a ben d or fold i n the fi n At other times .

,

partic ularly with the stro n ger co n ce n tratio n s of croto n oil a ,

blister forms aro un d the globule which u su ally disappears ,

withi n a few ho urs .

In the main lo n git u din al blood vessels of the tail an d i n the

-

larger vessels sit u ated n ear the m u scle edge the circ ulatory ,

chan ges described by early observers of i nfl amm at i on can be

, ,

followed the i n creased blood flow followed after a half hou r

—
,

or more by a slowi n g of the c urre n t an d a te n de n cy of the

leu cocytes to stick to the vessel wall In the capillary loops of .

the fi n these chan ges are n o t so eviden t owi n g to the n arrown ess ,

of the vessels which permits the passage of o n ly on e corp u scle

at a time In s u ch a capillary withi n the gra nular area the
.
‘
,
’

circ ulatio n u s ually ceases within an ho ur or two after the i nj ection

of croto n oil Wa n deri n g cells i n the n eighborhood are attracted
.

toward the i nj ected s u bsta n ce .

226 ELI O T R . CLARK AN D E L E ANOR LINT O N CLARK

fl uid an d is fi n ally excl u ded The heap of epidermal cells

,
.

coveri n g the glob u le is appare n tly throw n off at the same time .

A s soo n as the glob u le is extr u ded the co nn ective tiss u e cells ,

-

recover their n ormal shape and the circ ulatio n i n the blood
vessels of the i n j ured regio n is restored The statio n ary l eu .

Fig s . 1 to 5 A
k t h h wi g th g ser es l pi t
i of f i fl m
s e c es s o n e en e r a c u r es o an n a

m tia p d d by i j t i g d p f t
on ro u ce i l (10 p n ec t i p ffi i l )
n a ro o c r o on o e r c en n ar a n o

i t t h d l fi f H y l l v Th
n o e o r sa n ot iv t i a ll h wn i
a ar a . e c o nn e c e- ssu e c e s ar e s o n

so l id b l k t h p i g m t d l
ac ,
yt e d t t d d w d i g ll i
en e tli
eu c o c e s a re o e ,
an an er n ce s n ou ne .

X 1 33 .

F i g 1 Sk t h t t d M y 3ot h t
. e c A M i mm di t l y f t
s ar e th i j
a a . .
,
e a e a er e n ec

ti on o f t il Th b k li
c ro o n o i di t t h xt t f t h i j d
. e ro en ne n c a es e e en o e n u re a r ea a s

e v id d by t h g l t i f t h
en c e t iv t i
e r an u all Th p t h on o f m e c onn e c e— ssu e c e s . e a s o so e

o f t h mi g t i g l
e yt ra i di t d by d tt d l i
n eu co c d w
es areXm k n ca e o e n es an a rr o s . ar s

th le ti f
o ca il l on yt t o a P M L ym
sess l y m ph t i
e p ill y ;
eu c o c e a . . .
, a c ca ar

B V bl d v
. .
, l; O G i l gl b l
oo - e ss e . .
,
o o u e .

which had ass umed the form of small co nn ective tissu e

c o cyt e s , -

cells n ow draw i n their processes and res ume their n ormal

,

amoeboid shape However withi n an ho ur or t w o all of the .

, ,

leu cocytes of the regio n have become motio nless agai n b ut with ,

a spherical form (fi g .
 R E AC TI ON OF C E LL S TO C RO T O N 0 1 1. 22 7

Withi n a few ho u rs after the extru sio n of the glob u le of croto n

oil the opaqu e area which had occu pied the regio n of the croto n
,

oil begi n s to clear up The pigmen ted leu cocytes which had
.
,

crowded i n close to the croto n oil n ow scatter i n all direction s ,

an d wa n der wildly an d rapidly thro u gh the tiss u e spaces away

from the regio n A fter this scatterin g of the pigmen ted l eu

.

c o cyt es the area o n ce occ u pied by the croto n oil

,
is very thi n , ,

Fig . 2 Sk t h f t h
e c o e sa me re gi on as b g
i n fi gu re 1 , e un a t P . M .
,
May
3 ot h , sh o w i g t h w ll f l
n e a o eu c o c yt es , i p i
w t h r o c e sses , w h ch h as f o rm e d a t a sh o r t

di st an c e f m t h i l gl b
ro e o o ule .

an d there may eve n be a hole thro u gh the tail at this p oi n t

However a small opaqu e cl ump co n sisti n g of cell debris sur
, ,

ro un ded by pigme n ted le u cocytes remai n s i n the fi n an d is n o t ,

disposed of for several days .

D u ri n g the fi rst twelve ho u rs or more after the extru sio n of

the croto n oil the clear le u cocytes i n this part of the fi n remai n
motio n less an d spherical Duri n g this period n ew leu cocytes .

were see n to come out from the blood vessels an d to approach -

the site of i n j ury However when they reached the region

.
,
22 8 ELI O T R . CLARK AND E L E ANOR LINT O N CLARK

occ upied by the other leu cocytes they too became ro un d and ,

motio n less like the others .

Soo n after the extru sio n of the oil glob ule the co nn ective ,

tissu e cells o n the border of this torn area have all recovered
their n ormal stellate form and refractile appearan ce They .

begin to chan ge shape an d to wan der i n toward the i n j ured area .

D urin g this stage the processes of all the mesen chym e cells of

F i g 3 Sk
. et c h of t he sam e re gi on at P . M Ma y
.
,
3ot h . Th e oil gl b
o ul e

wa s xt d d
e ru e at P . M . A m a ss of o p q
a ue mat er i a l i s p resen t a t t he s it e

of in j ec t 1 on . T h e l eu c o c t e s h a y v e a ll i d
w th ra wn i p o e e an d b ecom
t he r r c ss s e

sph i l S v
er ca . e er a l r e d bl d oo -
c el l s ha v e es c a p d f o m t h b l o d v e l at
e r e o - ess

th i gh t f t h
e r o e fi el d .

the Vicin ity radiate toward the site of i nj ury D uri n g the .

s u cceedin g day the conn ective tiss ue cells co n ti n u e to move i n —

an d fil l u p the gap i n the tiss u e left by the extr u sio n of the croto n

oil .

The fin al repair of the i n j u ry an d ret u rn of the fi n to n ormal

con sumes several days Figu res 4 an d 5 show s u ccessive stages .

i n this process With the exceptio n of on e or two co nn ective

.

tissue cells which showed an extreme degree of i n j u ry an d which

,
23 0 ELI O T R . CLARK AN D E L E ANOR LINT O N CLARK

i n gest i t A week after the produ ctio n of s u ch an i n j u ry the

.
,

o nly sig n s of reactio n remai ni n g are a slight irregu larity i n the

edge of the fin the radiatio n of the processes of n ear b y c on
,
-

n ec t i v e tiss u e cells toward this defect an d the prese n ce of more

-
,

than the u s ual n umber of pigme n ted le u cocytes ( fi g .

Fig S m gi
. 5 a e re on o n e w ee k ft
i n j e t i n f o t n i l Th i n j u y
a er t he c o o cr o o . e r

is alm t p i d ly
os re a re n,
o a s ull i gh t i g
i t y i n t h d g f t h e fin i s n t i ceab l
r re ar e e e o o e,

to w d wh i h t h p
ar c e f th onn e c t i v
r o cesses o ti u
e c ell a di t A l i ght l y
e- ss e c s r a e . s

g r eat e r n u m b er o f w n d e i n g el l t h n i n a dj in in g
a r c s ag i n i s t i ll p n t
o re o s s rese .

The ge n eral pictu re of a micro i nflamm at i on illu strated i n —

,

fi g u r es 1 to 5 w a s obtai n ed by watchin g the same fi el d withou t

,

i n terruptio n for twe n ty fo ur ho urs or more after the i n j ection—

of a globule of croto n oil D uri n g the succeedi n g days the .

,

regio n w as observed o n ce or twice daily an d a rec ord made In .

other specime n s a certai n type of cell or tissu e was selected and

,

watched i nte n sively d uri n g the co u rse of the i n flammatio n .

These will be described separately .

 R EA C TI O N OF C ELLS TO C RO T O N O IL 23 1

LEU C O C Y TES
1 Clear leu cocytes

This is a large gro up of cells which i n cl u des wan deri n g cells ,

of the tiss u es a n d the leu cocytes which migrate from the blood
vessels an d lym phatics wi tho u t regard to their stai n i n g reac
,

tio n s S u ch cells were i n cl u ded i n on e gro u p i n the presen t study

.

for the reason that i n the livi n g an imal their morphological

, ,

characteristics are the same as is also their behavior toward the ,

croton oil .

A ll wan deri n g cells i n the vici n ity of the i n j ected glob ule
make their way toward the croto n oil soo n after the i nj ection .

A t the en d of a n ho u r the le u cocytes i n the blood vessels j u st

,
-

ou tside the gra nu lar zon e become adhere n t to the walls of the
‘ ’

vessels an d move directly toward the site of i nj ury This .

moveme n t and the chan ge i n shape of these migratin g leu cocytes

take place so rapidly that it is n ecessary to follow i n dividu al
cells co n ti nu o u sly In some cases leu cocytes which had already
.
,

migrated from blood vessels were observed to crawl from the

-

tissu e in to a lymphatic capillary an d the n to proceed rapidly

down toward the ti p by amoeboid moveme n t O n reachin g .

the tip of the lym phatic capillary the leu cocytes were seen ,

to crawl out thro u gh the wall again This observation was .

repeated several times i n differen t larvae an d i n some cases

several leucocytes i n s u ccession were observed to follow each
other i nto a lym phatic dow n to the tip an d o ut again By this
,
.

mean s the leu cocyt es reached the site of i n j ury by a mu ch more

,

direct an d speedy ro ute si n ce they were n ot forced to make their

,

way i n an d out amo n g the fi n e processes of the co n n ective tissu e

cells ( fig .

Figures 1 and 2 give the ge n eral pictu re of the migratio n of

the leu cocytes and the formatio n of a ri n g of sessile leu cocytes
with processes at a short distan ce from the globu le In some .

specimen s in divid u al cells were watched co n sec u tively for a

n u mber of ho u rs The series of sketches i n fi gure 6 shows the
.

chan ges which occ u r i n a leu cocyte which has migrated from a
blood vessel and has become statio n ary after approachi n g within
-
2 32 ELI O T R . CLARK AND E L E ANOR LINT O N CLARK

a short dista n ce of the croto n oil The fi rst processes se n t o ut .

are fi n e and hair like These streamers are withdrawn an d se n t

-
.

ou t agai n thu s chan gi n g the co n to ur of the cell frequ e n tly

, .

A fter a few ho urs these processes become red u ced i n nu mber ,

1 fig 1§ Q

Q
2 00

7
Fi g . 6 S i er es sh o w n i g c h an g es in a s n i gl e l e u c o c t e wh y i ch h as e mi g ra t e d
f ro m a bl d v
oo - esse l a nd b eco me st at i o n a r y n ea r the s it e of in j e ct i on . 2 1 X3 .

Fig . 7 S er i e s sh o w in g
d e g n by g
i l l u o cy t it u t ed
t he c h an es un r o e a se ss e e c e, s a

ne t h in j e t d i t
ar e f t im b f
c e snd e,f t t h xt u i n f t h e c t n i l
or a e e o re a a er e e r s o o ro o o .

At t h o i l gl ob u l w e v y u p fi i l n d t h nn t i v t i u c l l f
e as er s er c a a e co ec e- ss e e s o

th g i n h d b egun t sh w si gn f
e re o a v y At o m n y of t he n
o by
s o r e co er . a ea r

l u co y t
e c w mo b o id
es At
er e all t h el a l u y t h d un d d u p
. a e c e r e coc es a ro e .

T h p i g m n t ed l u c y t
e e w e e wan d i n g a un d
e co t iv ly
es T h co nn t i v
r er ro ac e . e ec e

t u
l ss ll w
e ce h n g in g n d m ov in g t w d t h
s e re c a i t f in j u y At
a t he o ar e s e o r .

cl l u
ear yt e f t h g i on w e t i ll mo t i n le an d p h e i l X1 33
e coc s o e re er s o ss s r ca . .

larger and more stable and the cell the n resembles a small
, ,

immatu re co nn ective tiss ue cell -

.

Metchn iko ff i n his acco u n t of i n fl amm at i o n i n the

tadpole s tail followi n g a b urn with silver n itrate described t h e
’
,
2 3 4: ELI O T R . CLARK AN D E L E ANOR LINT O N CLARK

blood vessels In cases i n which wa n deri ng cells were situ ated

-
.

very n ear the poi n t of i nj ectio n they stopped their amoeboid

,

moveme n t immediately after the i n j ectio n an d se n t out fi n e

processes .

Several specime n s were fix ed i n Bo u i n s fl u id an d stai n ed with

’

E hrlich s haematoxyli n an d co un terstai n ed with eosi n ora n ge G

’
, ,

a n d a u ra n tia an d the whole tail mo u n ted i n balsam ( fig

,
.

These specimen s showed le u cocytes i n stages of mi gratio n or i n

the stage i n which they were covered with fin e processes In .

these specimen s the polymorpho nu clear leu cocytes p red om i

n at ed b u t mo n o nu clear cells with an oval n u cle u s ecce n trically
, ,

placed an d a few cells resembli n g lymphocytes were also prese n t

,
.

The behavior of all the n on pigmen ted leu cocytes was the same
—

regardless of the kin d of nu clei they possessed .

The reactio n of le u cocytes and wa n deri n g cells toward this

i n j u riou s s u bstan ce differs from their behavior toward the
materials i n j ected i n former experimen ts With sterile paraffin .

oil there was very little migratio n of leu cocytes Wan deri n g .

cells of the tissu e moved toward the glob ule som et l m es fl att en i ng ,

o u t o n its s u rface an d the n movi n g away agai n The p araffin .

oil remain ed i n the tiss u e witho u t occasio n in g any distu rban ce ,

an d after the fi rst day or two n o cells reacted toward it i n an y

observable way Whe n carbo n and carmi n e gran u les previo u sly
.
,

sterilized were i nj ected i n to the fi n they were phagocytized by

, ,

wa n deri n g cells which migrated toward them In the case of .

i n j ected fat ( i n the form of olive oil oleic acid cream and yolk, , ,

of egg ) le u cocytes migrated from the blood vessels goin g directly -

,

toward the i n j ected s ubstan ces an d actively en gul fin g small

glob ules of fat In this case o nly part of the wa n deri n g cells
.

were attracted toward the in j ected s u bstan ces .

A ny explan atio n of the exact n at ure of the reactio n of these

le ucocytes and wan deri n g cells toward an i n j u riou s agen t su ch
as croton oil wo uld be mere spec u latio n However the ob ser .
,

vat i on s yielded ab un da n t evide n ce that the formatio n of this

barrier of statio n ary le u coc yt es with processes has a n eu tralizing

effect on the oil and forms an effi ci ent method of localizin g the
i nj ury It was n oted i n every case that co nn ective tissu e cells
.
—
,
 RE A C TI O N OF C ELLS TO C R O TO N O IL 23 5

which had previo u sly shown all the sig n s of i nj u ry cleared up ,

a n d r eg ai n ed their n ormal appeara n ce as soo n as a ri n g of these

‘

sessile le u cocytes with processes had formed betwee n them

an d the glob u le of croto n oil Moreover i n c o n trast with the
.
,

appearan ce j u st described we observed o ne case of i n flamm at i on

,

i n a very yo u n g tadp ole i n which o n ly a few wan deri n g cells

were prese n t i n the tiss u e spaces and o n ly an occasio n al white
blood cell i n the blood vessel In this case the granu lar area
- -
.

was very exten sive Co n n ective tiss u e cells and e n dothelial

.
—

cells became swolle n and vac u olated an d the circ u latio n i n the
blood capillaries stopped i n a regio n three or fo u r t i mes as large
as the o n e i n volved i n the case of a similar i n j u ry i n a n older
tadpole i n which the migratio n of le u cocytes had bee n abun da n t .

In the later stages after the extru sio n of the glob u le these
, ,

clear leu cocytes were see n to act as phagocytes pickin g up ,

extru ded red blood cells —

.

2 . P i gmen ted leu cocytes

These cells are n ormally prese n t i n the tissu e spaces of the

fi n of Am phibian larvae an d they are also fo un d occasio n ally
,

i n side blood vessels and lym phatics They are relatively large
—
.

cells an d co n tai n brown and black pigme n t granu les of varyi n g

sizes Their origin an d their relatio n if an y to the large bran ch
.
, ,

i n g chromatophores of tadpoles have n o t bee n st u died In .

s tai n ed specime n s they may be see n to possess a si n gle n u cle u s ,

u s u a ll y ro un d or oval i n shape M ax i m ow an d F ri edsoh n .

c on sider the pigme n ted leu cocyt es to be a special class of cells

withou t an y homologu e amo n g mammalian blood cells -
.

The pigmen ted le u cocytes are stro n gly attracted toward the
c roto n oil an d those located n ear the site of i n j u ry wa n der
,

toward it soo n after i n j ectio n These cells are remarkably .

resistan t to i n j ury They wan der i n thro u gh the gran u lar area
.
‘
,
’

past the li n e at which all the n on pigmen ted le u cocytes have -

s topped an d make their way directly i n to the opaqu e regio n

,

immediately surro un din g the croto n oil an d even fl at t en o ut ,

d irectly again st the oil glob u le In some cases pigme n ted .

2 36 ELI O T R . CLARK AN D E L E ANOR LINT O N CLARK

le u cocyt es were fo un d co n tai n in g small refractile glob ules of oil

an d i n o n e i n sta n ce
,
s u ch a pigmen ted cell w a s followed as it
,

approached a mi n ute droplet of oil an d proceeded to i n gest i t .

These cells co n ti n u e to s u rro un d the glob ule of croto n oil

d u ri n g its soj o urn i n the fi n an d they occ u py the site of i n j ectio n
,

u n til after the glob u le has bee n extr u ded Two or three hou rs .

after the extru sio n they begin to scatter i n all directio n s and to
,

wan der away from the sce n e of the i n j ury The cleari n g up .

of the opaqu e area which occ u rs at this time is chi efly d u e to

‘

the dispersal of these pigme n ted leu cocytes Most of them .

wa n der away thro u gh the tissu es while some of them were ,

observed to crawl thro u gh the walls of the n eighbori n g lymphatic

vessels A number of pigme n ted le u cocytes remain at the site
.

of i n j u ry d u ri n g the s u cceedi n g days an d appear to be i n st ru

me n tal i n the disposal of the small m ass of debris which remai n s
i n the tail after the extru sio n of the glob u le This opaqu e .

material grad u ally dimin ishes i n amoun t du rin g the n ext few
days an d at the en d of a week the place of i nj u ry is marked
, ,

o n ly by o n e or two of these pigmen ted leu cocytes .

L ike other leu cocytes these pigmen ted cells were also see n to
,

act as phagocytes of extravasated red blood cells -

.

3 . The conn ec ti ve— ti ssu e cells

Th e ge n eral picture of the reactio n of these cells toward the

croto n oil has already bee n give n In on e specime n this class .

of cells was followed with partic u lar care In this case a record .
,

was made before i n j ection of a selected portio n of the tail an d

, ,

every mese n chyme cell i n the regio n w a s draw n By u sin g two .

pigme n t cells as markers it w as possible to i n sert the globu le

,

of croto n oil i n the exact positio n desired O n examini n g this .

regio n immediately afterward i n the observatio n chamber it

was foun d that the glob ule had bee n i n serted witho ut disturb
i n g any of the cells of the tail The co n n ective tissu e cells were
.
-

n umbere d an d observed co n ti nu o usly for twelve hou rs O n .

acco un t of the mild n ess of the i n flammatio n prod u ced i n this

S pecime n it was possible to follow i n divid u al cells throu ghou t
,

the process .
 2 38 ELI O T R . CLARK A ND E L E ANOR LINT O N CLARK

Fig . 8 Sh o ws a g ro u p o f c on n e c t iv i
e -t ssu e c e l l s b fe o re (A ) an d i mme di a t el y
a ft er ( B) t he inj i
ec t o n o f c r o t on oil . Th e gl b l (0
o u e . G ) w as i n sert e d p r o x i
.

m all y t o t hese c el l s . The p iti

os on l at er o c cu p i d b y th
e g l b ul
e oi l o e d tt d
is o e

i n t h e fi rs t s k et c h . The sa me c ell s are show n in b th d wi g (

o ra n s seenu mb e s ) r .

T he c el l p ro c esses bm e sh or t er an d t h
ec o i ke
c r an d t h wh l e gi o e re on h i nks r s .

T he i n c rease in an ast o m o ses et w een t he b p ro c e sses o f t he di ff er en t c el l s sh ow n

i n t he k et ch m d af t t h
s a e er e i nj c t i n
e o i pp
s a ar en t r at h er t h an r e al , s n c e i in th e

nor m l t ad p l e m n y f t h
a o a o e b nhs
ra c e f th p
o e ro c esses ar e t oo fi n e t o di t i n gu i sh
s

W i t h ou t t h o i l i mm i on
e ers . X400 .
 REA C TI O N OF C ELL S TO C RO T O N O IL 23 9

Fig . 9 S i ill er e s i gle c onn e t i v e t i ssue cel l i tu at ed

u st rat n i g t he c h an g es in a s n c -
s

ne t h g l b u l e f c t n o il
ar e I llu st at es i u i n j u y nd comp l et e e ov e y
o o ro o . r ser o s r a r c r .

A S k et h m d e Jun 4t h 1 P M j u st b ef o
. c a i n j e t i on B S k t h m d Jun e
e ,
. .
,
re c . . e c a e

4t h ,
P M n e h l f h ou f t i n j t i n f c t n oi l P oces es sh t ened
. .
,
o -
a r a er ec o o ro o . r s or ,

nu c l eu s v i ib l e v c u l s p e s nt C Sk t h m ad Jun e 4t h t 4 P M p
, a o e es r se . . e c e a . .
,
r oc ess

m t l y w i t h d wn m
os v u ol p ent D Sk et h m ad e Jun e 6t h
ra , ore ac es res . . c ,

PM . ab u t tw en t y fiv
. h ou s af t e t h
o xt u i on f t he i l gl bul C ll h as
- e r r e e r s o o o e . e

e ov e ed p di t t w d t h si t e of i n j u y E Sk et h m ad e Jun e
—
r c r r o cesses ra a e o ar e r . . c

7 t h 4 P M t h ee d y
,
f t r t he i n j e t i on f c t n i l n d tw o d y s f t t he
. .
,
r a s a e c o ro o o a a a er

e xt u i n r f t he g l b u l e
s o C el l h com p let el y eg i n ed i t n m l app ea n c
o o . as r a s or a ra e .

A, X200 = B-E X300 , .

Fi g . 10 S i er es of s i v e t i ssu e ell si t uat ed v y n e r

k et c hes o f an o t h er conn ec t — c er a

t t he i t
o f inj t i n
s I l lu t t e x t m e g d f i n j u y T h i i n f t h e
e o ec o . s r a es re ra e o r . s s o e o

f w e x am p l es b se v ed i n w h i c h t h e cell did n ot e ov
e o rA Sk et h m d e Jun e r c er . . c a

4t h 1 P M b f e i n j ec t i n f
,
. .
,t n oi l B Jun e 4t h
e or P M fif t n m i nu t s o o cro o . .
,
. .
,
ee e

af t e t he i n j e t i on P o e ses sh o t e n d t h i k
r c N u l eu s V i s ib l e
. C Sk t c h
r c s r r a c er . c . . e

m d e Jun 4t h
a P M eP s W ith d wn v u ol i z t i n
,
D Sk et h . . ro c es se ra ,
ac a o . . c

m d e Ju n e 5t h
a P M A b out t n h u aft e ext u i n f t h i l gl obu l
,
. . e o rs r r s o o e o e .

P t i ll y l l t h t he onn t i v e t i ue l l of t h eg i on h av e
rac ca a e o ve d r c ec - ss ce s e r rec o re .

T h i s c ll i t h i n w ll d c n t i n in g v u l
e s a g an ul
-
N u l eu s d e n t
a e sa co a ac o es o r r es . c o s o

O u t h f l l w i n g d ay ( Jun 6) t h i c ll w s st i ll p h d wy
'
sh w o . e o ont a e s e a r ese as s a o

o ut l in e n t i n i n g a f ew g an u l
co i n B wn i n m o v m n t
a O n J un 7 t h n t e r es ro a e e . e o rac

of i t ul d b e f un d
co A X200 ; B C D X300 o .
, , , , .
2 40 ELI O T R . CLARK AND E L E ANOR LINT O N CLARK

The vac u olizatio n of the last two types has bee n described
rece n tly by L ewis “
i n st u dies of the dege n eratio n gra n u les
an d vac u oles
” of cells i n tiss u e c ult ures In o ur st u dies it is .

n oteworthy that cells showi n g su ch marked dege n erative ‘ ’

chan ges co u ld recover so rapidly an d completely A s soon as .

the ri n g of sessile leu cocytes had formed the vac u olated c on ,

n ec t i v e tiss u e cells
—
outside of this barrier became clear an d
, ,

refractile the cell bodies became sle n der an d were occ upied
,

solely by the nucleu s an d the lon g bran chin g processes again

,

exte n ded to their n ormal le n gth A fter the extru sio n of the .

croto n oil practical ly all of the co nn ective tissu e cells ret u rn ed

,
-

to n ormal an d their recovery w a s amazi n gly rapid O n ly on e .

or two cells were lost an d these were always cells which had
previo u sly shown the most extreme type of i n j u ry O n e of .

these cell s watched co n ti nu ou sly became paler an d paler and

, ,

fi n ally remai n ed as a n o u ter rim with a few gran ules i n sid e

before disappeari n g en tirely (fi g 1 0 D ) O n the other han d a .
,
.
,

nu mber of cells which had show n the most extreme type of

reactio n toward the irritan t were followed thro ugho ut the i n

flammation an d were seen to recover completely soo n after the
extru sio n of the oil (fi g .

This vac u olizatio n and roun din g up of the co n n ective tissu e -

cells i n co n trast to the behavior of the le u cocytes is a p urely

, ,

passive reactio n However i n the later stages of the process

.
, ,

after the extr u sio n of the croto n oil these cells play a very activ e ,

part They wan der actively toward the gap i n the tissu e left
.

by the extru sio n of the oil an d some of them disappear from ,

View temporarily i n the opaqu e regio n s at the site of the i nj u ry

,
.

D u ri n g this period of migratio n the processes of the co nn ective ,

tissu e cells radiate towards the i n j ured regio n The en ds toward .

the gap are bru sh like i n appearan ce while the opposite e n ds

-
,

are ro un ded (fig O n e of the aut hors ( E R Clark

. 1 2) . .
,
’

had previou sly st udied the co nn ective t i sssue cells i n the n ormal —

tadpole an d fo un d by observi n g the same cells for sever al

, ,

weeks that they are n o t fix ed b ut co n stan tly chan ge their

, ,

positio n i n most cases grad u ally wa nderi n g i n the directio n of

,

the fi n margi n However after the extru sio n of the oil the c on
.
,
2 42 ELI O T R . CLARK AND E L E ANOR LI N T O N CLARK
T H E BL OO D -V E SS E L S

The blood vessels sit u ate d i n the area i n which the c o n n ective
-

tissu e cells became vac u olated showed d efin i t e sig n s of i n j ury .

The en dothelial cells became vac u olated an d the nu clei became

more promi n e n t withi n a few mi nu tes after the i n j ectio n The .

vessels became co n stricted an d the circ ulatio n ceased abo u t a

half ho ur later At some poin ts these vessels co n stri cted un til
—
.

they con sisted of a solid hyali n thread At other poi n ts they still .

"
o=3 0 a m 7

2 : 30
pm .

M
10 0 0
P. .

Fi g . 12 D r aw n i gs sh ow n i g t h r ee su c c e s s ive st ag es i n t he mo v em en t of t he
tw o c el l s g an d h, w h i c h are sh ow n i n t h e first t g
s a e A M ) i n fi gu r e 1 1
. . .

T h e m ar k d d t i i l g th f p
e re uc on n en o r o c e ss es du r ng th is ( r el at i v el y ) rap id m ov e
men t i s p t i ul l y t i b l i t h
ar c ar no ce a e n e l ast st a g e P . M an d 1 0 P M resp ec
. . .
,

t i v el y ) . 0 G . i l gl b l
.
, X320 o o u e . .

co n tai n ed red blood cells an d leu cocytes There was n o migra —

.

tio n of white blood cells from the vessels within the granu lar —
‘ ’

area O utside this regio n the blood vessels showed the typical
.
,
-

chan ges of i nflamm at i on acceleratio n i n the fl ow followed after ,

about an ho ur by a slowi ng of the c u rre n t and diapedesis of

le u cocyt es I n side the granu lar area the frequ e n t occ urren ce
.
‘ ’

of hemorrhage gave fu rther evide n ce of i nj u ry to the vessel wall .

An d twe n ty fo u r ho u rs or more after the i n j u ry a retractio n of

-
,

on e or more of these blood vessel spro u ts or loops located withi n -

 REA C TI O N or C E LL S TO C RO T O N O IL 2 43

this granu lar area w as always observed O u the co ntrary t h e

‘ ’
.
,

vessels j u st o u tside this zo n e u s u ally showed a ten de n cy to se n d

o u t n ew spro uts d uri n g this period A day or t w o after the
.

extru sio n of the glob ule d urin g the stage of repair the vessels of ,

the i n j u red area began to se n d ou t n ew S prou ts an d formed n ew

loops o n ce more .

The reaction of the blood vessel e n dothelium toward the

—
.

croto n oil appeared to be e n tirely passive In a nu mber of larvae .

records were kept of the nu clei of the blood vessels both i n side -

an d o u tside the i n j u red area b ut n o u n u s u al cha n ges were n oted

,
.

There was n o sign of any proliferatio n of en dothelial nu clei to form

wan derin g cells or en dothelial leu cocytes O ccasion ally leu co .

cytes were observed to flatten out on the i n terior of the wall of a

blood capillary an d to pass thro u gh the wall sideways D urin g .

this process there were momen ts at which s uch a le ucocyte

greatly resembled a nu cleu s protrudi n g in to the l u men of the
vessel and a little later it appeared very mu ch like a nu cle u s
,

detachin g itself from the wall However i n the co n sec u tiv e

.
,

observatio n s of livin g specimen s it w as always possible to follow

the i n dividu al e n dothelial nu clei an d to observe that the phen om
e n o n co n sisted of the emigratio n of a leucocyte thro u gh the wall ,

rather than the formation of a leu cocyte from an en dothelial

cell.

T H E L YM PHATI C S

The lym phatic e n dothelium plays b oth an active and a passive

part i n the i nfl amm at i on set up by the presen ce of a drop of
croton oil The lym ph capillaries withi n the granular area
.
‘ ’

show the chan ges characteristic of i n j ury displayed by the

conn ective tissu e an d epithelial cells The areas s urro un din g
.

the nu clei of the lym phatic became swollen opaqu e an d va ou , ,

o l at ed partic ularly toward the tip S u ch V essels u s u ally

, .

re tracted at abo ut the time of the extru sio n of the oil A day .

or two later they sen t ou t n ew spro u ts i n to the rege n eration area .

The vessels o utside the gran ular area showed the fin e processes
‘ ’

characteristic of fun ctio n al activity (E R Clark 09 ) an d eve n

. .
,
’

co n ti nu ed to in crease i n le n gth d uri n g the period of active i n flam

2 44 E LI O T R . CL A RK AND E L E ANOR LINT O N CLARK

matio n S u ch growth o n the part of blood vessels and lym

.
-

p h a t i c s however did n o t appear to be i n excess of the n ormal

, ,

rate of growth .

The mann er i n which lymphatic capillaries served as paths

by which the migratin g le u cocytes were e n abled to reach the

Fi g . 13 Sk e t c h es o f c el l s f r om a fix ed s p im
ec en . Fiv e h ou r s a ft er t he i nj ec
t i o n o f c r o t o n o il , th i s l ar v a w as e x mi d
a ne an d f o un dt o b e i n t he st a g e i n wh i ch

a r n i g o f sess il e l eu c oc t es w as i n y p r oc ess o f f o rm a t i on . T he t a dp ol e w as fi x ed

in B i ou n s
’
i n d i n l um h m t y h n unt e t i n d
fl u i d , t h e w h o l e t a l w as st a i e a ae a ox , co rs a e

in ineos ng G nd ,
u nt i
ora d hyd t ed l d i n xy l l m u n t d i n d m
e ,
a a ra a, e ra , c ear e o ,
o e a ar .

Th k t h w m d w i t h t h i l i mm si n l n A A m b id l eu y t
e s e c es er e a e e o -
er o e s . . oe o coc e .

B nd C S il l u yt W ith p
a . ess e D N m l
e nn
coc t iv t i ue ll
es ro c esses . . or a co ec e- ss ce

ou t id i nj u d
s e E C nn t i v t i u
re ll n
ar ea th
. t n i l gl b u l
. F o ec e- ss e ce e ar e c ro o -
o o e . .

C nn t iv t i u
o ec ll i n l o e— p ximit y t t h
ss e ce t n il h wi n g xt m t yp
c se ro o e c ro o o ,
s o e re e e

of inj u y ( h m tinr tt d t h u g h t h ll ) XGOO

c ro a sc a ere ro e ce . .

sce n e of actio n more q uickly has already bee n described There .

are steps i n the migratio n of s u ch a leu cocyte thro u gh the wall

of a lym ph vessel i n which it is easy to be deceived i n to the
-

belief that the proliferatio n of an e n dothelial le u cocyte is taki n g

place for the le ucocyte ofte n comes o u t of the vessel n ear the
,

poin t where a nu cleu s is located But co n ti nu o u s observatio n .

2 46 ELI O T R . CLARK AND E L E ANOR LINT O N CLARK

while those o utside the area co n ti nu ed to se n d ou t fine processes

an d to grow i n a n ormal man n er b u t i n n o case do w e fi n d an y ,

evide n ce of e n dothelial proliferatio n to form wan deri n g ce lls .

In a number of cases hemorrhages occ u rred from blo od capil

,

l a ri es located n ear the croto n oil The extravasated red blood .

corp u scles were taken up by the lymphatic vessels as well as by ,

le u cocytes of both the pigmen ted an d clear varieties The .

method by which the occasio n al extru ded red blood cell foun d —

i n the n ormal t ail is take n up has bee n described by o n e of the

au thors (E R Clark . .In this case the process was the
,

same b ut o n a mu ch larger scale I n stead of on e S pro ut growi n g

,
.

toward a si n gle blood cell a num ber of processes appeared o n

-
,

both sides of the lym phatic an d by mea n s of these several red , ,

blood corpu scles were picked up and simu ltan eo usly taken in to
the l umen of the vessel where they were seen to move alo n g ,

the lymph capi llary to the mai n lon gitudi n al lym phatic A fter .

two or three d ays s uch an area would be almost clear of blood

,

cells a n d the maj ority of these wo u ld have been salvaged by the

,

activity of the lymphatics .

In a n umber of larvae an edema developed i n the i n j u red

region This u su ally occ u rred soo n after the extru sion of the
.

globule althou gh it was sometimes n oted o n the followi n g day

,
.

The edema may be demo n strated by measu rin g the thickn ess
of the tail by mean s of the fi n e adj u stmen t screw The appear -
.

an ce of the tail is altered i n s u ch areas ; the regio n becomes

clearer the co n n ective tiss u e cells more widely separated an d
,
-
,

all the leu cocytes i n cl u din g the pigmen ted o n es ro u n d up i n to

, ,

spheres The lymphatic S pro u ts of the area become markedly

.

diste n ded at the tip This e n largeme n t is eviden tly d ue to an

.

i n crease i n the fl u id co n te n t of the vessel for the endotheliu m ,

is represe n ted merely by a thi n li n e This differs from the .

e nlargeme n t of the lymphatic tips which occ u rs soo n after the

i n trod u ctio n of the croto n oil and which is d u e to a swelli n g of
the e n dothelial cells proper Figu re 1 5 shows su c c esssi v e records .

of a lymphati c spro u t i n an i n fla m matio n area which has b ecome

edemato u s after the extr usio n of the glob ule of croto n oil Com .

pariso n of the diff eren t drawi n gs with the measu remen ts of this
 REA C TI O N OF C E LLS TO C RO T O N O IL 2 47

regio n of the tail recorded at i n tervals shows that the lymphatic

capillaries expan d with an in crease i n the thick n ess of the tail
an d co n tract agai n whe n the tail res u mes its n ormal dime n sio n s .

A bsorptio n thro u gh the lymphatic is evide n tly active d uri n g

i nfl amm at i on a n d appears to i n crease with an i n crease i n the
amo unt of fl ui d prese n t i n the tiss ue o u tside .

Fig . 15 Sk et c h e s i g
y p i p u t w i th in t h
sh ow n the r eac t o n s i of a l m h at c s ro e

ar e a aff t d by t h ec e t n il In t h i p im n n d m d v l p d
e inj t d ec e cro o o . s s ec e a e e a e e o e

a f t t h e x t u si n f t h g l b u l R d f t h t h i k n
er e r o o f th t il t th
e o e . ec o r s o e c ess o e a a e

p in t un d b v t i n w m d by m
o er o ser f t h fin
a dj u t m n t
o w
er e A a e ean s o e e- a s e sc re . .

Sk t h m d M y 29t h
e c a M e i m m di t l y f t t h i n j t i n T h i k n
a ,
P . .
,
e a e a er e ec o . c e ss

o f t h t i l 55 y
e B 3 a M W ll f t h l ym p h t i w l l n n d g n u l t t h
, . . P . . a o e a c s o e a ra ar a e

t ip T hi kn
. f t h t i l 66
c 6 L ym p h t i
ess o t eM ( gl b ul
a xt u d d
,
u
,
. . a c a P . . o e e r e

a t E n d th l i l w ll f l ym p h t i m t h n d t h i n lu m n n l g d
o e a a o a c s oo a ,
e e ar e .

T h i k n s f t 1 1 00 p D 6 M L ym p h t i n t w id b u t t i l l l g d t
c e s o a , . . P . . a c o so e, s en ar e a

t h tip
e T hi kn . f t il 74 u E M y 30 t h 1 0 A M
c es s o L ym p h t i p i l l y i s
a ,
. a ,
. . a c ca ar

n m l in iz
or a nd nt u nd h
s t
e a t d T hi kn
co f t i l 57 y
o X 178
r, a as r e r ac e . c ess o a .
.
2 48 ELI O T R . CLARK A ND E L E ANOR LINT O N CLARK
DIS C USSI O N
The s u bj ect of i n flam m at i on has bee n st u died so exte n sively
a n d the literat u re is so vol umi n o u s that it has seemed i n advisable

to attempt an exte n sive review i n this paper and we shall merely ,

i n dicate b ri efly some of the diverge n t Views Ma ny of the early .

st u dies of C ohnhei n Metch n ikoff an d A rn old were made on

, ,

livin g material and the ge n eral feat ures of i n flamm at i on an d

,

man y of the sp ecifi c details as di sc overed by them are accepted , ,

by most pathologists With regard however to the derivatio n .

, ,

an d possible tra n sformatio n of the cell s taki n g part i n i n fl a m

matio n the widest differe n ce of opi n io n may be fo un d amo ng

,
—

b oth pathologists and an atomists The qu estio n as to the .

s p ec ifi ci t y of this gro u p of cells i n n ormal growth has n o t yet , ,

bee n settled by an atomists O n the side of n on sp eci fi ci ty are .

—

fo un d M ax i m ow D an ch ak off W ei denrei ch and partic u larly

, , ,

Mollier who altho ugh their st u dies have bee n c onfin ed

,

main ly to early stages of embryo nic develop me n t believe to a , ,

greater or less exte n t i n the i n terchan geability of e n dothelial

,

cells leu cocytes mese n chym e and retic ul um Schulte

, ,
has ,
.

gathered together data an d opi n io n s which appare n tly s upport

this View an d presen ted them i n the form of a pla u sible tho u gh
n ot e n tirely co n vi n ci n g arg ume n t The spe ci fi ci t y of the cells of .

the gro u p has bee n advocated partic ularly by Mi n ot Marchan d , ,

M ac Callum an d Mall an d his cowork ers Sabin E van s E R

,
—
, ,
. .

Clark altho u gh Mall appare n tly recognizes an exceptio n i n

—

the retic ulo e n dothelial cells of the spleen liver an d en do

‘
-
’
,

cardium and E va n s ,
i n the formatio n of free phagocytic
cells from the Kupffer cells of the liver followi n g i nj ectio n s of ,

t ubercle bacilli .

In st u dies made o n i n flam m at i o n the vari o us views as to the ,

possible tran sformatio n of o n e type of cell i n to an other have

been s upported by the followi n g : a ) co nn ective tiss u e cells may —

be derived from le u cocytes : R ibbert Metch n iko ff

M ax i m ow 03 ) b) e n dothelial cells from le u cocytes : R ibbert
—

c ) co nn ective tiss u e cells from e n dotheli um : R ibbert

-

M ax i m o w i n i n te n se i nfl amm at i o n ; d) e n dothelial cells

2 50 ELI O T R . CLARK A ND E L E ANOR LINTO N CLARK

seaso n i n order c hi efly to follow the same le u cocytes thro u gho u t

, , ,

the process This we s u cceeded i n doin g an d fo un d i n every

.
,

in stan ce that the le u cocytes revert after the extru sio n of the ,

oil to typ ical ro un ded or amoeboid le u cocytes We feel morally

,
.

certain that had we see n these leu cocytes i n sectio n s we sho uld ,

have labelled them co nn ective tiss u e cells -

.

A gain whe n leu cocytes were fi rst seen emer g in g from l ym

,

phatic capillaries they often gave the appearan ce of p u llin g

,

away from t h e endothelial cells an in terpretation which might —

very readily be made from sectio n s However con tinu o u s .

,

observation of lym phatic capillaries showed that the leucocytes

always e n tered the lym phatic vessel from the ou tside that they , ,

moved a variable distan ce alo n g the l um en and the n passed

o ut thro u g h the e n dothelial wall In their moveme n ts amo n g .

the co nn ective tissu e cells agai n leu cocyt es frequ e n tly pass over
—
, ,

them so closely that o n ly co n tinu o u s observatio n shows that

they are n ot bein g formed i n some way by the co nn ective tissu e -

cells .

While we hesitate to ge n eralize from a set of observatio n s

restricted to a sin gle species of a lower vertebrate still our ,

res ults are so defini t e an d so positive that it seems fair at least to , ,

advan ce the hypothesis that similar tissu es i n other ani mals

react similarly Certai nly on e mu st be very skeptical toward
.
,

co n clu sio n s as to cell tran sformatio n s which are based on st udies

of fix ed material especially whe n s u ch st udies yield s u ch a
,

variety of c onfli ct i ng Views .

O n e matter remai n s to be cleared up by further st udies ,

n amely the possibility of the tran sformatio n s of on e type of

,

leu cocyte i n to a n other O wi n g to the great di ffi culty i n seein g

.

the o utli n es of the nu clei i n the cells stu died we did n ot attempt ,

to disti ng uish i n the livi n g be twee n the differe n t types of

, ,

amoeboid cells except for the pigme n t cells and o ur st udies

, ,

of total mo unts of fi x ed S pecime n s are still i n complete Some .

i nvestigators hold that polym orpho nu clear leu cocytes may be

tran sformed i n to mo n o nu clear cells (J an owski Metchn ikoff ) ,
.

A s u spicio u s fact i n this co n n ectio n is the oft repeated ob ser -

vatio n that early i n i nflamm at i on polymorpho nu clears predom

, ,
 REA C TI O N OF C ELLS TO C RO T O N O IL 2 51

i n at e ,while mo n o nu clears are mu ch more numero u s later This .

is explai n ed by man y as due to the destru ctio n of p olym orph o

nu c l ears In o u r st u dy we saw very little if any destru ctio n of
.
, ,

le u cocytes M oreover the roun din g up of all n on pigme nted

.
,
-

le u cocytes after the extrusio n of croto n oil an d rete n tion of this

shape for several days has led us to wo n der whether i n this
phase the nu clei also ro un d up This poin t we hope to take up .

i n fu t ur e st u dies .

S UMMAR Y

The reaction s and chan ges un dergon e by the cells i n the tail
of Am phibian larvae i n respo n se to i n j ection s of minute glob ules
of croto n oil are as foll ows :
1 Co nn ective tiss ue cells man ifest all grades of i n j u ry
.
-
,

depen din g u pon their n earn ess to the oil In the n earest cells the .

bran ched process es are withdraw n the body of the cell becomes ,

m u ch swollen an d vac u olat ed an d the nu cle u s becomes easily ,

Visible L ater the vac u oles run together an d the cell becomes a
.
,

delicately walled sac co n ta i ni n g a few granules s u spen ded i n

,

fl ui d .L ater the o utlin es of the cell are lost completely Com

,
.

p l et e destr u ctio n however is limited to a very few cells

, ,
A .

little further away cells become swollen and vac u olated processes
, ,

become mu ch shorter b u t are n ot retracted e n tirely an d the

, ,

n u cle u s becomes visible a n d may be i n de n ted by vac u oles

,
.

S u ch cells may recover completely the vac u oles disappear the —

,

cell becomes smaller the nu cle u s less disti n ct n ew processes are

, ,

se n t ou t and moveme n t of the cell resum ed Still further away .

,

the cell s may show merely slight swelli n g with granu latio n .

The cells on the border zo n e of the area affected react by movi n g

toward the oil at a rate mu ch faster than the n ormal rate of
—

moveme n t of these cells In this moveme n t the number an d .

,

len gth of their processes are mu ch red u ced un til they resemble ,

the typical fil oblast .

2 a ) N o n pigme n ted le u cocytes ( i n cl u di n g the wan deri n g cells

.
—

o u tside the blood vessels ) respo n d immediatel y by movi n g toward

-

the globu le of croton oil When abo ut half w ay betwee n the .

-

o u ter limits of the a ff ected co nn ective tiss u e cells a n d the oil -

,
 ELI O T R . CLARK AN D E L E ANOR LINT O N CLARK

they become statio n ary an d se n d ou t man y fi n e processes .

These processes are withdraw n an d re n ewed co n ti n u o u sly ,

tho u gh te n di n g to become more and more stable At this stage .

they resemble very closely small co nn ective tiss u e cells They -

.

are n ot tran sformed i n to co nn ective tissu e cells however for —

, ,

eve n t u ally probably correspo n din g to a diminu tio n i n the

—

i n ten sity of the croto n oil action the processes are retracted
- —

an d the le u cocytes res u me amoeboid powers A fter the oil has .

bee n extr uded the le u cocytes remain statio n ary for several day s
,

i n a spherical co n ditio n after which they scatter thro u gh the

,

tissues Wan deri n g cells free i n the tissue S paces an d le u cocytes

.

which emigrate from the blood vessels do n ot di ffer i n thei r -

reactio n L e u cocytes may en ter lym phatic vessels an d emerg e

.

again b ut we saw n o evide n ce whatever of the origi n of le ucocyt e s

,

from lym phatic or from blood vasc u lar en dothelium —

.

b) Pigme n ted le u cocytes move toward the croto n oil glob ul e -

an d are more resistan t to its actio n tha n the other le u cocytes ,

for they may eve n come i n co n tact with the oil glob ule withou t
showin g an y diminu tio n i n their motility .

3 Blood capillaries n ear the glob ule show thicke n i n g and

.

vac u olizatio n of the en dothelium an d later n arrowin g of t h e , , ,

l um en with i n man y cases retractio n of e n dothelium Ex t rav

, , ,
.

a sat i o n may occ u r thro u gh the i n j u red part of the wall .

.4 L ymphatic capillaries n ear the glob ule also show vac u oli
z at i on of the e n dotheli u m a n d their l um e n may become m u ch
,

diste n ded partic ularly i n case an edema develops The i n j ured

,
.

capillary may be retracted b ut may grow out again later after ,

the oil globu le has been extruded .

.5 Thro u gho u t the i n fl amm at o ry reactio n each type of cell ,

le u cocyte co nn ective tiss u e cell blood vessel e n dothelium an d

,
-
,
-
,

lymphatic e n dotheli um main tai n s its sp ec ifi ci t y and there i s

—

n o evide n ce for the tra n sformatio n of o n e type of cell i n t o

a n other .
2 54 EL O I T R . CLARK AND E L E ANOR L INT O N CLARK
A 1 9 03 E n t s t e ll un g S t kt u ,
ru r, uud Ve ra n d e r u n g d e s N a rb en g c
w ebe s . Z i e gler ’
s B i t g Bd 34
e ra e, ,
1 9 03 .

Bc nt ra g c z u r H i s t o l o g l e ( l c r e i t e r i en E n t z u n d u n g
1 9 05 Z i e l e r s Be i g . g ’

t r a g c , Bd 3 8 , S 3 0 1 . . .

1 9 10 U
e b e r e m b r y o n a l e En t w w k c l un g d c r l u t e l l e n b e i S c l a c h ic r n B z
u n d A m p h l b l en V e rh a n d l d e r A n a t G e se l l s c h a f t . F 1 s c h e r J en a . . .
,
.

M E T CH N l K O F F , E 1 89 3 L eco n s su r l a a t h o l o e c o m a r e d c l m fl a m m a t i o n p gi p é ’
.

P i ar s, 7 th L e co n , p . 111 .

MI N ERVI NI ,
R . 19 1 1 Ub e r di e N b i ld g
eu un von Bl u t g e fa s sc n . Vi rc h A rch ,

Bd 204 S 7 5 .
,
. .

MI NO T C S ,
. . 1912 The d v e e lo pm en t of t he b lo o d . In H u m an E mb y r o o l gy ,

ed . by Ke i b e l an d M a ll ,
vo l . 2, p . 498
M O LLI E R S ,
. 191 1 Ub er d en B a u de r c a p i l l a re n M i l z v c n en ( M l l z s m u s \ Al ‘

Cl l l V .

f u r m i k r o sk . A n at . un d En t w Bd 76 S 608 .
,
.

RI BBE RT 1 89 0Ub d er ie Be t h e i l u n g d c r L e u k o c y t e n an de r N b i ld g
eu un des
Bi d g w b
n e e e es . Ce n t ra b l f . . P th
a ll g e m a . u .
path . A n at Bd 1 , S 667 . . .

SA IN B ,
F . R . i g i n an d e e o e n
1 9 13 Th e p or y d v l pm t o f t he l y m ha t i c s s tem .

J o h n s H o p k m s H o s p i t a l R e p o r t s M o n o g r n ew s e r i e s n o 5 ,
.
,
.

S HUL
C TE , H VO N W . 1 9 1 4 E a l y s t a g es o f v a s c u l o g e n e s i s n t h e c a t
. M e m o i rs
r l .

o f Th e W i s t a r I n s t i t u t e o f A n a t o m y a n d B i o l o g y n o 3 ,
.

ZI E G L E R ,
E 1 9 05 L eh r b u c h d e r a l l g e m e m e n P a t h o l o g i e u n d d e r p a t h o l A n a t .
,

1 1 t h ed G F i s c h e r J en a S 3 68
.
,
.
, ,
.
THE AM E R I C AN J OU R N A
L OF AN x r omr vo n 27 NO 3
J U LY
.

, 1 920
R esume n por el au tor ,
J ames W . Pepez .

M u sc ulat u ra de los atrios del coraz on .

El obj eto del prese n te trabaj o es la d em o st raci on de los fas

c i c ul o s m u sc u lares de los atrios ( au r i c ulas de la termi n olog i a an

tigu a N del ,
. L a mayor parte de ellos se origi n an en el ta
biqu e i n teratrial L a ban da i n teratrial y los f asc i cul o s exter n os
.

del atrio derecho se origi n an en el n odo si n o auric ular 5 trio -

. .

derecho : L a ban da i n teratrial arran ca del n odo y se extien de

hasta el apé n dice izqu ierdo L os fasc i cul o s exter n os tambié n .

arran ca n del n odo exte n dié n dose sobre el atrio L a cresta an .

t eri o r derecha se origi n a en el tabiq u e y da or i ge n a los m uscu l os

’ '

p e c t i n a d o s a n teriores L a cresta posterior derecha arran ca del

.

tabiqu e dan do or i ge n a los muscul os p ec t i n ad o s posteriores .

Se n o ve n oso : El f asc i cul o i n t erc av al se origi n a en el tabiqu e ,

exte n dié n dose sobre el atrio derecho L os f asc i cul o s de la ve n a .

cava s uperior se origi n a n en el n odo y tabiqu e y rodea n a la ve n a ,

cava L a hoj a derecha del tabiqu e sec u n dario se ori g l n a en el

.

tabiqu e y se extie n de sobre la ve n a cava i nferior L a hoj a de .

recha del tabiqu e primario arra n ca del tabiqu e pasan do a las ,

V alvulas de la ven a cava i n ferior A trio izqu ierdo : L a cresta .

a n terior izqu ierda procede del tabiqu e y ba n da i n teratrial L a .

creta posterior del mismo lado procede tambié n de la ban da y

tabiq u e El f asc i cul o sep t op ul m o n ar se origi n a en el tabiq u e y
.

c u bre el techo del atrio L a hoj a izqu ierda del tabiqu e secu n .

dario se origi n a en cl tabiqu e exte n dié n dose sobre la sup erfi c i e ,

posterior del atrio El fasc i cul o sep t o at ri al arran ca del septo

.

exte n dié n dose sobre el techo del atrio L a hoj a izqu ierda de .

tabique primario n ace tambié n del septo pasa n do alrededor de ,

la base del atrio .

T ra ns l at i on by J o se F N o ni d c z
C o r nell U m vc rs x t y Me di ca l C o l le g e, N Y .
2 56 JAM E S w . PA PE Z

co mmen ces in this n ode They have also s uggested that t h e

.

excitatio n process S preads from the n ode thro u gh both right

an d left atria alo n g several m u sc u lar paths The right posterior .

crest ( tae nia termi n alis ) has bee n co n sidered the pri n cipal pri
m ary pathway for the excitatio n process i n the right atri um .

The i nterat rial ban d has bee n co n sidered the primary pathway
for the excitatio n to the left atri um ( G Bachman n It is .
,

the p u rpose of this paper to show the gross arran gemen t of the
mu sc u lar b un dles of the atria and to i n dicate how this arran ge
men t may serve to spread the excitatio n process thro u gh them .

M ATERIAL AND M E TH O D

For this work the method of clean in g an d strippi n g the mu s

c ul at u re has bee n fo u n d most u sef ul H uman beef and dog .
, ,

hearts i n ab un dan ce were u sed .

The cardiac o rifi c es were tied an d the hearts were distende d

with a 1 0 per ce n t sol utio n of formali n un der a pressure of abo ut
3 0 cm of the fl uid
. The fl uid was allowed to act for a day or
.

two d epen din g upo n the size of the heart The hearts were then
, .

washed i n runn in g water for a day an d tran sferred to alcohol .

To preserve the diste nded shape and facilitate extern al dis

section s the atria were the n st uffed with absorbe n t cotto n To
,
.

display the b un dles the epicardium an d en docardium were

removed .

S I N O -A UR I CU L A R NO DE AND SEPTAL R A P HE
From the preparation s at han d it i s clear that all of the pri n ci
pal mu sc ular b un dles of the atria radiate from on e cen tral area
which s urro un ds the o rifi ce of the s uperior ve n a cava This .

area is for the most part b uried i n the anterior part of the atrial
sept um b ut i n fro n t and to the right of the orifi ce of the ve n a
,

cava it comes to the extern al s urface The portio n that appears .

i n the groove betwee n the ve n a cava an d the right atri um has

bee n design ated the sin o auric ular n ode by Keith an d Flack
—

it is the seat of imp ulse form atio n for the atria i n the
n or m ally beati n g heart The portio n that is b u ried i n the atrial
.
 ATRIAL M US C ULATU R E 2 57

sept u m will be here called the septal raphe ; it provides an ap

pare n t mechan ical s upport for man y of the larger m u sc ul ar
b un dles of both atria My preparatio n s show that some of the
.

extern al b un dles commen ce i n the si n o auric ular n ode an d that —

other deeper b un dles comme n ce i n the septal raphe .

The sin o au ric ular n ode was fir st described by Keith an d Flack

—

They state that it co n sists of delicate pale an d pale ,

stai n in g primitive un di ffere n tiated striated mu scle fib ers plexi

, , , ,

form i n arran geme n t W ith well marked elo n gated n u clei The
—
, .

fi b ers resemble those of the atrio V e n tric u lar n ode of Tawara

-
.

They are embedded i n de n sely packed co nn ective tiss u e The .

fi b er s lead somewhat abr uptly i n to the s u rro un di n g atrial m u s

c u l at u re that forms the exter n al strata They men tion n erve .

termin als i n this n ode The sin o au ric ular n ode is recogn izable
.
-

as a thin strat u m of fin e pale m u sc ulat u re ben e ath the epicardiu m

i n the s ulc u s betwee n the s uperior ve n a cava an d right atri u m .

I have n ot bee n able to fi n d macroscopically any n erve filam en t

passin g to the sin o au ric ular n ode In co ntrast to this the
—
.

ab un dan ce of n erve filam en t s on the septal ( left ) side of the

o rifi c e of the s u perior ve n a cava si n ki n g i n to the regio n of the

septal raphe is si g n ifi can t Here those authors have described

.
,

t iss u e of the same character as forms the s i n o au ric u lar n ode —

.

Moreover they believe that the tiss u e of the atrio ven tric u lar
,
-

n ode of Tawara represe n ts a part of the same ri n g of u n d i ff er

en t i at ed m u scle tiss u e that s u rro un ds the si n o a u ric u lar j u n ctio n -

.

E xperime n tal work has c o n firm ed the View that the sin o
au ric ular n ode is the seat of imp ulse formatio n i n t h e n ormally
beatin g heart That the excitatio n process for at least the
.

sup erfi c i al b u n dles of the rig h t atri u m an d for the i n teratrial

ban d comme n ces i n the head of this n ode has bee n amply demo n
st rat ed by W yb au w T L ewis a n d his associates
.

a n d has bee n c o n fi rm e d by Eyster an d M eek

Several extern al b un dles commen ce i n the si n o au ric ular n ode -

regio n : the extern al bu n dles that cover the u pper an d lateral

s urface of the right atriu m (fi g 3 the circ ular b un dles of the
.
,

s uperior ve n a cava ( fig s 3 and 5 6a) and the i n teratrial ban d

.
, ,

( fig 5
.
,
The delicate fi b ers of the n ode cann ot be regarded
as givi n g these b un dles mechan ical s upport .
2 58 JA ME S w . PA P E Z

The septal raphe (fi g s 1 an d 2 R ) is the portio n of the atrial

.
,

sept um i n fro n t of the oval fossa i n which most of the large mu scle
b un dles of both of the atria commen ce I have design ated it .

the septal raphe becau se the term s u ggests that this is the seam
alo n g which the atria are k n itted together It is best seen o n .

the i nter nal s urface of the atria when the e ndocardiu m h as bee n
removed b ut its anterior margin appears i n a vertical groove
,

o n the a n terior s u rface of the atria below the i n teratrial ban d

as show n i n fi g ure 5 .

The septal raphe as shown in fi gures 1 and 2 R forms an

, , ,

irregular lin e i n the septu m that extends from the fro n t of the
atrioven tric ula r n od e of Tawara to the fro nt of the s u perior
’

ve n a cava where it t u rn s to the right to j oi n th e head of the s in o

auric ular n ode This co n tinuity is alon g the extern al or anterior
.

s u rface of the right posterior crest O n the right s ur face of

the sept u m ( fig 1 ) the prin cipal portio n of the raphe is abo ut
.

cm . above and i n fro n t of the atriove n tric u lar n ode regio n

a n d is covered by sup erfi c i al strata of right posterior crest ( 4)

an d right leaf of the sept u m sec un d um which have a lower

an d variable origi n These strata have a variable origin i n
.

di ffere n t hearts an d obsc ure the raphe From the right side .

of the raphe arise the right an terior crest right posterior

crest right leaf of the sept u m sec un dum ri ght leaf of
the sept u m primum sep t o c o ron ary b u n dle i n t erc av al b un dle
,

an d the left radiate b un dles of the s u perior ve n a cava ( 6b) .

O n the left s urface of the sept u m ( fi g 2 ) the raphe as seen from

.
,

the inn er s urface exten ds alon g a lo n g s shaped lin e The upper

,
-
.

en d of it is sit u ated j u st behi n d the i n teratrial ba n d which is ,

o n the an terior s u rface of the atrial j un ctio n T hi s portio n of

.

the raphe gives origin to the sept opulmon ary b un dles ( 1 2 ) and
the u pper portion of the i nt ercaval b un dle (5) an d the left leaf
of the sept u m sec un d um The middle portion of it is
sit u ated i n front of the oval fossa adj acen t to the prin cipal
portion on the right side an d gives origin to the large left septo
atrial b un dle The lower portio n of it is situ ated below
the oval fossa opposite the region of the atrioven tric ular n ode
close to the j un ction of the atrial and ven tric ular septa an d gives
origi n to the large left leaf of the sept um prim um
2 60 JAM E S w . PA PE Z

the sin o a u ric ular n ode is abolished is the seat of imp ulse form a
-

tio n for the atria t ran sferred to the atriove n tric ular n ode .

By the m ethods employed I have n o t been able to determin e

what co nn ectio n s exist between the septal raphe an d the sin o
a uric u lar an d atriove n tric ular n odes It is con ceivable that .

rem n an ts of sin o au ric u lar tiss u e exist i n the raphe or that the
—

b un dles that arise fro m the raphe are d irectly co nn ected with
both of the n odes by mean s of similar tiss u e It is clear that .

the septal raphe co n tai n s fib rou s tissu e which serves as a poi n t

of attach m e n t for the m u scle b un dles that arise i n the sept um .

The qu estio n is raised whether or n ot the pacemakin g f un ctio n

for the b un dles of the left atri um an d the deep b un dles of the
right atriu m exists i n the septal raphe If n ot what 18 the .
,
’

relatio n of these b un dles to the head of the sin o au ric ular n ode -

o r how is the excitatio n process co n d u cted to the m ?

.
If this
fun ctio n does exist i n the septal raphe what is the relatio n of ,

th e r a p h e to the atriove n tric u lar n ode of Tawara ?

M US C ULATURE O F TH E RI G H T ATRIUM
1 . The i n tera tri a l ban d

The i n teratrial ( i n tera u ric ular ) ban d as show n i n fi g u re 5 1

’

, , ,

is a stro n g exter n al m u sc ular b un dle o n the an terior s u rface of

the atria It exte n ds from the si n o au ric ular regio n to the left
.
-

appe n dage It is represe n ted i n Toldt s A tlas fi g u r e 9 58 Keith ’

“
.
.
,

an d Flack design ated it a co n stan t ba nd passi n g fro m t h e

si n u s m u sc ulat u re to the vestib u le of the left a u ricle
” T L ewis . .

an d others have adopted this n ame for i t L ewis fo u n d .

,

i n traci n g the ex c i t at at i o n from the si n o a u ric u lar n ode that the—

,

rate of propagatio n w a s greater alo n g the ban d than elsewhere

.

i n the atriu m G Bachman n

. . showed that altho u gh , ,

perhaps n o t the o nly it is the primary or more direct path for

,

the co n d u ctio n of the excitatory process from the right to the

left au ricle .

This ban d (fi g 5 1 ) lies o n the extern al an d a n terior s u rface

.
,

of the a tria i n fro nt of the left sept opulm o n ary b un dle

‘
which
for ms the deeper an d m ore exte n sive strat um i n this regio n .
 AT R IAL M US C ULATU R E 2 61

It arises by its deep s urface from the head of the si n o au ric ular -

n ode i n fro n t of the o rifi c e of the s u perior ve n a cava This .

origin exte n ds alo n g a diago n al li n e the left en d of which termi

,

n ates i n the atrial sept u m The b ulk of the fi b ers arises n ear
.

the sept um below the ban d Some of the b un dles arise above
.

the head of th e n ode on the fro n t of the ven a cava an d are dis
t i n ct ly traceable i n to the fi b er b un dles of the n ode These .

u pper fi b er b u n dles from the u pper part of the ban d exte n d to

, ,

the left atri um an d form a small part of the left posterior crest
(fi g 4 1 1 ) e n circlin g the base of the left atrial appen dage Be
.
,
.

hin d them is the large left sep t op ulm on ary b un dle (fi g 5 .
,

The larger portio n of the fi b er b un dles that arise from the head
of the sin o au ric u lar n ode alon g the diagon al lin e exten ds to the
-

fron t of the left atrial appen dage an d divides to en circle the

appen dage midway between the tip an d t h e b ase Here it .

prod u ces a sharp co n strictio n cau sin g the left atrial appen dage
,

to be ben t u pward The right end of the ban d exten ds in to the

.

fro n t of the right atrial appen dage and divides to en circle it n ear
its base i n a mann er similar to that on the left side b ut the ,

co n strictio n is slight an d m ay be wanti n g In the left atri um .

it gives origin to the middle or apical gro up of pecti n ate mu scles .

Both en ds of the ban d give origi n to a thin sup erfi c i al strat um

of fi b er b u n dles that e n circle the atrial appe n dages O n the .

right side this strat um is well d efin ed on the left side it is sparse ,

an d ofte n abse n t so that the d eeper pecti n ate strat u m is disti n ctly

seen on the s urface This is the case i n the dog heart o n both
.

sides so that the deeper pectin ate strat u m is disti n ctly sup erfi ci al .

The e n circlin g fi b ers which arise i n the co n ce n tration area ‘ ’

(fi g 3 2 ) of T L ewis i n the dog heart mu st n o t be co n fu sed with

.
,
.

sup erfi c i al circ u lar strat u m fo u n d i n the h u man an d bovi n e

hearts The sup erfi ci al fi b ers are seen well i n beef hearts
. .

2 . The ex tern a l bu n dles of the ri ght a tri u m

T he extern al b un dles ( fig s that e n circle the right

. 3, 2, an d 5, 3 )
atriu m are derived from the septal raphe head an d tail of the ,

n ode an d from the in t er cav al b un dle

,
In the h u man heart they .
2 62 JAM E S W . PA PE Z

fo rm a thin more or less un iform strat um over the s urface of the

,

right atrial appen dage an d right s u rface of the base of the atri um .

In the bovi n e heart they are stro n gly developed In the dog .

an d cat hearts they form a thi n strat u m more di ffi c ul t to di ffer

en t i a t e
. O wi n g to their diff u se origin an d termi n atio n they ,

can n ot be acc urately described The followin g descriptio n is

.
,

therefore an approx imate on e from which there are variatio n s

,
.

a . The origi n of those from the septal raphe is below the right
limb of the in teratrial ban d It is covered by the origin of the
.

ban d as seen o n the extern al s urface in fi g ure 5 3 They pass ,

.

forward toward the atrioven tric ular rin g and are n othi n g more
than the sup erfi ci al strat um of the right an terior crest They .

are spread o ut and in serted i nto the atriove n tric ular rin g t o the
right of the aortic o rifi ce .

b The origi n of those from the head of the n ode is hidden by

.

the right limb of the in teratrial ban d (fi g 5 They pass .

,

forward an d e n circle the base of the right atrial appen dage O n .

the lateral side of the atrium they divide in to two limbs O ne .

of these en circles the base an d forms a thin strat um over the

lower an d right side of the atrium The other sweeps upward .

aro un d the base of the atrial appen dage coverin g the middle
clu ster of the an terior pecti n ate m u scles and forms a distin ct ,

con strictio n i n the lower lateral part of the base of the atrial
appen dage which is seen especially well i n dog hearts as shown ,

i n fi g ur e 3 .

Those that arise from the tail of the n ode form a forward
'

0 .

and a lateral rad iatio n an d a posterior radiatio n ( fi g 3 The .

,

forward radiation is d istin ct on ly i n the dog heart T L ewis . .

has called it the co n ce ntratio n area It aris e s from the fro n t

‘
.
’

of the tail of the n ode and passes alo ng the upper margin of the
right atrial appe n dage Together with the right limb of the
.

i nteratrial ban d it gives rise to the extern al circ u lar b un dles of

,

the right atrial appen dage It covers the in terlaceme n t of the

.

apical pecti n ate mu scles divides and is diffu sed over the lateral
, ,

s urface of the right atrial appendage In the human heart this .

radiation is gen erally n ot recog n izable b ut i n some in stan ces a ,

sm all disti n ct sup erfi ci al b un dle abo u t 1 cm i n len gth occ urs i n .
2 64 JAM E S W . PA P E Z

u lar ri n g givi n g rise to the right an terior pecti n ate m u scles

,
.

Its a n terior or lower margi n is attached to the thin fib rou s atrio

ve n tric ular ri n g Between the origin s of the right an terior an d
.

posterior crests there is formed a thin spot The right an terior .

crest e n circles th e base of the right atri um an d n ear the mou th

of the coro n ary si nu s i n terlaces W ith the termin al pectin ate
col um n s of the right posterior crest From here they are c on .

t i nu ed as a thi n strat u m to the atriove n tric u lar n ode The right .

an terior pecti n ate m u scles ( fig 7 ) nu mber n i n e to fou rtee n

. .

T he o n es i n the apex of the right atrial appe n dage are large .

The others are smaller They arise from the an terior crest
.

si n gly or i n three radiati n g cl u sters T he trun ks of the pecti n ate

.

m u scles bran ch o ut an d the bran ches in terlace with those of the ‘

posterior pecti n ate col umn s The u pper pecti n ate cl u ster arises
.

as a stro n g col um n from the an terior portio n of the atrial sept u m

i n fro n t of the origi n of the posterior crest Its pectin ate m uscles .

exte n d toward the apex of the atrial appen dage The middle .

cl u ster is u s u ally d efin i t e It forms an extern al con striction

.

below the base of the atrial appen dage Keith an d Flack .

design ated the an terior crest as the annu lar fi b ers of the au ricle
‘ ’

an d M c M u rri c h i n Pi erso l s A n atomy speaks of them as an siform

’

fascic uli
.

F rom the experime n ts of T L ewis M eakin s an d Wh ite it

.
, , ,

wo uld appear that the excitatio n process is con d u cted by the

extern al b un dles of the atriu m from the si n o au ric ular n ode in to —

the an terior pectin ate m u scles an d the n ce i n to the an terior crest .

However the fat an d vessels i n the coron ary s ulc u s raise the
,

epicardi um from the an terio r crest to s u ch an exte n t that direct

applicatio n of electrodes can n ot be made .

4 The ri ght p osteri or

. cres t

The right posterior crest ( fig s 7 4 and 1 4) is a stro n g mu s

.
, , ,

c ul ar col um n called by His the termi n al crest becau se it forms

, ,

the bo un dary between the sin u s ven os u s an d the right atriu m .

It is seen on the i ntern al s u rface of the atri um and gives rise to

the posterior set of right pecti n ate m u scles Extern al b un dles .
 ATR IAL M US C ULATUR E 2 65

derived directly from the si n o au ric u lar n ode e n ter i nto its —

compositio n The in tern al bu n dles however form a large part

.
, ,

of the crest They arise ( fig 1 R 4) i n the an terior part of the

. .
—

, ,

atrial sept um from the septal raphe They c u rve u pward an d .

to the right i n fro n t of the orifi c e of the s uperior ven a cava an d

the n dow n ward o n the right of both ven ae cavae They are .

overlaid by the sin o au ric ular n ode A lo n g its co urse the crest
-
.

gives rise to abo u t t en posterior pecti n ate mu scles ( fig an d .

as these are given o ff the crest diminishes i n size O n e or .

two upper pecti n ate mu scles are large an d exte n d in to the atrial
appen dage The s u cceedin g on e s are progressively smaller In
. .

the wall of the atriu m the pectin ate mu scles divide in to n umerou s
bran ches that i n terlace with those of the an terior pectin ate
mu scles The lower pectin ate mu scles are small an d exten d to
.

the lower right part of the right atrioven tric ular rin g The .

lowermost on e e n circles the orifi c e of the coro n ary si nu s an d is

j oin ed by the right leaf of the sept u m primu m ( fig 7 8 ) which .
,

exten ds throu gh the ( right ) valve of the in ferior ven a cava .

The extern al b un dles have already been described un der the

extern al b un dles of the right atri um The u ppermost o n es run .

parallel to the sin o auric ular n ode an d commen ce in it as att en u

-

ated fi b ers They spread out with the pectin ate mu scles which
.

they cover extern ally Below they are rei n forced by the i nter
.
,

caval b un dle which spreads down ward to the lower part of the
right atrioven tric ular rin g .

Keith an d Flack co n sidered that the right posterior

“
crest probably belo n gs to the sinu s ve n osu s sin ce it exten ds ,

ben eath the en docardium of the atrium from the positio n of on e

ve n o us valve to the other ” In the same artic l e however they
“
.
, ,

state that the auricles ( atria ) are o utgrowths from the dorsal
wall of the auric ular ( atrial) can al ” The latter seems to be the .

best View Hen ce the an terior an d posterior crests may be

.
,

regarded as derived from the ann ular mu sc ulat u re of the atrial

can al A s the crests become separated the in terven in g mu s
.

c ul a t ure becomes arran ged i n the form of an an terior an d poste

rior set of pectin ate mu scles .

2 66 JAM E S w . PA PE Z

The electrocardiographic res ults of T L ewis M eakin s an d .

, ,

White show that the excitatio n process spreads from the

sin o auric u lar n ode to the right atrium alo n g the superfi ci al
-

m u scle b un dles an d alo n g the posterior crest The mass c on .

tractio n of the atri um is eviden tly due to the con tractio n of the
pecti n ate m u scles and their co nn ectio n s with the septal raphe
thro u gh the anterior an d posterior crests .

M US C ULATURE O F TH E SINUS V EN O S U S

5
. The i n terca va l bun dle

The i n t er cav al b un dle ( fig s 3 an d 6 5) is a large fiat on e that

.
,

passes obliqu ely i n the posterior fi b rous wall of the si n u s v en os us .

It arises (fi g 7 5) i n the fro n t an d left side of the orifi ce of the

.
,

s uperior ve n a cava from the head of the s in o auric ular n ode an d -

from the u ppermost li mit of the septal raphe and i n man y cases
a large slip arises i n fro n t of the orifi c e of the right u pper pul
m o n ary vei n as show n by Keith an d Flack ( O7 fi g 5 f )
,
The’
,
.
,
.

u pper margi n of the i n t er c av al b u n dle is i n series with the b un dles

that e n circle the orifi c e of the s uperior ve n a cava the lower ,

margin is i n series with the right leaf of the sept um sec un d um

( fig 1
.
,
It c u rves obliq u ely backward an d down ward and
to the righ t i n the fi b rou s posterior wall of the sinu s ven os u s .

It forms an obliq u e co n striction betwee n the orifi ces of the ve n ae

cavae i n the regio n of t u bercle of L ower Crossi n g the sin o .

au ric ular j u n ctio n and right posterior crest it S preads out as a ,

fi b ro us expa n sio n over the extern al s u rface of the lower portio n

of the right atri um where it forms a sup erfi ci al strat um A .

portio n of it covers the posterior and right side of the ori fi ce

of the in ferior ve n a cava as shown i n fi g ure 6 5 Where it
, ,
.

crosses the tail of the n ode it may receive accessio n s from the
n ode Its i ntimate attachme n t to the posterior crest and the
.

n ode re n der its relatio n to these diffi c ul t to determi n e Some .

of its b un dles overlap the right s urface of the o ri fi ce of the i n ferior

ve n a cava an d are i n serted i n to an extern al fi b rou s sheath The .

more proxim al of these pass aro un d the o rifi ce to the regio n of

the coro n ary si n u s The exte n t to which this m u sc ulat ure
.
2 68 JAM E S w . PA PE Z

fi g u re 3 , 6a , there is sometimes a disti n ct b un dle that arises

obliqu ely from the n ode an d sweepi n g u pward over the right
,

s urface of the ve n a cava divides to en circle the vessel In th e

,
.

dog heart the m u sc ulat u re of the s uperior ve n a cava is qu ite

exte n sive an d s urro u nds also the orifi ce of the azygos vein
“
.

Eyster an d Meek believe they have shown that the excita

tio n u s u ally reaches the i n t erc av al regio n an d the s uperior ve n a
cava before an y other regio n s ” The electrocardiographic res ults
.

of T L ewis Meak i n s an d White

.
, ,
i n dicate that there is n o
mate rial di fferen ce of the rate of co n du ctio n from the sin o
au ric u lar n ode to the ven a cava They have show n that the .

excitatio n process i n the dog s heart exten ds radially upward

’
.

The variable rate which they obtai n ed is d ue as they s u ggest , ,

probably to the obliqu ity an d S piral arran gemen t of the ven a

caval mu sc ulat u re .

The radial b un dles (65) appear partic u larly in the left side and ,

i n some hearts also i n the posterior side of the s u perior ve n a ,

cava They are covered by the circ ular set The left on es
. .

arise from the u ppermost limit of the septal raphe (fi g 5 6b) .

,

an d at the o rifi c e of the ven a cava and radiate i n an u pward an d

posterior directio n In the wall of the h um an an d dog hearts

.

they form an exceedin gly thi n strat um The posterior radial .

set of m u scle b un dle s is comm o n ly absen t i n the hu man heart .

When presen t they are feeble They arise from the tail of the
,
.

si n o auric ular n ode and sweep diago n ally u pward and to the
-

left over the posterior s urface of the ve n a cava which they te n d

to s urro un d i n a circ ular directio n ( fig . They are sit u ated
j u st above the i nt ercaval b undle from which they are diffic ult ,

to dis ti n guish The posterior s u rface of the s uperior ve n a cava

.

is more extens ive than the an terior on accoun t of the obliqu ity
with which it j oin s the sinu s ve n os u s E xcepti n g the circ ular .

b un dles and i n some i n stan ces; the posterior radial b un dles

, ,

it is fi brou s between the ve n a caval m u sc ulat ure an d the obliq u e

i n t erc av al b un dle .

The bun dles arou n d the i nferi or ven a ca va The i nf erior ve n a .

caval b un dles (fig s 4 6 and 7 ) are derived from the sept um

.
, ,

primum (8 ) an d from the i n t ercaval b un dle They exten d .

 ATR IAL M US C ULATURE 2 69

down ward over the ven a cava b ut do n ot form cir c ular b un dles ,

aro un d i t The right side of the ori fi c e is limi ted by the right
.

posterior crest The lower portion of the in t erc aval b un dle (5)
.

spreads down ward over the posterior and right s u rface of t h e

in ferior ven a cava and termi n ates i n a fibrou s expan sion The .

u pper portio n of the right leaf of the sept u m prim u m ( fi g 7 8 ) .

,

spreads over the left and posterior sides of the in ferior ven a
cava an d termi n ates i n a fib rou s expan sion The posterior .

s urface of the in ferior ven a cava between the right leaf of th e

septum prim um and the i nt erc aval b un dle is u su ally fi b ro us .

Th e lower portio n of the right leaf of the sept um prim um forms

a stron g ban d i n fro nt of the orific e of the in ferior ven a cava
alon g the bas e of the ( right ) valve of t he in ferior ven a cava .

Thi s frequ en tly forms a sharp fold between the orifi c e of the
in ferior ven a cava and that of the coro n ary s i nus .

Keith and Flack fi g 3 ) have ill u strated the b un dles on

.

the dors al s urface of the in ferior ven a cava .

7 . The r i ght leaf f the sep tum

o secu n du m

The septum sec un dum (fi g s 1 7 2 1 3 and 7 7 ) or lim bu s of .

, , , , ,

the oval fossa is a stron g musc ular b un dle whi ch arises i n the
septal raphe (fig 2 R ) chi efly on the right side and arches
.
, , ,

backward over the oval fossa It is divisible i n to a right an d .

a left leaf b ut the separatio n of the leaves is to some extent an

,

arbitrary matter .

The right leaf of the sept um sec un d um (fig s 1 7 an d 7 7 ) .

, , ,

belon gs to the sinu s ven osu s It arches over the left side of the.

o rifi c e of the in ferior ve n a cava above an d posterior to the o rifi c e ,

of the coro n ary sinu s There it is in serted i nto a fi brou s ex pan

.

sio n over the in ferior ve n a cava A bove it is i n series with the .

,

i n t erc av al b un dle b u t it is sit u ated deeply i n the atrial j un ctio n

,
.

A portio n of this b u n dle t u rn s to the left o n to the o rifi ce of the

coro n ary sin u s .

TH E AM E R I C AN J OU R N A L OF ANA O M Y V O L
T , 27 , NO 3
 JA ME S w . PAP E Z

8 . The ri ght leaf f

o the sep tu m p ri mum

The sept um pri mu m (fi g s 2 8 and 7 8 ) is a large b undle of

.
, , ,

m u scle tiss u e that form s the lower bo un dary of the oval fossa .

It is j oin ed by fib rous tissu e to the ven tric ular septu m It is .

divisible i n to two distin ct portion s the right leaf for the sin u s ,

ve n os u s and the large left leaf for the base of the left atrium .

The two leaves have in depen den t ori gin s an d are almost en tirely
separate Sin ce the left leaf belo n gs to the left atrium o n ly
.
,

the right leaf will be described here .

The right leaf of the sept um prim um (fi g 7 8 ) arises from .

,

the septal raphe on the right side i n fro n t of the oval fossa and ,

passes down ward and to the right i n the ( right ) valve of the
in ferior ven a cava that exten ds alon g the fron t of the orific e
,

of the vessel and in termin gles with the termin al expans io n s

,

of the right crests and i nt ercaval b undle Thi s leaf passes .

,

therefore between the orifi ces of the in ferior ven a cava an d

,
'

coro n ary sinu s to both of which it co n trib utes mu sc ulat ure

,
.

Some of its bun dles sweep u pward and form the posterior border
of the oval fossa .

The lower portio n of this b un dle that may be called the ,

s ep t o c oro n ary b un dle arises from the lowermost portio n of the

,

septal raphe an d spreads out i n a fan shaped mann er i n fron t of -

the coro n ary sin u s coverin g the right side of the atrioven tric ular
n ode (fi g 1 A V N
.
,
. The b undles that form its right s urface
.

appear to be attached to the fib rou s upper exte n t of the in ter

ven tric ular sept um b ut its deeper b un dles sta n d i n i ntimate
,

relatio n to the atriove n tric ular n ode T L ewis has shown that . .

mu sc ular co ntraction i n this regio n appears early as is i n dicated ,

by an early e x trin sic defl ect i on on the cardiogram E yster an d .

Meek have shown that thi s regio n receives the excitatio n process
early before it has spread fully i nto the right atrium .

Keith and Flack have shown that the mu sculatu re of

the ven o u s valve which is here called the right leaf of the sept um
,

prim um is formed by an i n foldin g of the atrial wall at the sin o

,

auric ular j un ction To the right this mu sc ulat u re S preads out

.

an d j oin s f reel y with that of the ri ght atriu m .

272 JAM E S w . PA PE Z

Si n ce the left an terior crest is i n large part a co n ti nu atio n of

the i n teratrial ban d it is clear from the experime n ts of Bach
, ,

man n that the excitatio n process reaches this regio n of the

l ett atri um chiefly alo n g the ba n d .

11 . The left p osteri or cres t

The left posterior crest ( fig s 5 an d 8 1 1 a 1 1 5) is a broad .

, ,

b un dle the s uperficial portio n of W hich (fi g 5 1 1 a ) is a c on

,
.
,

t i n ua t i o n of the u pper part of the i n teratrial ban d an d the deep ,

portio n of which ( fi g 8 1 1 ) arises from the sept u m i n series

.
,

with the sep t oat ri al b un dle It passes to the left aro un d the
.

base of the left atrial appe n dage i n fron t of the orifi ces of the
left p u lmo n ary vei n s It gives origin to the small posterior set
.

of pecti n ate m u scles Posteriorly it i n terdigitates W ith the left

.

an terior crest formin g the trian g ular area which is attached to

,
,

the i n ferior margi n of the left atriove n tric ular ri n g It lies .

i n fro n t of the sep t op u l m o n ary ( fi g 8 1 2 ) an d sep t o at ri al (fi g .

,
.

8 1 4) b un dles an d forms a disti n ct co n strictio n aro un d the base

,

of the left atrial appe n dage The left posterior pectin ate m u scles
.

are several i n n umber Their bran ches i n terlace with those of

.

the an terior set The left atrial appen dage is a n arrow forward
.

prolo n gatio n of the left atri um The left extremit y of the i n ter .

atrial ban d exte n ds alo n g the a n terior margi n an d midway ,

betwee n the base an d apex it divides and embraces the appe n dage
This gives rise to an immediate set of pecti n ate m u scles that
o cc u py a s u perior positio n In the h uman heart the posterior
.

set of pecti n ate m u scles is above the an terior set b ut i n the beef ,

an d dog hearts they are posterior For morphological reaso n s .

this design ation has bee n retai n ed for the h uman heart .

Bachmann has i ndicated that the spread of the excitatio n

process to the left atrial appe n dage is alon g the in teratrial ban d .

This forms the sup erfi ci al strat um of the left posterior crest .

The deeper strat um however arises from the septal raphe

, ,
.
 AT R IAL MUS C ULATU R E 273

12 . S ep top ulmon ary bun dles

S ep t o p ul m on ary b un dles (fi g s 4 6 an d 8 ) are the large flat

.
, ,

extern al b u n dles that cover the u pper an d posterior s u rface of

the left atri um betwee n the p ulmo n ary vei n s an d e n circle the
o ri fi ce s of the vei n s They arise deeply from the left s u rface
.

of the septal raphe (fi g 2 1 2 ) behin d the in teratrial ban d an d

.
,

c u rve upward an d to the left chi efly i n fro n t of the orifi c e of

,

the u pper right p ulmon ary vein an d above the i n teratrial ban d
, ,

where they come to the extern al s u rface They spread out as .

six ban ds over the u pper an d posterior s u rface of the left atriu m
,
.

The an terior sep t o pul m on ary b u n dle ( 1 2 a) arches above the

i n teratrial ban d exten ds to the left i n fro n t of the left s uperior
,

p u lmo n ary vein as the sup erfi ci al strat u m coverin g the left
posterior crest Below the o ri fi c e of the vei n i t curv es backw ard
.
,

an d su p erfi c i ally over the back of the atri u m betwee n the left

i n ferior p u lmo n ary vein an d the coro n ary sin u s It covers the .

left leaf of the sept u m prim um .

The left i n t erv en ou s sep t op ulm on ary b un dle ( 1 2 h) exten ds

u pward as a deep strat u m of the foregoi n g an d to the left betwee n

the left p u lmo n ary vein s an d j o in s the an terior sept opulm on ary
b un dle .

The left posterior sep t opulm on ary b un dle ( 1 2 0 ) arises with

the foregoi n g an d spreads backward an d to the left over the left
atri u m between the left an d right p ulmo n ary vein s Behin d .

the left lower p ulmo n ary vein it spreads o ut i n to the an terior

sep t op u l m o n ary b u n dle an d c u rves to the left below the lower

left p ulmo n ary vei n Some of the fi b ers of the foregoin g b un dles
.

exten d o n to the left p ulmo n ary vein s an d give origin to the fi b er

b un dles that en circle the vein s i n a radiate an d somewhat circ u lar
mann er .

The right posterior sept opulm on ary b un dle ( 1 2 d) like the ,

other exte n ds backward over the upper s urface of the left atrium
,
.

Spreadin g o u t over the posterior s urface of the atrium it c u rves ,

to the right below the right lower p ulmon ary vein It overlies .

the left leaf of the sept u m sec un d u m an d sept um primum A .

portio n of it exte n ds to the back of the in ferior ven a cava bridgin g

over the atrial j un ction in terlacin g with the septal b un dles
,
.
 JAM E S W . PA P E Z

T he right i n t e r v en o u s b un dle ( 1 2 e) arches

s ep t o p u l m o n a r y

over the o ri fi ce of the right u pper p ulmo n ary vei n Betwee n the .

vei n s it t urn s d ow nward toward the atrial j un ctio n it spreads

,

over the left leaf of the sept u m sec un d u m T his an d the forego .

i n g b un dle give rise to b u n dles that e n circle the o ri fi c e of the

lower right p ulmon ary vein .

The right sep t opu lm on ary b un dle (fig 1 1 2f ) arises i n the septal .
,

portio n of the n ode an d exten ds u pward on the right wall of the

(

u pper right p u lmo n ary vei n an d c u rves dow n ward alo n g it s

right wall It gives rise to circ ular fi b er b un dles that s u rro un d

.

the o ri fi c e of the u pper ri g ht p ulmon ary vein It m ay be re .

garded as the u ppermost portio n of the left leaf of the sept u m

sec un d um With the right i n t erv en ou s b un dle it gives origi n
.

to an obliq u e ban d that exten ds to the right on to the posterior

s urface of the i n ferior ve n a cava bridgin g over the atrial j un ctio n .

This m u sc ular bridge ( fig 6 I Ze) is n ot always presen t an d m ay

.
,

be fi b ro u s .

Keith and F lack have fi g ured these b un dles They .

co n sider them a part of the i n teratrial ban d The origin of these .

b un dles from the septal regio n behi n d the ban d however is , ,

clear ( fig 2
.
, It is also appare n t that they belo n g to the
same sup erfi ci al strat um as the ban d (fi g 5 .
,

13 . The left leaf f

o the sep tu m secun du m

The left leaf of the sept u m sec un d um (fi g s 2 an d 6) comme n ces .

i n septal raphe (fi g 2 1 3 ) an d exte n di n g backward forms the

.
, , ,

right or septal wall of the left atri um i n fro n t an d below the

o ri fi c es of the right p u lmo n ary vei n s Below the lower vei n .

it t u rn s to the left sup erfi c i ally to the left leaf of the sept um
,

prim um (fi g 2 1 5) W ith W hich it i n termi n gles an d becomes

.
,

spread over the posterior s urface of the left atri um ( fi g s 4 an d 6 .

,

14 . The left sep toatri a l bu n dle

The left sept o at ri al b un dle (fig s 2 an d 8 1 4) is a large flat

.
,

b un dle on the inn er s u rface of the left atrium It rises i n the .

septal raphe i n fro n t of the oval fossa an d below the origin of

27 6 JAM E S w . PA PE Z

crests Its lower m argi n is attached to the atriov entric ular

.

ri n g where it appears o n the extern al s u rface an d gives off some

,

lo n git udi n al b un dles to the coro n ary si nu s (fi g 4 The .

,

s ep t o p ul m o n ary b u n dles overlie a n d i n termi n gle with its u pper

margi n The left sep t oat ri al b un dle ( fig 6 1 4) j oi n s the deep

. .
,

portio n of its u pper margi n with which it appears to be co nti nu o u s .

Keith an d Flack have fi g u red this b un dle on the posterior

s u rface They co n sider it as a portio n of the ann ular b un dles
.

of the atrial can al .

E xperime n ts to determi n e primary n egativity over this regio n

an d over the posterior an d s u perior wall of the left atri u m have

bee n n egative (Eyster an d Meek ) It is clear that there is n o .

imp ulse formin g tiss u e i n this regio n .

A NO TE O N TH E N E R V E S U PPL Y O F TH E ATRIA
The atria are s u pplied by both the right an d left v ag osym p a
thetic n erves a O n the right side u s u ally three n erves r each
. .

the heart These pass dow n i n fro n t of the right p ulmo n ary
.

artery The lower two un ite above the right p u lmo n ary vein s
.

an d termi n ate i n a ga n gliform e n largeme n t i n the sept u m o n the

left side of the o ri fi c e of the superior ve n a cava From this .

termi nu s bran ches s upply the i n teratrial ban d i n t ercav al b un dle

, , ,

an d other b u n dles that radiate from the septal raphe O n e or .

two slen der fil am en t s pass to t h e left an terior crest alo n g the

left limb of the i n teratrial ban d .

I) The left vagosympathetic cardiac n erves term i n ate sim i

.

l arly i n the left atri um above the base of the left atrial appe n dage
an d the upper p u lmo n ary Vei n F il am en t s from this termi n u s
.

s upply the left an terior an d the left posterior crests septo ,

p u lmo n ary an d left sept oat r i al b un dles A small filam ent may
,
.

pass to the right in the in teratrial ban d as far as the sept u m .

M u ch of the literat ure that deals with the s ubj ect is con cern ed
with experime n tal work The fi g ures of Q u ain Porier and
.
,

Sharpey Toldt Keith an d Flack an d Sp al t eh ol t z show extern al

, , ,

Views of the atrial mu sc ulat u re .

 ATR IAL M US C ULATU R E 27 7

BIBLI O G RAPH Y
B A C H M A NN 1 91 6 Am . J o u rn . f P hy i l
o l 41 s o .
, vo .
, p . 30 9 .

EY A ND M K
S TE R EE 1 9 14 H t e ar , v ol 5, p 1 1 9
. . .

G A NTER A N D Z A H N 191 2 A hiv f

rc . d Ges h . . P y i l Bd s o .
, . 1 41 , S 3 35 . .

K I H A N D F A CK
E T L 1 90 7 J o u rn . of A n at . an d P hy i l s o .
,
v ol . 41 , p . 172 .

L WI
E S 1910 1 1 —
H e art , v o l . 2, p . 23 .

L WI
E S, ME A KI N S A N D W HI TE Phi l os . T r an s a c t . of R . S . of L d
on o n , s er . B, v ol .

20 5, p . 375 .

L WI A N D O N H I M R 1 9 10 H
E S PPE E E e ar t , v o l .2 , p 1 47 . .

M K A N D EY S TER 1 9 1 4 H
EE t e ar , v ol . 5, p . 227 ; A m J o u rn . of P hy i s ol .
, v ol . 3 4,
p . 368 .

S CH L O M O VI T Z , EY R A ND M
S TE EE K 1915 Am . J o u rn . of P hy i s ol .
, v ol . 37 , p 1 77 . .

S P A L TE H O L Z an H d Atl
as , v o l 2 , fi g s 41 3 , 41 4 . . .

T D
OL T A
t l a s fi g s 9 58 , 959
,
. .

W Y BA U W 1 9 1 0 rc h A
n t e rn a t d . I
h s ol , T 1 0 , p 7 8 . . P yi . . . .
 PLATE 1

EX LANA I N
P T O OF F I G UR ES

1 B dl
un es t h at ra di at e f m th pt
ro e se al r a ph e i n t he r i g ht f
s u r ac e o f t he a t rl a l

se pt um o f a h u m an h e a r t . The en d o c ar d i um h as b een re mo vde .

2 B dl
un es t h at ra di at e f rom t he se pt al r a ph e i n t he le t f f
s u r ac e o f t he i
a t r al

se pt um of a h u m an h e ar t . T he en d o c ar di u m h as b een re m o vde .

N i nt i b nd
erat r al a i v nt i
atr o e r cul ar n o d e

NM e x t n l b u d l f i ght t i u m
er a n es o r a r t i v t i
a r o en r c u l ar r n i g
Q
Q O
\
r i gh t t i t
an e r o r c re s O S . .
,
n
c o ro y i u ar s n s

R
Q
r i ght p t i t
os e r o r c r es inf i v n er o r e a ca v a

i nt v l bundl M O mit l ifi
I
J
Q
M er c a a e . .
,
ra or ce
b
Ow s up i v
er or v l bu dl
en a c a a n es 0 F . .
, ov lfa o ss a

Q r i ght l f f p t u m u d m
ea o se s ec n u P A . .
,
p lmu o n ar y a r t er y
O
O v r i g h t l f f p t u m p i mu m
ea o se r P . M .
,
p tiec n at e m u sc l es
9, mu u l t u sc f th
a y i
re o e c o ro n a r s nu s P V . .
,
p lmu o n ar y v i
e n

1 0, l f t nt i
e a t
e r o r c r es B .
,
ra ph e

1 1 , le t f p i
o s t e r o r c re s t R A . .
,
r i gh t i
at r u m

12, s ep t o p u l m o n a ry b un d le i i
s n o —au r c u l ar n o d e

1 3 , l e t l ea f f o f se pt u m se c u n d um p i v
su er o r en a c a v a

f
1 4, l e t se p t o a t r i a l b u n l e d TO . .
, t i p id fi
r cu s OIl
’

CG

f f
1 5, l e t l e a o f s e t u m r m u m p pi Ve n t , v ti l
en r c e
 PLATE 2

EX LANA I N
P T O OF F I G UR ES

3 Ex t e rn al bu dln es o f th i gh t t i
e r a r um of t he d o g h eart , vi ew e d f ro m t he
ri ght id s e . In t h i h
s e art t he
‘
c o n cen t r at o n i are a
’
of T . L wi
e s w as i k i n gl y
st r

d ev l p ed
e o . An u nu s u a l b u n d le e xt den s o nt o t he r i ght f
su r a c e o f t he sup i
er o r

v en a c a v a .

4 Ex t e rn al b un d l es o f t he l e t f i
at r u m o f a hu m an h eart , vi ew e df
m t he l e t
ro f
s id e . T he vi s c era l p i
e r c ar d ium an d t he c o ro n ar y v v
essel s h a e b een r em o e vd .

T he l ef t at r a l ai pp d g
en a e is O p en e d al o n g i t s m ar gi n t o sh o w t h e n t e rm eiat e di
g ro u p of p i
e c t n at e m u sc l es .
AT R IA L MU SC U A T U R E
L
P LAT E 2

J AM ES P AP E
W . Z

S AH
.

IV O
. .

S VC. .
 PLATE 3

E X LANA I N
P T O OFF I G UR ES

5 Ex t b u n d l es seen o n t h e a n t e r o r s r a c e o f t h e h u m an h eart
e rn a l i uf . Th e
v i sc era e r c ard i u m an d c o ro n ary v essel s h av e b een rem ov e d T h e at r i a
l p i . ar e in
a di s t en d e d o r di a s t o l i c c o n d i t i o n .

6 Ex t er n a l b u n d l e s se en o n t h e p o s t e r i o r su rf a ce o f t h e hu m an h e ar t . T he
vi sc e r al p i
e r c ar di u m a n d c o ro n a r yv ess el s ha v b e ee n r e m o vd
e . T he i
a t r a ar e in
a di s t en d d
e or di t l i
as o c c on diti on .
 PLATE 4

X L A N A I N F I G UR
E P T O OF ES

7 I n t ern a l b d l f t h i gh t t i m f t h h m
un es o e r a r u o e u an h e ar t , p i
os t er o r vi ew .

T he v en a c a v hv b
ae a p d t h gh t h i p t i
e ee n o en e rou e r o s er o r w al l s . T he i nf i
er or

v en a c a v a is s p d w id l y p
r ea Th
e d
o di m h b
en . e en o c ar u as e en r e mo vd e .

8 I n t e rn al b u n d l es of l ef t i
at r u m o f t h e h u m an h e ar t , p i Vi
o st e r o r ew . T he
p ost er o r i w a ll of t he i
at r u m h as b een c u t an d W id el y o p en e d . T he en d o c ar di u m
h as b e e n r em o vd
e .
MU S C U LAT U R E P LATE 4

W P AP E /

IV G .
R es u me n por a u tor Harvey Er n est J orda n
el , ,

Un iversidad de V irgi nia .

N u evos est u dios sobre la m éd ula osea roj a .

I . Experi m e n tos II Citolog i a c o n especial m en c i on de los

. .
,

datos qu e i n dica n la existe n cia de u n a actividad i n tracel u lar

h em o c i t og én i c a e n las cél ulas giga n tes y la si g n i fi cac i on de
l as llamadas fi g u r a s m i t ot i c a s en estas cél u las .

L a m ed ula osea roj a del co n ej o co n tie n e tres variedades pri n

c i p al e s de cél ulas gigan tes : m eg acari o c i t o s formas m on onu cl ea ,

das derivadas a co n sec u e n cia del crecimie n to excesivo del hemo

blasto ; p oli m o rfo c ari o ci t o s for m as c o n n ucleo polimorfo o lob u ,

lado derivadas del m eg ac ar i o c i t o por di v i si é n directa i n completa

del n ucleo ; y p oli c ari o c i t os formas m ul t i nu cl ed as derivadas de ,

formas c o n n ucei lob ulado a co n sec u e n cia de la d i v i si é n completa

de dicho n ucleo qu e da l u gar a la f o rm aci é n de n ucl eo s d e me n or
,

tamano V ariedades cel ulares comparables a las me n cio n adas

.

existe n tambié n en el saco Viteli n o del em b ri on de cerdo Al gu .

n a s cél u las giga n tes de la variedad m u l t i n u c l ea d a ret i en en en -

peq ue no grado la capacidad del h em o b l ast o a n cestral origi n ario

para tra n sformarse en eri t r o c i t os i n tracel u lares L as células g i .

ga n tes de la médu la n ormal y saco Viteli n o carece n de fu n ci on

fa g o c i t i c a L os g ranul o c i t o s p ol i m o rfo nu cl ea do s d e tipo eo si n é
.
_

fi l o especial q u e aparece n c o n mayor o me n or ab u n da n cia de n tro

del protoplasma n o debe n co n siderarse como u n a di fer en ci aci on
i n tracel u lar o u n a i n gest i o n f ag o c i t i c a si n o qu e debe n i n t erpr e ,

tarse como i n vasio n es pasivas sec u n dar ias debidas al caracter

relativame n te me n os resiste n te del citoplasma de la cél ula giga n te ,

c u an do se compara c o n el del p aren qu i m a ge n eral de la méd ula ,

co n tra la presi é n ej ercida por u m foco de l eu c o ci t o s em V i as de ac

tiva p rolifera ci é n o como i n vasio n es activas por parte de fago
,

c i t o s polimorfos de u n citoplasma en V i as de desi n t eg rac i é n

,

Estas cél u las giga n tes n o se divide n por mitosis L os grupos de .

p a rt i c u l as c ro m at i c a s qu e se e n c u e n tra n alg un as veces sim u la n do

placas de cro m osomas en h u sos mu l t ipl o res so n simpleme n te
agregacio n es cas u ales de cromati n a c arac t eri st i c a s de procesos ,

dege n erativos L as agregacio n es n o v a n ac o mp a fi an d as de fi bras

.

a r c o p l asm i c as Estas fi g u ra s so n simpleme n te sim u lacros de

.

p rocesos m i t ot i c o s q u e represe n ta n estados fi n a l e s de la d esi n t e

g rac i on de las cél ulas giga n tes .

T ra n s l a t i on by J os éF N o m d ez
C C ll g
.

o r n ell U m v erst t y Me di c a l o e e, N Y
2 88 H . E . JOR D AN

work w as u n der w ay desig n ed to test the hypothesis that the

gian t cells of n ormal red bo n e marrow represen t p ot en t i al m ul
- -

tiple h em ob last s like those of the yolk sac by a st udy of the -

,

marrow an d the splee n d u rin g the rege n erative stages followin g

experime n tal hemorrhage This work has n ow progressed to a .

poin t where a report of res ults seems warran ted The resu lts do .

n o t disprove the esse n tial acc u racy of this hypothesis b u t n either ,

do they give to it any support The sectio n s of this p o st hem or .

rh ag i c marrow were s u bseq u e n tly employed together with the ,

co ntrol material for a st u dy of the si g n i fic an ce of the alleged

,

mitotic fi g u res i n the gian t cells -

.

DES C RI F TI O N
Two rabbits were u sed i n this st u dy They were deprived of .

an amo un t of blood drawn from the femoral vein equ al to 2

, ,

per cen t of their weight Five days later they were killed an d .
1
,

specim en s of spleen an d of marrow from the femu r were fix ed

severally i n a mixt u re of 1 0 parts of formalin to 1 00 parts of a
sat urated n ormal salt sol utio n of corrosive su blimate an d i n
-

H elly s fl u id The stain ed sectio n s of marrow showed exten sive

’
.

hemopoietic activity b ut the gian t cells wer e n ot appreciably

,
-

differe n t from n or more n umero u s than those of n ormal c on

, ,

trols the marrow from which w as similarly fix ed an d stain ed

,
.

The spleen also showed n o i n crease i n the nu mber n or any

chan ge i n the character of the very few relatively small gian t
cells .

M ore favorable resu lts were to be expected from the marrow

of the guin ea pig for here larger nu mbers of the gian t cells are
-
,
-

of the multinu cleated type It was assumed that these more .

exten sively lob ulated and mu ltinu cleated gian t cells were at -

least on e step n earer the stage where un der appropriate hemo

ge n ic stim uli i ntracell ular erythrocyt es co u ld differe n tiate an d ,

that in co n sequ e n ce the relatively i nt en sifi ed posthemorrhagic

hemopoiesis wo uld i n volve these pote n tial m ultiple hem ob l ast s .

1
I n t h e se e xp i m t I
er i v d v l b l id f
en s r ec e e a ua e a ro m P f ro . J . H . N ff
e and Dr .

D C
. . Smi th ,
o f th U iv i ty f V i gi i H p it l
e n ers o r n a os a s t af f .
 GIANT C ELLS -
OF B ON E —
MARROW 28 9

Bu t the gian t cells of the red marrow of the fem u r of a g u in ea

-

pig of 400 grams weight from which 1 0 cc of blood had bee n

’
,
.

taken directly from the heart were after fi ve days of rege n era ,

tive activity on the part of t hi s marrow n ot essen tially d ifferen t

‘

from those of n ormal marrow .

I then resolved to t ry the effect of a twice repeated bleedin g —

.

From a g uin ea pig weighin g 42 0 grams 1 0 cc of blood was

-
, .

taken directly from the heart on D ecember 23 rd O h J anu ary .

8 t h an other 1 0 cc was removed Before killin g this an imal

. .

fou r days later 1 2 cc of blood w as removed i n order to free the

.
,

marrow as far as possible from circ ulatin g blood Hemopoietic .

activity w as in ten se i n the femu r b ut the gian t cells agai n gave ,

-

n o evide n ce of active participatio n i n the eryt h ro cyt o g eni c

process .

It wa s then decided to experimen t with pigeo n s The special .

poin t of valu e of this material seemed to b e the absen ce of

gian t cells i n the red marrow It seemed reason able to suppose
-
.

that possibly the in creased posthemorrhagic hemogen ic deman ds

might stimu late the poten tial gian t cell progen itors to develop -

in to genu in e megakaryocyt es with eryt hro cyt o g eni c capacity , .

Two pigeo n s of 300 grams weight were aspirated ; from on e 5 cc ’

.

of blood w as removed from the other 8 cc The fi rst w as killed,

.

fo u r an d on e half days later the secon d fiv e days later The

—
,
.

marrow from these femu rs showed i n ten se hemopoietic activity ,

b u t n o gian t cells were discern ible

-
.

From the resu lts of these ex perim en ts I draw the co n cl u sion

that suble thal experimen tal hemorrhage is n ot an adequ ate c on
ditio n to st i mu late in tracellular eryt h rocyt og en esi s i n giant
cells of the red bon e marrow of the g u in ea pig an d the rabbit
- —
,

n or to in d u ce gian t cell formatio n i n marrow n ormally lacki n g

-

su ch elemen ts as i n the pigeon App aren tly the inten se hemo

,
"

poietic activity obtain in g n ormally i n t he yolk sac is n ot the -

sp eci fi c factor which i n d u ces the slight eryt h ro cyt o g en i c activity

of the gian t cells of this hemopoietic organ The fact that a

-
.

similar activity on the part of these cells occ u rs i n certain patho

logic marrows for example typhoid m arrow s u ggests that the
, ,

causative factor m ay be the presen ce of a toxic agen t followi n g

r m: A M ERIC A N J QU R N A L o r ANA O M Y V OL T , . 27, NO . 3
2 90 H . E . JOR D AN

regressive an d disi n tegrative chan ges in the atrophyin g yolk sac —

an d the morbid marrow respect ively The experimen tal evi

,
.

den ce s u ggests that the i n tracell ular eryt h ro cyt og en i c activity

of the hemoge n ic gian t cells is of n egligible physiologic si gn i fi
‘ ’

can ce .

The vario us microscopic preparation s of the red bo n e marrow

of the rabbit an d the gu in ea pig o ffered a favorable material -

for the st u dy of the e ni gmatic an d disp u ted si g ni fi c an ce of the

so called
-
mitotic fi g ure s vario u sly atypical and apparen tly
,

m ultipolar of the gian t cells an d of the data su ggestin g an

,
—
,

in tracellular eryt h rocyt og en i c f un ctio n of these cells It will .

co n d u ce to a si m p lifi c at i on of the an alysis of the complete prob

lem to co n sider the latter matter fi rst .

II C YT O L O G I C
.

I N T RA C ELLUL AR ER YTH R O CY T O GEN ES I S

R evi ew o f li tera ture

This phase of the s ubj ect i n volves the qu estio n of the origi n
of the m ultiple n u clei of the p olyk aryo cyt es an d the si g nifi can c e
of the occ urren c e of gran ul ocytes an d erythrocytes i n the gian t
cell cytoplasm A rn old ( 1 2 ) describes a mode of nu clear
.
,

divisio n i n the rabbit s marrow which he terms i n direct frag

’ ‘

men tatio n This he claims is followed by a divisio n of the

.
’

cytoplasm which prod u ces separate mo n o n u cleated and poly

,

m o rph onu c l eat ed le u cocytes A ccordin g to A r n old therefore

.
, ,

these cells divide by a method essen tially amitotic an in ter ,

,

m ediate phase of which process is represe n ted by a m ul t i nu

cleated giant cell Heide n hai n ( 8 ) recogn izes amitotic n u clear
-
.

mul tiplication b ut regards m ul tipolar mitosis un accompani ed by

, ,

cytoplasmic divisio n as the more importan t an d the prepo n der

,

an t mode of n u clear proliferatio n H e i n terprets the prese n ce .

of granulocytes as the res u lt of an i nvasio n n ot phag ocyt osis , .

P ugliese ( 1 5) in terprets the prese n ce of erythrocytes an d leu co

cytes i n the gian t cell cytoplasm as the expressio n of an endog e
-

n o u s h em o c yt o g e n i c capacity F o a ( 7 ) also describes an in tra

.
2 92 H . E . JOR D AN

e n doge n o u s blood cells illu strated i n D enys fi g ures however

-
’
, ,

o n ly two or three have the nu clear characteristics of an erythro

cyte The practically excl u sive type of en doge n o u s cell is the
.

p o l ym o rp h o n u c l eat ed gra n u lar le u cocyte pres u mably the special ,

gran u locyte with fin e eosin ophilic gran u les These cells fill the .

gian t cell cytoplasm mean while compressin g co n sumin g an d

-
, , ,

eve n t u ally destroyi n g the gian t cell n u cleu s un til the origi n al -
,

cell becomes a mere cyst fill ed with the e n doge n o u s le u cocytes .

E ve n t u ally the cyst is said to r u pt u re and to free the n ew formed -

eosi n ophilic granu locytes In a sectio n of on e s u ch cyst eighty

.

cells c an be co un ted ; the complete cyst may well have i n cl u ded

several hun dred le u cocytes D e nys co n cl u des that this series of
.

eve n ts si g n ifi es a n ormal hemogen ic fun ctio n on the part of these

gian t cells in volvi n g a genui n e amitotic divisio n Heiden hain
-
,
.

wo u ld in terpret these phen omen a as the resu lt of the i n vasio n

of the gian t cell cytoplasm by phagocytic leu cocytes ; D ickso n as
-

sign ifyi n g phagocytosis of polymorphs on the part of the gian t

cells D en ys co n clu sio n will be fu rther disc u ssed below
.
’
.

Descri p ti on

This chapter will o n ly i n cl u de a descriptio n of those cells i n

my preparatio n s which relate to the process which D e nys d e
scribes as divisio n by ste n ose a process i nterpreted as leadin g
‘
,
’

to the formatio n of i n tracellu lar leu cocytes In fi g ure 1 is ill u s .

‘

t rat ed a cell which represe n ts the begin n i n g of the gian t cell -

series It is esse n tially an e n larg ed hemoblast It traces back

. .

to the typical hemoblast of hemopoietic tiss u es ; an d forward by ,

small gradation s to a cell like that ill u strated i n figure 2 The

,
.

latter cell is m u ch larger an d co n tai n s a re n iform or more deeply

cresce n tic frequ en tly i nten sely stain in g nu cleu s These two cells
,
.

are ge nu in e megakaryocytes The n u cleu s of cell fi g ure 2 is

.

simply a m odifi cat i on of the spheroidal n u cle u s of cell fig ure 1 .

It seems desirable to emphasize the poi n t that these cells were

st u died i n serial sectio n s It is obvio u s that s imilar appearan ces
.

wo uld resu lt from possible sectio n s thro u gh very di fferen t an d

m u ch larger nu clei as for example sectio n s of cells fig ures 3
, , ,
 GIANT C E LLS
—
OF B ON E MARROW
-
293

an d 4 . these two cells (fig s 1 an d 2 ) and many others have

But .

app ro x irn at el y as simple a nu cle us as represe n ted i n these ill u s

t rat i on s The cell of fi gure 2 can be traced thro u gh small grada

.

tio n s to one l i ke that of fig ure 3 a p olym orph ok aryo cyt e The, .

n u cle u s has become more complex an d lob ulated A c en tral .

cytoplasmi c area ( e n doplasm of Heiden hai n ) co n tai n s a c on

‘ ’

S p i c u o u s g ro u p of ce n trosomes There is however n o i n dica

.
, ,

tio n of archoplasmi c fib ers .

By co n strictio n s at the levels between adj acen t lob ules the ,

n u cl eu s of t h e p ol ym orp h ok ary o cyt e be com es divid ed i n to smal l er

n u cl ei prod u ci n g th u s a polykaryocyt e (fi g
,
This cell also .

co ntain s a con spic u o us pluri corp uscul ar cen trosome i n the en do

plasm Appearan ces of discrete nu clei lik e that of fi gure 4
.
’
, ,

could co n ceivably res ult from tan gen tial sectio n s throu gh poly
morph nu clei lik e that of fi gure 3 b ut i n this partic ul ar c as e ,

a g ain and i n many others it coul d be proved by study of serial

, ,

sectio n s that t h e gian t c el l act u all y co n tai n s i n c ertai n in st ance s

-

several or many discret e nu clei .

In fi gure 9 is illu strated a g iant cel l i n which thre e n u cl ear -

b u ds have separated from the main polymorphou s nu clear mass .

Tw o of the n u clear b uds ( a an d e) hav e t he feat u r es of nu cl ei of

t h e marrow erythroblasts ; an d abo u t n u clear mass 0 a lay er of
cytoplasm has separated from the gian t cell ex oplasm thu s -
‘
,
’

givin g origin to an eryt hroblast which n ow appears to li e in a

vac u ole Figur e 8 illu strates i n simpl est co n ditio n the amitoti c
.

mode of intracell ul ar hemocyto g en esis described by D e nys as ,

above o u tli n ed T hi s cell co n tain s a polymorphou s n u cleu s with

.

scattered c en trosomes i n the endoplasm At t he right periph

‘
.
’

ery lies a p olym orph onu cl eat ed eosin ophi lic granu locyte In my .

preparation s any of the cells illu strated i n fig ures 3 4 7 8 9 , , , , ,

1 0 1 1 1 6 and 2 0 may co n tai n on e or several eosi n ophilic gran

, , ,

u l ocyt es Bu t o n ly mu lti nu cleated cells lik e those of fi g ures 4

.

an d 9 co n tai n erythroblasts .
2 94 H . E . JOR D AN

Di scussi on

The above sketched ge n etic series of eve n ts for these giant

-

cells agrees with that previo u sly o utli n ed for these same cells
both i n the yolk sac of the pig embryo ( 1 0 ) an d i n the red bo n e
—

marrow of the rabbit This developme n tal history is i n

s ubstan tial agreeme n t also with that give n by D e nys for the rab
bit s marrow and with Heide n hai n s statemen t that the gian t
’
,
’

cells of the rabbit s bo n e marrow arise from leu cocytes by i n ter

’
-
‘

c u rren t mitoses which n ever lead to division b ut res u lt in

’
,

nu clear ref u sio n s followi n g the mother star fi g u r e

‘ ’

I have show n i n a previo u s paper that i n the yolk sac of the —

1 0 mm pig embryo certain h em ob l ast s e n large a n d become bi

-
.

p ol ym orph o or m u ltinu cleated In the bi n u cleated co n

ditio n which is comparable to a polykaryocyte these smaller
, ,

gian t cells exte n sively differe n tiate e n doge n o u s erythrocytes

-
,

freq u en tly on e occasio n ally two These eryt hrocytes subse

,
.

qu en tly break thro u gh the peripheral shell of the origin al gian t

cell cytoplasm O ccasio n ally also i n the q u adri nu cl eat ed co n di
.

tio n e n dogen o u s erythrocyte formatio n occ u rs There is some -

,
.

slight evide n ce that qu adri nu cl eat ed (the u su al mu lti nu cleat ed

con ditio n i n the yolk sac ) gian t cells occasio n ally divide i nto
- -

b i or mo n o n u cleated forms before erythrocyte differe n t i atio n

occ u rs These giant cells of the yolk sac are multiple hem ob l ast s
.
- -

or small blood islan ds ( figs 1 7 9 1 0 1 1 2 4 2 5 2 6 an d 3 5 of

-
.
, , , , , , , ,

my article o n the yolk sac of the pig embryo ( 1 0) an d fi g s 1 to

-

2 4 of my article on gian t cells A similar occasio n al en

—

d o g en ou s origi n of erythrocytes was described also for pathologic

( typhoid ) human bo n e marrow -
Figure 9 above described , ,

also admits of n o other obvio u s explan atio n than that of endog

en o u s eryt h ro cyt og en esi s Si n ce these gian t cells are e n larged
.
-

an d m o d i fi ed h em o b l ast s it seems plau sible that they sho u ld

,

mai n tai n to some degree their origin al capacity for erythrocyte

differen tiatio n un der certain co n ditio n s However this phen om .
,

e n o n of in tracell ular eryt hro cyt ogen esi s i n gian t cells occ u rs -

i n so slight degree that it does n ot seem reaso n able to ascribe

to it any essen tial or eve n importan t r61e It seems more n early .
2 96 H . E . JOR D AN

However whe n we co n sider the gra n ulocyte i n cl u sio n s the

, ,

case seems q u ite di ffere n t In the fi rst place su ch cells do n ot

.
,

occ u r i n the yolk sac gian t cells Here the cells are qu ite widely
- -
.

scattered i n the blood S paces and n o t closely crowded as i n the

,

bo n e marrow F urt herm ore these gra nu locytes m ay occ u r i n

—
.
,

practically any type of older gian t cell i n red bo n e marrow b ut - -

,

most ab un dan tly i n those cells whose n u clear con ditio n i n dicates
dege n erative chan ges i n these cells ; wit n ess the fi gures of D e n ys .

If the prese n ce of the eosin ophilic polymorphs si g n ifi ed gian t

cell phagocytic activity then o n e wo uld expect all types of mar
,

row cells to be in gested i n approximately eq u al n u mbers But .

the polym orphs with fi n e eosin ophilic gran ules very greatly pre
po n derate Whe n on e con siders the fact that the polymorphs
.

of the rabbit with fi n e eosi n ophilic gran ules ( special gran ulo
cytes ) are the represe n tatives of the n e utrophilic poly morphs of
most mam mals an d that the latter are the predomi n a n t phago
cytes Heiden hai n s s u ggestio n that these gran u locytes have i n
,
’

vaded the gian t cell appears very plau sible This co n cl usio n is
-
.

i n agreeme n t with the fact of their greater ab u n dan ce i n o b

v i o usly dege n erati n g an d disi n tegrati n g cells .

The observatio n s of D e n ys as illu strated i n his fi g ures 1 6 to 2 3

are absolu tely un iqu e N o o n e else as far as I have been able to
.
,

learn has described anythi n g closely comparable i n degree with

,

c o n ditio n s he describes for his sectio n s of rabbit s red bo n e mar

’
-

row H e does say that great variatio n s occ u r i n regard to these

.

co n ditio n s i n di ffere n t i n divid u als b ut he does n ot specify any ,

pec uliar circ u m stan ce abo u t the i n divid u al from which this
specime n of m arrow was derived In my st u dy of preparatio n s .

of at least a score of differe n t i ndivid u als of rabbit and g uin ea

pig I have n ever see n an ythi n g like the co n ditio n ill u strated by
D e n ys i n for example his fi g ures 2 2 an d 2 3
, ,
D ickso n wo uld .

explai n these co n ditio n s as phagocytosis ; Heide nhain as the re

s ult of i n vasio n However correctly i n terpreted the con ditio n
.
,

of this degree m ust be extremely rare Besides the arg ume nts .

above give n agai n st an i nterpretatio n i n terms of phagocytosis

o n the part of the gia n t cells may be me n tio n ed this additio n al
-
,

an d appare n tly co n cl u sive co n trave n i n g fact as applied to the

 GIANT C ELLS -
OF B ON E MARROW
-
297

ill ustratio n s of D e n ys : It is in co n ceivable an d qu ite witho u t ,

parallel how a cell co uld co n tin u e to fun ctio n ph ag o cyt i cally

,

an d co n ti nu e to load itself with i n gested cells to the poi n t of

r upt u re lo n g after its nu cleu s had practically disappeared throu gh

,

press u re exerted by the extran eou s i n gested cells O n the o ther .

han d D e n ys evide n ce is far from co n cl u sive that the co nta i n ed

,
’

le u cocytes are act u ally differe ntiatio n prod u cts of the nu cleated
areas of the polykaryoc yt e cytoplasm The cr u cial li n k i n the .

evide n ce for s u ch co n clu sio n is lacki n g : he c an give n o satis

factory tran sitio n stage betwee n the p olym orph ok aryocyt e and
the gian t cell cysts We desire a cell showin g at least a score
-
‘
.
’

of isolated n u clei as a tra n sitio n to the cysts co n tain in g at least

a hun dred le u cocyt es i n terpreted as derived from n u clear b u ds
,
.

This tran sitio n stage is lackin g i n D e nys series of fi gures A t ’

.

most he can o n ly show a cell with two isolated b u ds Moreover .

,

the in tracell ular leu cocyt es themselves appear at approximately

d efi ni t i v e stages of diff ere n tiatio n n ot at widely differe n t stages
,

of diff ere n tiatio n as wo uld be expected if they act u ally developed

i ntracell u larly .

Whe n on e co n siders D e n ys fi gures 1 7 to 2 3 o n e gets the i m

’
,

pressio n of a mass in vasio n of gran u lar polymorphs in to these

gian t cells The n u cle u s is p u shed to on e side ; it is greatly com
-
.

pressed and fi n ally broke n up an d destroyed Th e s ug gestio n .

presen ts itself very persisten tly that these mass in cl u sion s of

p yo l m o r p h o n u c l e at e d gra nu locytes sig n ify an 1 n v a s1 o n of these
cells du e to the close j uxtapositio n of areas of in ten sely prolifer
,

atin g gran u loc yt es an d less resistan t areas of closely packed ad

j ace n t gian t cells There remai n s n o acceptab le evide n ce that
-
.

the hemoge n ic gian t cells of red bo n e marrow of the rabbit pos

- —

sess any i ntracell ular l eu c o cyt op oi et i c fu n ctio n There is how .

,

ever a residu u m of rigidly tested evide n ce which s u ggests a

,

slight eryth ro cyt op oi et i c capacity on the part of these cells ,

both i n the yolk sac ( 1 0 ) an d i n the red bo n e marrow

- -
29 8 H . E . JOR D AN
THE SI G NIFI C AN C E O F TH E SO -C A LL E D M I T O TI C FI G URES
R evi ew o f li ter a ture
U n der this head also the i n vestigatio n of D e n ys is i n this c on
n e c t i o n by far the most importa n t D e n ys (4 ) alo n e gives a .

detailed acco unt with adequ ate ill u stratio n s of what he believed
, ,

to be mitotic divisio n of the hemoge n ic gian t cells of red bo n e -

marrow A rn old s ( 1 2 ) acco un t obvio u sly relates to degen er

.
’
,

ative chan ges as already su ggested by D ickso n D ickso n ( 5)

,
.

desires his ow n acco un t of mitosis i n these cells to be accepte d

o nly as a prelimin ary sketch B un tin g ( 3 ) merely refers to the
‘
.
’

mitotic divisio n of these cells withou t givi n g any detailed de ,

scriptio n or illu stratio n s Heidenhain gives u s o n ly meager de

.

tails of what he regards as m u ltipolar ( mu ltiple mitoses of these ‘

gia nt cells (pp 62 2 62 5 . The stateme n t by V an Bamb ek e

—

a n d V an der Stricht ( 1 6) that the lob u lated nu cle u s of the mega

karyocyte is formed by f u sio n of a large n umber of small dau gh

ter n u clei d uri n g the telophase of man y repeated m u ltipolar
-

mitoses is hardly more than a s u ggestio n N evertheless these

,
.
,

less complete acco un ts of an alleged mitotic process will be b ri efly

co n sidered after a detailed a n alysis of D e n ys evide n ce for mitosis ’
.

Mitosis is said by D e nys to be i n itiated by the rearran geme n t

of the chromati n i n to a de n sely wo u n d thread like the close spi ,

reme of bi n ary mitotic divisio n (fig s 2 3 24 1 8 an d S u b se .

, , ,

q u e n tly this spireme segme n ts in to short rods ( chromosomes ?)

which arran ge themselves first i nto a spherical mass (fi g .

a n d later i n to what is described as a basket str u ct ure with polyg

‘

o n al meshes Figu re 1 5 ill u strates well the several varieties of

.
’

this latter stage as fi g ured by D e nys This stage is said to rep .

rese n t that of the equ atorial plate of bin ary mitosis j u st preced ,

i n g the lo n git u di n al divisio n of the chromosomes D e n ys de .

scribes the shape of the alleged chromosomes at this stage as

havi n g the form of a U with the ben d of the U directed toward
the cen ter of the nu cle u s Prese n tly they are believed to s u ffer
.

a lo n gitu di n al splitti n g after which the dau ghter gro ups of

,

chromosomes separate and take pos itio n s as the polar gro ups of
m u ltipolar mitoses ( fig These gro ups are the n said to re
.
3 00 H . E . JOR D AN

are give n regardi n g the sp ecifi c r 6le of the ce ntrosomes i n this

process n or does he give any fig ures showi n g astral rays or
,

spi n dle fib ers accompan yi n g the m ultiple mitoses ‘

.
’

B unti n g describes pec u liar complex mitotic fi g ures i n

'

‘ ’

‘
dividi n g m eg al okaryocyt es of the rege n erati n g red bo n e mar ’
—

row of the rabbit after experime ntal i n toxicatio n with sapo n in

, ,

rici n or t urpe n tin e H e fi nds n o evide n ce however of a divi

,
.
, ,

sio n either of the chromatin masses or of the giant cell cyt oplasm —
.

N o me n tio n is made of either ce n trosomes or S pi n dle fi b er ; it

may therefore be i n ferred that s u ch were n ot see n i n co nn ectio n
with these mitoses H e s uggests that these appare n tly i n com
‘
.
’

p l et e an d m u ltipolar mitoses lead merely to an i n crease i n

, ,

the complexity of the polymorpho u s n u cle u s an idea very like —

that expressed by Van Bam b ek e an d Van der Stricht .

The seriatio n described by D ickso n seems o n its face qu it e

improbable The relatively small size of his cell at prophase
.
‘
,
’

fi g ure 2 ( fi g an d of the cells at a n aphase (fi g s 1 4 an d 1 5)

‘ ’
'
. .

is n oteworthy D ickso n also makes n o me n tio n of ce n trosomes

.

or S pi n dle fib ers i n these ass umed mitotic phases It sho uld be .

stated also that two of the fi gures i n this series ( fig s 1 0 an d 1 7 ) .

occ ur o n ly very i n frequ e n tly i n my material D ickso n s fi g ure .

’

correspo n din g to my fig ure 1 0 co n tai n s a spireme of co n sider ‘ ’

ably lesser girth D ickso n te n tatively describes mitosis i n

.

these gian t cells as follows : -

The t e i n thi s c om p li c ate d divi si on see m s t o be an agg reg a

fi rst s ag
ti on of the b asket like nu cle us ( fig o f the g i an t cell alre ady d e
-
.
—

s c ribe d i n t o a m ore c o m p ac t so li d m ass su ch a s i s sh own i n the C ell 1

, , ,

of the fi gure ( fig This process of c on den sati on m ay c on ti nu e

.

u ntil the wh o le nu cle us be c om e s m u ch re du c e d i n t ot al b ulk an d d u r ,

i n g whi ch p r o cess the st ai n i n g re a c ti on of the chr om ati n bec om es m ore

i ntense (Note It i s possible the st ag e fig u ed i n C ell 2 ( fig 1 7 )
. .
—
r .

m ay be a s om ewh at l ater on e i n the pro c e ss an d m ay re pre se nt the ,

ph ase whi ch f oll ows d ivi si on an d re arr an g e m e n t o f the chr omo some s ) .

The n u cle u s n ow pr oc ee ds t o arr an g e it self i n t o o n e l on g c onti nu o us , ,

c o n v ol ute d thre ad like str uctu re Whi ch g radu ally be come s thinner
,
-

a n d m o re c o n d e n se d an d whi ch sh ows the chr o m ati n be co m i n g ag g re

,

g at e d i nt o l ittle r o un d e d m asse s d o tte d at re g u l ar i n terv al s al o n g i t s

4
The f
r e e r e n c es t o fi g u r es , gi v en in b ra c k et s, in th i qu
s i
o t at o n a r e in ser t e d by
m e, an d d i gn
es at e i
fi g u r es l l u s t ra t i g thi
n s ar t c l ei .
 GIANT C ELLS -
or B ON E -
MARROW 30 1

e ntire length the red nucle ol ar st aini ng havi ng n ow di sappeare d

,

( spirem e or p ro ph ase) (fig Thi s thread o r spire m e when the

.
,

c o n de n sati on of the chrom ati n 1 s c o m plete breaks up (m et a

ph ase) i n t o sh ort seg m ent s re sem bl i n g chr om osom e s whi ch
be com e rearrang e d into a den se r oun de d g l obe i n the c enter of the
c ell ( an aph ase) ( fi g . after whi ch it i s pr ob able th at the re su lti n g
b asket nu cleus i s rebuilt u p on a m ore c omplex pl an by a proce ss ,

resembli ng that by whi ch t he orig i n al un derwent the transform ati on

in to chrom osom e s i e by st age s sim il ar t o th ose j u st de scribe d b ut
,
. .
, ,

o c cu rri n g i n the rev erse or der Thi s i ntric ate proc ess of
.

nu c le ar r ea ran gemen t r ather th an di vi si on is n ot ac com p anie d o r f ol

r

l owed by corre sp on din g divi si on of the cell b ody or tel oph ase the pro -
,

t o pl asm si m ply i n cre asi n g i n b u lk as the n u cleu s be c om es l ar g er an d

m ore com pli c ated .

Descr i p ti on

The earlier steps ( fi g s 2 3 and 4) i n the developme n t of t h e

, ,

several varieties of the gian t cells from the e n larged hemoblast

-

”
(megakaryoc yt e fi g 1 ) have already bee n co n sidered above
,
. .

P olyk aryo cyt es arise from p ol ym o rp h ok aryoc yt es thro u gh the

divisio n of the lob u lated nu cleu s of the latter Certain of the .

isolated n u clei of the polykaryocyt e may fun ctio n as ce n ters of

i n tracell ular diff ere n tiatio n of erythrocytes (fi g Poly .

karyocyt es moreover s uffer dege n erative chan ges characterized

, ,

by the aggregatio n of the chromati n of the separate nu clei i n to

peripheral masses sim u lati n g equ atorial plates of chromosomes
(fi g. The origin ally distin ct gro u ps s ubsequ e n tly become
mi n gled a process see n at the begi nn in g at the left of fi gu re 5
, ,

an d fi n all y become scattered thro u gho u t the cell ( fi g The .

l atter may occasio n ally show an arran geme n t of the chromati n

segmen ts ( chromosomes ?) lik e that of fi g ure 1 5 simulati n g a ,

mu ltipolar mi totic fi g ure lik e that of certai n n eoplastic cells ;

b u t n either ce n trosomes n o r sp i n dle fi bers are here discern ible .

The chromatic rods of fi g ure 6 su bsequ en tly become pale an d

granul ar an d appear to dissolve within the disin tegratin g cyto
plasm .

The gian t cell with basket n u cle u s (p olym o rph ok aryo cyt e)
-

also passes throu gh early stages of dege n eratio n by gradu al

steps as ill u strated i n fi g ures 7 8 9 an d 1 1 The nu cleu s i n t h e
, , ,
.

last of these stages becomes compacted i nto a spheroidal mas s

30 2 H . E . JOR D AN

w ith very irreg ular an d chromatic co n to u r The chromati n .

has collected i nto spheroidal a n d bacillary droplets of greatly

varyi n g size S ubsequ e ntly the nu clear wall disappears an d
.

th e chromatic material collects i nto a complicated meshwork

simulati n g somewhat the loose and segme n ti n g spiremes of
normal mitosis ( fi g . The latter stage passes i nto one char
ac t eri z ed by partially isolated masses of chromati n sim u latin g

th e segme n ted spireme which may pass before dissol utio n

,

thro ugh a stag e characterized by the aggregatio n of still fi ner

chromatic gra nu les i nto gro u ps showi n g fo u r or more co n de n sed
li n ear areas thu s givi n g the impressio n of a mu ltipolar mi totic
,

fi g ure b ut witho u t i n dicatio n of spi n dle fib ers

,
Cells like that.

of fi g u re 1 5 disappear thro u gh sol utio n of the ch rom at i c b o di es

'

W ithi n the disi ntegrati n g cytoplasm .

The gian t c ell s m ay disin tegrate by still an other dege n erative

-
.

ro u te R epeated amitotic n u clear divisio n of a polykaryocyte

.

may lead thro u gh stages represe nted by fi gures 4 1 6 an d 1 7 , , .

The relatively small n u clei of cells like that of fi g ure 1 7 may

the n s uffer a peripheral co n de n satio n of their chromati n fol ,

lowed by f usio n (left of fi g which steps lead to a co n dition

.

like that show n i n fi g u re 1 9 ; after this stage the n ow n aked

nu cle u s loses its stai n i n g capacity a n d fragme n ts Fin ally the .
,

polymorpho u s n u cle u s of certai n gian t cells may begi n to stai n

-

very deeply givi n g to the whole a homoge n eo u s appearan ce

,

(fi g. S u ch a cell has e n tered u po n a stage of active cyto

plasmic disi n tegratio n The latter progresses forcin g a c o n
.
,

de n satio n u po n the nu cle u s (fi g The nu cleu s persists

.

for a while lo n ger as a compact n aked body (fi g b u t eve n .

t u ally disappears by fra g me n tatio n Fig u res 23 an d 2 4 ill us

.

trate two additio n al variatio n s of nu clear chan ges d u ri n g dege n

c ratio n of the type of cell represe n ted i n fi g ure 1 8 .

D i scu ssi on

The foregoin g description shows that the several later stages

i n the alleged mitotic divisio n of the med u llary gian t cells rep re —

se nt i n fact the termi n al stages i n at least fo ur differe nt modes

304 H . E . JOR D AN

phase an d metaphase gro ups of chromosomes of m ultipolar

S pi n dles
. They have ass umed a correspo n de n ce with the typ
ical m u ltipolar mitoses of n eoplastic tiss u es But n o one with .
,

the exceptio n of D en ys has act u ally claimed a co n s u mmated

,

mitotic divisio n The other i nvestigators have o n ly ve n t u red

.

to i nterpret these mitotic simu lacra as a method by which the

p yo l m o rp h o n u c l eat ed n u cle u s of the gian t cell becomes still -

more complex N o o ne has claimed the prese n ce of spi n dles

.

i n co nn ectio n with these fi g u res N or has any o n e appare n tly .

see n the si g nifi can ce of the red u ced amo u n t of cytoplasm of

the cells co n tain i n g these alleged mitotic fig ures ; n or taken
acco un t of the obvio u sly dege n eratin g n aked nu clei i n their
seriatio n s of stages i n the life history of these cells The abse n ce
—
.

of spi n dles alo n e however marks these fi gures at o n ce as some

, ,

thi n g di ffere n t from the m ultipolar mitoses of n eoplasms an d of ,

morbid and experime n tal ly m odi fied tiss u es .

Si n ce Wright ( 1 8 ) p ublished his more complete paper with ,

co n vin ci n g colored illu stratio n s o n the origi n of b lood

platelets from the cytoplasm of the medul lary gian t cells we -

,

have bee n furn ished a n ew basis for the i nterpretatio n of these

vario u s nu clear m odi fi cat i on s Wright s co n clu sio n regardin g .
’

the ge n esis of blood platelets has n ow had confi rmat i on at the

-

hands of at least three s u bsequ e nt i n depe n de n t i n vestigators ;

B unti n g ( 3 ) D ow n ey (6) an d J ordan ( 1 1
, ,
Blood platelets ,
-

arise by a process of pse u dopod co n strictio n s an d by cyto ,

plasmic fragme n tatio n from hemoge n ic gian t cells The former

,
-
.

method of origi n is ge n erally restricted to the cells with t he

more vesic ular nu clei ( fi g s 1 to the latter method to those
.

cells havi n g deeper stai n i n g more complex an d less regular ,

nu clei ( fi g s 8 to 1 1 an d 2 0 to
. It w as show n i n previo u s
st u dies that blood platelets arise by a similar method from sim
—

il ar cells an d eve n from the pare n t h em o b last s i n the yolk sac

, ,
-

of the l O mm pi g embryo
-
. a n d that le u cocytes ge n erally ,

both lymphocytes an d gra nulocytes i n the red bo n e marrow of ,

-

the frog possess the capacity of co n strictin g off platelet like -

bodies The co n cl u sio n was there stated that platelet

formatio n is a b y prod u ct of the n ormal activity of le u cocytes
-

an d of the disi n tegratio n of dege n erati n g le u cocytes .

 GIANT C E L LS
-
OF B ON E -
MARROW 305

A n other fact m u st be take n i nto co n sideratio n i n this co n n ec

tio n n amely the divisio n of the ce n trosome i n to a p l uri co r
, ,

p u sc u l a r body a n d n ext the

,
scatteri n g a n d fi n all y the disappear
a n ce of these eleme n ts co n comita ntly with su ccessively later
steps i n the dege n eratio n process as j u dged by the co n ditio n of
the n u cle u s Whe n on e keeps i n mi n d the fact that the gian t
.

cell s u ffers a co n ti nu al di m i nu tio n of its cytop l asm thro u gh

the formatio n of blood platelets the i n variably smaller size
—
,

( see also D e n ys fi g u res) of the cells with the later n u clear

’

dege n erative chan ges sim ulati n g gro ups of chromosomes beco m es
i n telligible .

The q uestio n arises as to whether the n u cle u s u n dergoes

dege n erative cha n ges becau se the cytoplasm i s grad u ally bei n g
elimi n ated i n the formatio n of platelets or whether cytoplasm
is bei n g discarded an d platelet s i n cide n tally formed becau se t he
nu cle u s s u ffers dege n eratio n or wh ether both eve n ts are the
,

combi n ed e ffects of the sam e fun dam e n tal c au se The most .

plau sible i n terpretatio n i n V iew of all the facts i n cl u din g the

, ,

data previo u sly derived fro m a st u dy of gian t c ells of the yolk -

‘

sac an d of frog s m arrow is as follows : A s the res u lt of the

’
,

operatio n of some u n k n ow n fac t o r certai n h em o bl ast s e n large

_ ,

to form the simplest type of gian t cells This e n largeme n t dis —

.

t u rb s the opti m u m n u cleocytoplasmic relatio n In an atte m pt .

to recover this origi n al optimu m relatio n pse u dopods are pro ,

j e ct ed ; these co n strict an d break up i n to platelets an d the cyto

plasmic vol u me relative to the n u cle u s is th u s red u ced T he .

same en d may be served by the nu cle u s thro u gh i n crease of its ,

b ulk an d its s u rface area by e n largeme n t lob ulatio n an d fi n ally

, ,

direct divisio n The dist u rba n ce of optimu m n u tritive co n di

.

tio n s operati n g mea n while works an u n toward effect u po n the

ce n trosome which i n co n sequ e n ce fragme n ts i n to a p luri c orp u s
c u l a r ce n troso m e F ailu r e to cope with the factors e ffecti n g
.

i n terfere n ce with the optim um n u cleocytoplasmic relatio n

i n itiate gross dege n erative cha n ges expressed on the part of the
cytoplasm by m ass fragme n tatio n an d o n the part of the nu cle u s ,

( as i n fi g 4 ) by a n aggregatio n of the chrom ati n i n peripheral

.

drop lets (fi g S u bse q u e n tly the nu clear membran e di sap

.

TH E AM E R I C A J OU R N A
N L OF ANA O M Y V O L
T ,
27 , NO 3
 30 6 H . E . JOR D A N

pears the chromoso m e like masses beco m e m i n gled ( fi g

,
—
and .

u ltimately the e n tire cell disi n tegrates A ll the co n spic u o u s .

sig n s of dege n eratio n as they relate to nu cle u s ce n trosome

, , ,

an d cytoplasm are appare n tly the co mm o n e ffect of the sa m e

u n derlyi n g cau se of which platelet formatio n is a mere b y prod

,
—

u ct. T he n u clear alteratio n may be of vario u s sorts as ill u s ,

t ra t ed i n fi g u re 1 1 to 1 5 1 7 to 1 9 a n d 2 0 to 2 2 T hese dege n, ,
.

e ra t i o n phe n ome n a prod u ce appearan ces simu lati n g m u lti

polar S pi n dles , both i n the yolk sac (fi g 1 3 ) an d i n the marrow -
.

( figs . 1 2 1
,
4 a n d ,
A s the cytoplas m becomes red u ced i n
amo u n t the n u cle u s i n co n se q u e n ce becomes more an d more
compressed (fi g s 2 3 2 4 1 8 an d 1 9 a n d fi g s 2 0 to
.
,
a n d u lti
, , ,
.

mately o nly n aked compact n u clei remai n The latter ar e n ow

,
.

of s u ch red u ced size as to be n o lo n ger mechan ically barred from

the i n itial med u llary blood vessels an d i n co n sequ e n ce may hn d -
,

.
. their way i n to the peripheral blood stream an d become lodged
i n the capillaries of the l un g before fi n al dissol u tio n .

There is i n short n o adeq u ate evide n ce i n dicative of ge nu i n e

, ,

m u ltipolar mitoses i n these gian t cells ; the mitotic sim u lacra c an —

all be reaso n ably i n terpreted i n terms of n u clear disi n tegratio n .

Th u s is explai n ed o n e of the most p u zzlin g phe n ome n a relati n g

to these cells These cells wo u ld seem to be of the n at u re of
.

atypical cell gia n ts witho u t fun ctio n other tha n the on e of

‘
,
’

prod u ci n g blood platelets i n cide n tally to their disi n tegratio n

—
.

If a liberal i n terpretatio n of the n at u re of a chromosome as ,

simply a m ass of chromati n witho u t sp ec ifi c co n stit u tio n or

f un ctio n may seem to permit the cl assifi cat i o n of the chromatic
,

bodies formed by peripheral n u clear co n de n satio n s of the ch ro

mati n of the gian t cells i n the category of chromoso mes it
-
,

remai n s tr u e n evertheless that the mitotic arran geme n t of

, ,
‘ ’

these masses un associated with S pi n dles o n ly represe n ts a n

, ,

abortive atte m pt at divisio n .

The above i nterpretatio n of the si g nifi can ce of the alleged

mitotic fi g u res of the gia nt cells is f urther s upported by eve n ts -

i n the gia n t cells of the e n amel p ulp ( fi g s 2 5 to

-
The three .

cells here ill u strated are from the j aw of the new bor n cat -
.

They r eprese nt three s uccessive sta g es i n the disi nte grative proc
 H . E . JOR D AN

In fi g u re 2 7 we m eet with a still later stage i n this same dege n

e rat i v e process T he cytoplas m
. at certai n poi n ts o n the p e
r i p h ery appears to be disi n tegrati n g Mea nwhile the chromati n .

masses have become scattered thro u gho u t the cytoplasm N ear .

the ce n ter m ay be see n what appears to be the origi n of paired

chromatic rods by o u tflow of chro m ati n from a n uclear vesicle .

T hese chro m osome sim u lacra m ay by cha n ce become arra n ge d

—

i n certai n portio n s i n to c o n fi g u ra t i o n s s u ggesti n g m u ltipo l ar

S pi n dles of ca n cer cells In certai n portio n s they become more
.

closely massed i nto a fi ne meshed chromatic n etwork Certai n

—
.

of the chromatic ro d let s fragme n t an d certai n masses lose their ,

deep stai n i n g capacity Eve n t u ally all of this chromati n dis

-
.

solves withi n the disi ntegrati n g gian t cell cytoplasm The close -
.

correspo n de n ce betwee n the phagocytic sy n cytia of the e n amel

p ulp a n d the m ulti nu cleated gia n t cells of red bo n e marrow is - -

striki n g These two types of gian t cells have howe ver an

.
, ,

e ntirely differe n t origi n an d f un ctio n b u t i n th eir later di si nt e ,

g rat i v e stages they prese nt comparable aggregatio n s of ebro

m o som e sim ulacra derived i n very similar ma nn er i n both
-
, '

i n sta n ces an d clearly i n both cases i ndicative of dege n eratio n

,
.

S UMMA R Y

1 . The red bo n e marrow of the rabbit and the g ui n ea pig

- —

co n tai n s three chief varieties of gia nt cells : megakaryocytes the —

,

m o n on u cleated forms derived by excessive growth from the

hemoblast ; p olym o rphok aryo cyt es the forms with p olym o r ,

pho n s or lob ulated n u cle u s derived from the megakaryocyte by

i n co m plete direct divisio n of the nu cle us a n d p olyk aryocyt es , ,

the m ulti nu cleated forms derived fro m forms with lob ulated
x v
n u cle i by c o m p l et l o n of the c o n st r l c t i o n s l n the co m plex mu c l eu s

to prod u ce separate smaller an d spheroidal n u clei Comparable .

varieties with si m ilar ge netic history occ ur also i n the yolk sac -

of the pig embryo .

2 Certai n gia n t cells of the m u lti n u cleated variety c h i e fly

.
-
,

b i or q u ad ri nu c l eat ed retai n to a small degree the capacity of

,

their origi nal he m oblast a n cestor of prod u ci n g i n tracell ular

erythrocytes In these gia nt cells erythrocytes may di ffere n
.
-

t i a t e i ntracell ularly abo u t isolated n u clei .

 GIANT C ELLS
—
or B ON E MA R ROW
-
309

3 . T he gia nt cells
-
ormal marrow a n d i n the yolk sac are
in n -

devoid of phagocytic f un ctio n The prese n ce of p olymo r

.

p h o nu cl ea t e d gra n u locytes of the special eosi n O p hi li c variety

i n greater or less ab u n da n ce withi n the cytoplasm do n o t sig n ify
i ntracell u lar di ffere n tiatio n n o r phagocytic i n gestio n b u t are to ,

be i n terpreted as seco n dary passive i n vasio n s d ue to the rela

t i v ely less resista nt character of the gia nt cell cytoplasm as -

compared with the ge n eral marrow pare n chym a agai n st the

press u re exerted by an area of actively proliferati n g le u cocytes ,

or as active i n vasio n s by special p ol ym o rph onu cleat ed phago

cytes of a disi n tegrati n g cytoplasm .

.4 The gia n t cells do n o t u n dergo mitosis either complete ,

cytoplasmic or i ncomplete n u clear The gro u ps of chromatic

.

particles simu lati n g plates of chromosomes of m u ltipolar spi n dles

of pathologic tiss u es are simply chan ce aggregatio n s of chroma
t i n d u ri n g the dege n erative process These aggregatio n s are
.

no t acco m pa n ied by archoplasmic fi b ers They represe n t .

termi n al stages i n the several modes of dege n eratio n of the

p yo l m o rp o nu cl eat ed an d m u lti n u cleated varieties of gia n t cells
h -
.

.5 The lob u latio n an d direct divisio n of the gia n t cell n u cle u s ; —

the accompan yi n g partitio n of the ce n trosome i n to a pl uri

corp u sc u lar eleme n t ; the later dissol u tio n of the n u clei and the
formatio n of chromati n masses an d gro u ps of s u ch masses ; an d
the coi n cide n t prod u ctio n of blood platelets from the cytoplasm
-
,

fi rst by seg m e n tatio n of pse u dopods an d later by mass f rag m en

t at i o n of larger cytoplas m ic areas are all to be i n terpreted i n
,

terms of the actio n of some commo n u n k n ow n factor worki n g

toward the eve n t u al disi n tegratio n of these cells .

.6 There is n o adeq u ate evide n ce to s upport the claim that the

hemoge n ic gian t cells of red bo n e marrow have a phagocytic
- -

f un ctio n or that a ge n ui n e mitotic mechan is m un derlies the

,

i n crease i n co m plexity or the segme n tatio n of the polymorpho u s

n u cle u s . O n ce havi n g passed beyo n d the stage with feat u res
characteristic of the hemoblast their f u rther history o n ly leads
,

thro u gh progressive steps toward disi ntegratio n i n volvi n g ,

termi n ally i n some cases n u clear appearan ces simu lati n g mu lti
polar mitotic fig ures .
3 10 H . E . JOR D AN

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, ,

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e s o o

s t o ly t i c f o i
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s o o -

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, . . .
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’
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,

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 13 Simil a r c el l a t a c o m p a ra b le st a g f e, ro m t he a n gi b l o a st la y er of t he y lko

sa c o f a 10-m m . pig em b ry o . Z en k er fi x t i a o n ; he m a t o xyli n a n d e os n s t a n s i i .

14 a nd 15 S u c c e s s w e l y l a t er sta g es i n t he p ro c ess o f d i i nt g t i
s e ra on m i t i a t ed
i n fi g u re s 1 1 a n d 1 2 . I n t he i i
ser a t o n o f m i tot i c sta g es d ib d f
escr e or t h e se cel l s

by D y
en s, fi g u re 1 5 re p r es e n t s a n i n t erm ed i a t e sta g e b e t w ee n fi g
u re 6 a nd fi g u re 5 N ithe er c e n t r o s o m es n o r s p i n d l e

fi b e rs a re d i s c e rn ib l e .

16 A v i ty
p ly k y y t
ar e whi h m
o f y f th o l i l t iv ly
ar oc e in c an o e n uc e a re r e a e

s m ll a T hi l l r p d w t h t g 1 m m i t i d i b d b y D i k
. s ce co r es on s 1 s a e os s as esc r e c son .

17 V y v i t y f p lyk y y t i w h i h l l f t h
er r are ar e l i o f o ar oc e, n c a o e n uc e a re o

a pp x i m t l y if m i z d l t i v l y v y m l l T h i t y p f l l
ro a e un or s e an re a e er s a . s e o ce c o r re

d wit h t g 2 f D i k i ti f t p i mit i
’
s p on s s a e o c s on s s e r a on o s e s n os s .

18 L t t g t h di i t g t i v p
a er s a e in h w b gi i g i fig 1 7
e s n e ra e ro c e s s a s s o n e nn n n ure .

A w f m ll
ro o l i s t i ll d i
a ib l t t h i g h t A t t h l f t t h h m t i
nu c e is s s c e rn e a e r . e e e c ro a n

o f th l h
e n u c e1 l dt f m v y as co a esce d i g l h m ti tw k
o or a er c o ars e a n rr e u ar c ro a c ne or

or p i m T h i typ f l i lm t b
‘
S re e .
’
s f yt p l m e o n u c eu s s a os a re o c o as .

1 9 St ill l t t g i th p a er s a f di mt g t i f thi gi
e nt ll T hi e r o c e ss o s e ra on o s an -
ce . s

t yp f e o l q t k d Th yt pl m h b m
n u c eu s 1 8 ul d p i th p
e na e . e c o as as ec o e u se u n e ro

d ti
uc f blon o d pl t l t oo -
a e e s .

20 A v i t y f p l ym p h k y y t i w h i h t h l b l t d u l
ar e o o or o ar oc e n c e o u a e n c eu s

a pp h m g
ea rs f
o th m o t p t
e n eo u s d t i v y i t ly T hi t p f
or e os ar an s a ns er n e n se . s y e o

l
n u c eu s ig fi b gi i g d mt g ti
s ni es S h
e ll p d
nn n pl t l t by ls e ra on . uc a ce ro u c es a e e s

f gm t t i
ra en f l g a on f t o y t pl m T hi t yp f ll
ar e a rea s p d o i s c o as . s e o ce c or r es on s

Wi th d ti t ph a ls f th l nc i i f th gi t l l
a se o d ib d by A l d e s en o ese an -
ce s as e sc r e rn o

as
‘
f r a g m en t a t i o n
m d i re c t .
’

2 1 a n d 22 S u c c e s s i v el y l a t er st a g es i n t he p r o c ess of di i s nte g rat on i of th i s

v a ri ety o f c el l l e a d i n g t o ,
a na kd e an d ul t im a t el y f gm t i g u l u
ra en n n c e s . T he
n u c l eu s i n fi g u re 2 1 i s en v l p d by
e o e onl y a n a rr o w la y f y t pl m f gm
er o c o as ,
ra en t n i g
p iph
er e ra l l y .

23 an d 24 S u c c ess uv e l y l a t e r s t a g es i n t he p r o c ess o f di i n t g
s e i
ra t o n o f o r i gin
all y p ly m p h o or o us n u c le i ,
co m p a c te d and m o d i fi e d t h ro u gh l o ss o f c y t op l a sm in
t he f or mat i o n of pl a t el e t s . Fi g u re 23 is p
c o m ara b l e t o 1 8 ,
and m ay b e d iv d er e

f ro m c el ls l ike t h o se of f g u re s
i 16 and 17 . Fig u re 24 i s al so c om p a ra b le t o fi g ure
18 , a n d re p re s e n t s a v i ty
i s t y p e o f n u c l eu s
ar e of th .

2 5 Mu l t i n u c l e a e an t d gi
c e l l f ro m t h e e n a m e l p u l p o f a n ew b o rn c a t
t -
T he se — .

c e l l s a r e f o r m e d by t h e f u s i o n o f o r g m a l l y d i s c r e t e m o n o n u c l e a t e d c e l l s o f t h e i ,

e n am e l o r g a n T hey m a y c o n t a i n g l o b u l es o f re s o rb e d en a mel T h e c y t op l a s m
. .

i s a c i d o p h i l i c m st ai n i n g r ea c t i o n Z en k e r fi x a t i o n ; h em at o x y l i n a n d e o s m s t a i n s
. .

2 6 E a r l y s t a g e m t h e d i s i n t e g r a t i o n o f t h ese e n a m e l g i a n t ee l l s T he c h r o -
.

m a t i n f t h e n u c l e i c o ll e c t s n s e v e ra l p e r i p h e ra l c r es c e n t i c m a ss e s a nd o c c a
o i

s o n a l sp he i c a l
r e lo n g a t e d d ro p l e t s
r or T h e n u c l e 1 b e c m e p a l e t h e i r m e m b ra n e . o ,

e v e n t u a l l y r u p t u r e s a n d t h e c h ro m a t i c m a ss e s a r e c a s t o u t i n t o t h e c y t o p l as m o f
,

t he d i s i n t e g ra t m g c e ll s w h e re t h e y i mu ls ate p ph
ro m o s o mes
a s e c h ro .

27 L at e s t a g e i n t h e d i s m t eg ra t i o n p r o c ess . T h e c h r o m a t c ma sse s ( c h r o m o i
some s im ul a c ra ) e v en t ual l y di ss o l v e w th i i n t he f g
ra m e n t n c y t o l asm i g p .
R es u me n por el a utor ,
Ch i k an o su ke O gawa .

L as fi n as ra m i fi c a c i o n es del p ulm on h u ma n o .

L os co n d u ctos alveolares del p u lm on ad u lto del hombre fu ero n

est u diados por el au tor por medio de rec o n st ru c c i on es L os c o n .
°

d u etos alveolares se divide n dos a n u eve veces a n tes de alca n zar

los sacos aéreos Como tipo de r am i fi c a ci én se e n c u e n tra n la
.

mo n opodia y la dicotom i a L os pla n os diverge n tes de los c o n.

d u etos frec u e n teme n te se cru za n e n tre si Se r a m i fi c an en su .

c esi é n frec u e n te y la d i sm i nu c i é n de su di ametro n o se ac usa

, .

L a regla de J u stese n n o p u ede aplicarse a l caso del p ul m é n h u

ma n o ad ulto El n umero de al v eo l o s co n te n idos en u n saco
.

aéreo var i a e n tre ci n co y vei nte El atrio de Miller es u n t er .

mi n o i nn ecesario por lo me n os en el caso del pu l m é n h u ma n o

,
.

L as paredes alveolares ( disti n tas de las de la base ) est on ge n e

ral m en t e formadas por la pared salie n te de los co n d u ctos alveo

la re s y los tabiq u es alveolares .

T 1 a n s la t l o n by J os éF N o m d ez
C m ne ll U iv it
n e rs y Me di c a l C o ll eg e , N Y
3 16 C H IKA N O S U K E OGAWA

4 . R eco n stru ctio n ( wax plate modeli n g ) -

.

To my k n owledge o nly M iller L ag u esse an d Keil have st u die d

, , ,
.

the l u n g by this method J u stese n shows this method to be .

based o n fo u r man ip ulatio n s a n d he believes that d uri n g the

excisio n a n d faste n i n g of the wax plates serio u s mistakes are
n ecessarily made i n the reco n str u ctio n of alveolar d u cts ; b u t

I a m of the opposite opi n io n .

In the reco n str u ctio n of wax models it is most importa n t to

decide whether the models will be made i n positive or n egative
c a st ( I mea n by the word positive a cast which prese nts the
.
‘ ’

l um i n a of the alveolar d u cts hollow s u ch as o ne fi n d s them i n ,

the act u al shape of the str u ct u re ) A fter carefu l co n sideratio n .

,

I e n deavored to co n str u ct both positive an d n egative models .

I will give a brief descriptio n of these M y materials co n sisted .

of two differe n t l u n gs : o ne ( cadaver n o 1 3 99 ) was take n from a .

thirty on e year old exec u ted m an an d w as fix ed by formol alcohol

— - - —

this was i n j ected i n to the trachea ; the other ( cadaver n o 2 8 54) .

was take n fro m a fift y hy e year old m an whose whole body was
- - -

fix ed by fo rmol alcohol i n j ected i n to the femoral artery

—
In .

this case the l u n g was fix ed i n n ormal positio n withi n the thorax .

I stai n ed the tiss u e i n toto by VVei g ert s iro n haematoxyli n an d ’

-

embedded it i n p araffin I u sed material n o 1 399 for the positive

. .

model a n d c u t it perpe n dic u lar to the l un g s u rface 2 0 thick u

,
.

For the n egative model n o 2 8 54 was u sed an d cu t 1 5 u thick .

,

parallel to the l u n g s u rface For the positive model I made .

plates from refi n ed beeswax i n which the p ar affin was mixed o n e

part to fo u r of wax T u rpe n ti n e oil was also added i n small
.

q u a n tity S u ch plates are solid n o t brittle a n d n o t cru mbly

.
, .

O rie n tatio n pla n e was n o t u sed for the model Th us I g ot a .

model m ag n i fied 8 0 times an d meas u ri n g cm i n height .

,

2 4 cm i n le n gth a n d 2 0 cm i n breadth
.
,
I c ut t he model i n . .

fo u r pieces parallel to the l u n g s urface by mea n s of a wire saw .

I made u se of u n refin ed beeswax i n the co n stru ctio n of the

n egative model m ag n i fyi n g it a h u n dred times so that it meas u red
,

8 cm i n height 1 2 cm i n le n gth a n d 8 cm i n breadth a n d it

.
,
.
,
.

co n tai n ed two air sacs ( figs 1 2 a nd —

.
, .
 Fl g . 1 C ut pl an e of p it iv
os e w ax -
pl at e mo d el . O ri gi n al mo d el 80 X e nl a r g ed .

Fig u r e re du d ce t o 50 X . C t p
u ar a ll el t o t he pl eu r a , 3 cm . a p a rt f ro m it . Pl eu r a l s id e .

Th e l a r g er ca vit i es s ho w n i n t he fi gu re a re a i r- s a c s . Th e s m a ll e r r e c e s ses i n t he i r

w a ll s a re a l v li eo .

Fig . 2 C t pl
u an e o f the p it iv
os e w ax -
pl ate mo d el . Mag n i fi c a t i on sa me as fig ure 1 .

3 cm . a p art f rom t he f or mer cut pl an e . Pl eu ra l s id e . The la r g er ca vit i es a re

3 18 CH I K A N O S U K E
.
OGAWA
DIVISI O N O F THE FI NER RESPIRAT O R Y TUBULES
A ccordi n g to B N A there are two divisio n s of the fi n er
. . .

respiratory passages n amely bro n chiol u s respiratoriu s an d du e

, ,

t u l u s alveolaris The space from which the en d bra n ches divide

.

themselves is desig n ated by Miller the atriu m This n ame is .

adopted i n several text books Several n ames are proposed for -

.

the en d bra n ches of the alveolar d u cts sacc ul u s alveolaris —

,

Fig . 3 N g t iv
e a e mo d l in
e cl u di n g t w o a i r- sa c s a dj a cen t to pl e u ra . Or i gi n al

mo d el 1 00 X e n l ar gd e . Fig u re re d dt
uce o 50 X .

i nfu n dib ul um sac air sac an d so o n Some au thors ( Cru

,
e n d- ,
—
,
.

V eill i er an d Brass ) u se the word i n fu n dibu lu m loosely for ‘ ’

d u ct u l u s alveolaris Fu rthermore N icolas an d Test ut rather

.
,

u n n ecessarily give the n ame vestib u le to the space betwee n the

bro n chiol u s respiratori u s an d the d uct u l u s alveolaris .

N ow I will co me to the detailed d escri ptio n of the alveol ar

duct s .
3 20 C H I KA N O S U K E OGAWA

Becau se the alveo l ar d u cts ofte n ramify s u ccessively a n d the

walls are irreg ularly shaped it is di ffi cu l t to ascertai n the type ,

of ra m i fi c a t i o n But after caref ul exa m i n atio n I recog n ize the

.

p rese n ce of both dichoto m y a n d mo n opody types predomi n ati n g

F i gs . 4 and 5 Di g a rams o f p j i
ro e c t o n o f the al v e ol a r du cts i n t h e p o s It i v e
mo d el . Ma g n i fi c a t i o n sa me a s a b o v e fi g ur e s . T hi n l i n es d eno t n i ga i r s a es
-
.

D o t te d lin d es en o t ni g co v e re dp a rt s .

in alveolar d u cts The accompa nyi n g diagram (fi g 6) shows

. .

the type of ram i fic at i o n i n my model D otted li n es represe n t .

the parts where the reco n str u ctio n is n o t complete In my .

diagram the a ngle an d le n gth of the bra n ches are n ot co n sidered

with care .
 H U MAN LUN G 32 1

A d i v erg m g pla n e of certai n bran ches of the alveolar d u cts

is i n d efi ni t e relatio n to the n ext i n series A ccordi n g to Waters .
,

i n the eve n t of a bra n ch o f the bro n chial t u be dividi n g twice

‘

i n s u ccessio n ,
divergi n g plan e of the fi rst is perpe n dic ul ar

to that seco n d J u st esen st u died this relatio n

. called
atte n tio n to the fact that the plan es sometimes cu t each other
un der an an gle of This stateme n t appears to refer to the
bro n chial system an d n ot to alveolar d u cts N ow i n my model .

the divergi n g plan es of alveolar du cts frequ e n tly cross each other ,

b u t they occasio n ally lie i n the sam e pla n e .

TH E AM E R I C AN J O U RNA L O F AN \ T O M Y V OL
, . 27 , NO . 3
322 C H I KA N O S U K E OGAWA
DIA METER AN D LEN G TH OF A LV EO L A R DU C TS
I mean by the term diameter the distan ce betwee n the ed g es
of two opposite alveolar septa and by the term le n gth th e le n gth , ,

of each bran ch of the al veolar d u cts By thu s measurin g both .

I i n ten d to show how freq ue n tly the bifurcatio n s i n alveolar

d u cts take p l ace .

L
TAB E 1

L ENG T H D I AM ET E R
M i ll i m et e rs N um b er o f t i m es Mil li m et ers . N um b er o f t i m es .

Av e 0 24. T o t al 39

A v e 0 32
. T o t al 40

A bran ch of alveolar d u cts is n ot always thicker than the

bran ches of the n ext order In other words on divisio n t he .
,

d ecrease i n the diam eter of the alveolar d u cts is n ot marked .

Schulze p uts the diameter of alveolar d u cts from mm to .

mm K( i lli ker gives the average of

‘

mm .
3 24 C H I KA N O S U K E OGAWA

wall is co n caved on: o n e side and protr udes correspo n di n gly on

the other side or it may be totally flat on both sides This
,
.

flat portio n is n at u rally n ot beset with alveoli .

8
Fi g . 6 Di g
a ra m o f ra v du t
m i fi c a t i o n o f t h e al eo l a r c s .

Fi g . 7 Di g
a v
ram o f c o mm o n w a l l o f t h e al eo l ar du tc s .

Fi g . 8 Di g
a i v i
ra m o f s ec t o n o f a l e ol .
 H UMAN LUN G 325

J U S TES EN S R ULE
’

In the embryo n ic an d adu lt ox J ust esen fo un d the followi n g ,

r ule co n cerni n g the n umber of bro n chial t ubes : the n umber of

side bran ches i n each case correspo n ds to those of the co nti nu
atio n of the tr unk an d its bran ches For i n stan ce take a bro n chi .
,

ol u s whose fi rst bran ch bears eight air sacs The n the total -
.

nu mber of air sacs attached to the co n ti nu atio n of the bro n chiol u s

-

are also eight H e gives a skillful expla n atio n of this fact

. .

N ow I have applied J u st esen s r u le to my model and fi n d that

’

it does n ot hold for the alveolar d u cts of the h u man l u n g .

T H E A IR - SA C
Sin ce ( accordi n g to O ppel ) R ossig n ol made u se of the term
i n fun dib u l um for the last divisio n s of the respiratory air t ubes ,

th e shape of this part has attracted the attentio n of man y

a u thors .

A ccordi n g to J u stese n the air sac ge n erally arises from the

,
-

atriu m with a relatively n arrow ope n in g an d the n it expan ds

remarkably O ppel agrees that the air sac takes an i nfun dib u lar
.
-

form O u the co ntrary He nle Stieda an d E b n er do n ot ack n o w l

.
, , ,

edge this L agu esse an d d H ardi v ill er poi n t out that the regio n
.
’

of the air sac is ofte n n arrowed I n otice at times a certai n

-
.

degree of exte n sio n i n the air sacs b u t I do n ot thi n k this exte n

-
,

sio n is e n o u gh to warran t the n ame i nf un dib ul um .

Miller J u stese n and Schulze have show n that the alveolar

, ,

septa are elo n gated to a certain exte n t i n the regio n of the air
sac an d divide it i n to two or three parts In case the regio n of .

the air sacs is separated i n these parts the elo n gated septa sta n d
-
,

parallel or perpe n dic u lar to each other O n the other ha n d .

,

whe n there is a partitio n i n to fo ur they cross each other The , .

alveoli may or may n ot be i n cl u ded i n these parts In the latter .

case the parts represe nt n othi n g more than deep alveoli .

N ow I will show the relatio n of the air sacs to the alveolar -

d u cts If i n my model I follow the alveolar d u cts forward they

.
,

u s u all y seem to divide themselves i n to two air sacs an d some -

,

t imes i n to three Bu t whe n the air sacs are se nt o ff as side

.
-
3 26 CH I KA N O S U K E OGAWA

bran ches there may remai n b ut on e R eferri n g to fig ure 6 .

,

on e may gai n a clear co n ceptio n of this matter A ccordi n g to .

Miller two to fi v e air sacs are se n t off from the atrium J u st esen
,
—
.

states that i n the 8 0 cm ex embry o fo u r air sacs an d i n the ad ult

-
.
-

three air sacs are se nt off from the atri um The nu mber of
-
.

alveoli co n tai n ed i n on e air sac is estimated to be from 1 0 to

-

20 by R ossig n ol My calc ulatio n is as follows

.

The nu mber of alveoli 5, 8 , 9 , 1 0 , 1 1 , 1 3 , 1 6, 1 9 2 0 ,

1 1, avera g e
The n u mber of times
.

1, 1, 2, 2, 2, 1,

ATRIU M
Sin ce Miller pr eposed the n ame atriu m i n n ome n clat u re for , ,

the n ew aspect he fo un d i n the respiratory t ubes of the do g

l un g by mean s of wax plate reco n stru ctio n man y authors have
—
,

bee n disc u ssi n g it witho u t comin g to a commo n un derstan din g .

I will first b ri efly qu ote Miller s statements ’

B etween e ach air sac and termi nal bron chu s there i s a c avity con
-
,

st an t i n all porti on s of the lun g whi ch I sh all term atri um The

, , .

term in al bron chu s is m ade up of sm o oth mus cle fib ers lined with ,

cyli n dric al epitheli al c ell s from the t i p of the termin al bron chus three
,

t o six ve stib u le s arise .

( Vestibules can be u sed i nter chang e ably with
a tri um be cause he with d rew the term ve stib ule i n a l ater pu b li cati on ) .

The di am eter of the atri um i s sli ghtly m ore th an half that of the ai r sac -

an d un like i t e a ch atri um h as three o r m ore o pe ni n g s The se O pen .

ing s c omm uni cate on the on e hand with the ai r sacs an d on the other -

with the termin al bron chus .

Si n ce Miller s so called termin al bro n chu s is li n ed by cyli n

’
-

d ri c al epithelial cells it o u ght to be the part of the bro n chial

,

t u be above th e bro n chiol u s respiratoriu s b ut accordin g to his ,

later pu blicatio n s it is said to be equ ivale n t to the alveolar du cts .

The fact that authors un derstan d the term atriu m i n di ffere n t

ways may be due to this i n acc urate descriptio n N ow B6hm .

D avido ff S p alt eh ol z J orda n Ferg u so n Morris Piersol Sh afer

, ,
-
, , , ,

an d others adopt this term i n their text books an d G ray co n -

,

siders it the same as an alveolar d u ct V E b n e r i n a w ay . .

opposes this as the term atrium has n o histological si g n ifi can c e

,

an d aro u ses o n ly co n f u sio n i n termi n ology O ppel opp osm g .

 C H I KA N O S U K E OGAWA

d ucts or the dividi n g parts of s u ch d u cts Si n ce the alveola r .

d u cts divide frequ e ntly alo n g their periphery the le n gths of ,

bran ches are accordin gly short These do n o t take the t u b u lar .

form In other words the formatio n of these n on t u b u lar spaces

.
,
-

is the i n evitable resu lt of frequ e nt ram ifi cat i o n I f urther see .

i n my model that some air sacs arise directly from alveolar d u cts
-
,

an d that o n the other ha n d

,
there are n o n t u b ular air spaces
,
—

i n the more ce n tral part of the alveolar d u cts .

Thu s I thin k t h e term atri u m lacks e n o u gh si g n ifi can c e to

j ustify its u se at least for the h uman l un g
, .

ALVE O LI
a . The shape of alveoli has bee n called polygo n al
S tru ctu re .
,

half spherical or irreg ular Alveoli are always polygo n al i n

, .

l un gs which have bee n adequ ately in j ected with fi x i n g fl u id by

way of the trachea or in j ected thro u gh the artery whil e the l ungs
are still i n positio n withi n the un ope n ed thorax Becau se of .

the irreg ular shape of alveoli it is ofte n difficult to determi n e ,

how man y su rfaces they have Schulze calls atte ntio n to the .

fact that i n several mammals the surfaces rarely n u mber over

twelve an d vary i n gen eral from five to twelve My models
,
.

show that the h u man l un g has alveoli with hy e s urfaces i n most

an d i n a comparatively few cases there are six A ltho u gh .

the shape of alveoli is irreg ular the formatio n of alveolar walls ,

seem to co n form to fix ed types I shall classify alveoli i n .

three types F ig ure 8 represe nts diagrams of sectio n ed alveoli

. .

Thi n li n es de n ote alveolar septa an d the thick o n es the alveolar

walls beside the septa .

Type A This ki n d of alveol u s is formed by the proj ectio n

.

of the walls of alveolar d u cts an d it lacks special septa .

Type B This ki n d of alveol u s is e n closed by alveolar septa

.

alo n e except at the base

,
.

Type C This type is an i ntermediate form betwee n the other

.

two an d prese nts variatio n s a ) O ne side may co n sist of the .

co n cave wall of the alveolar d u ct and the other of the alveolar

sept um 5) O n e side may co n sist of the co n cave wall a n d the
.
 HU MA N L U N G 32 9

alveolar sept u m while the other side may have o n ly the alveolar
,

sept u m c) The alveol u s may be e ntirely closed by both the

.

alveolar sept u m an d the co n cave wall of the alveolar d u ct

“ ”
.

( S u ch a n expressio n as the co n cave wall of the alveolar d u ct

may n o t be adequ ate b ut o n e can easily u n dersta n d what I
,

mean ) Type A an d type B are i n fact rarely see n the greater

.
,

nu mbers are of the C type Schu lze co n siders that the mo u th

.

is smaller than the l ume n b u t I n otice that the mo u th is some ,

times larger an d at other times smaller .

T ABL E 2 TABL E 3

D EP H N U M E R R
B
E A D H N U M E R D E P H R E
B
A D H
TI M ES
OF
TI M E S
OF
T B T T B T

mm mm mm

Av e . T o t al 61
Av e 0 1 5
.

Av e . T o t al 61

b. The si ze of a lveoli the alveoli

. A ccordi n g to K olli k er ,

collapsed after death are on e sixth on e te n th an d on e eight -

,
-
,
-

een t h the size of the n ormal Schu lze calls atte n tio n to the .

fact that alveoli are of differe n t sizes i n the vario u s parts of the
l un g n amely alveoli located n ear the s u rface of the l un g espe
, , ,

c i al ly i n the apex a n d edge are larger tha n those of the i n n er

part Sch u lze an d R ossig n ol say that the size of these alveoli
.

i n creases with age R ossig n ol gives the measu reme n ts of alveoli

.

accordi n g to ages as follows : mm for the ages between .

eightee n an d twen ty to for twe n ty fi ve years

,
-
,
33 0 CH I K A N O S U K E OGAWA

for thirty five to forty

-
to mm for seve n ty to eighty
,
. .

In Sch u lze s acco u n t he gives to mm for the n ew bor n

’
.
-
,

for middle age an d those to

,
mm i n breadth an d .

to mm i n depth for old age

. .

I measured i n my positive model from the thirty on e year old - - -

m an the alveoli i n several parts an d fo u n d the res u lts as give n

i n table 2 .

Thu s the alveoli i n my model o n the average measure mm .

i n both breadth an d depth .

The results obtai n ed from the measu remen t of the alveoli of

my n egative model from the fi ft y six year old m an are as show n
— - -

i n table 3 .

The alveoli here measu re mm i n depth an d mm . .

i n breadth A ccordi n gly these are larger than those of the

.
,

positive model Bu t I cann ot tell if this d iffere n ce comes from

.

age the part of the lun g stu died an d the degree of expan sio n of
,

the alveoli A ccordin g to R ossig n ol an d Miller the alveoli i n

.
,

the bases of air sacs are larger tha n the others I have also
-
.

n oticed this fact i n my model b u t this is n o t the i n variable case

,
.

SU MMAR Y
1 .The alveolar du cts divide two or n i n e times to reach the
air sacs
-
.

2 The a n gles of r am i fi c at i o n of the alveolar du cts are of

.

variou s grades ra n gin g from wide a n gle to ac u te A s type of .

r am i fi c at i o n both mo n opody and dichotomy are presen t i n

a n alveolar d u ct system
—
.

3 D ivergi n g plan es of alveolar du cts frequ e n tly cross each

.

other .

4 The alveolar d u cts bra n ch i n frequ e n t s u ccessio n the de

.
,

crease i n the diameter is n ot marked .

5 The mo u ths of the alveolar d u cts co n sist of alveolar septa

.
,

b ut u su ally the mo uths are e n closed partially by the wall of the

alveolar du cts itself .

6 J ust esen s r u le that the nu mber of side bra n ches correspo n ds

’
.

to those of the conti nu atio n of the tru n k and its bran ch is n ot ,

tru e for the alveo l ar du cts of t h e adult m an .

332 C H IK A N O S U KE OGAWA
MU LL E R 1 9 07 Z ur v gl i h d er e c en en Hi st o lo gi e d er L g un en u n se r er H i
a u s t e re .

A rc h f m ik
. A t mi r . na o c u . En t w .
-
g es c h c h t e , i Bd 69 . .

MER a 1 90 7 Egb i d A
r e n ss e er n at o m e u i . E n tw c i k l un g g s i
es c h c h t e , Bd 69 . .

1 9 02 A t h m u n g so r g a n e ( Ba rd e l eb en
’
s H dbu h an c d er A n a t o m i e d es

M e n s c h en ) .

O PPEL 1 9 01Egb i r e n ss e A t mi
d er E na o e u . n tw .
-
g i
e sch c h t e , Bd 1 0 . .

1 905 L h b e r uc h d v gl i h d
er er e c en en m i k ro s k . A n a t o m i e Bd 6 , . .

PI E RSO L H m A t
u an na o m y 5t h E di t i
,
on .

R o ss rc N O L 1 8 47 R eser c h e s su r la s t r u c t u re i nt im e du p
mo n d e l h o mm e et
ou
’

d es p i ip x
r nc au m a mm i f er e s ( O pp el : L eh r b u c h d er er l m i kro s k v g . .

A n at om i e d .

SCH UL Z E 1 87 1 Die L g un en ( S t r i c k er
’
s L eh r e v o n d en G ew eb en ) .

S CH KF E R T xt b k
e oo of m i c ro s0 0 p i c an at om y , Q ua in
’
s El em en t s of A n a t om y ,

vo l . 2, p . 1 .

S HUL
C ZE 1 90 6 B it g e A t mira e z ur na o e d e r Sa ug e t i e rl un g en . S it z g b i un s er c h t

kgl . p Ak d m i d Wi
r eu ss . a e e . ss .

S I DA
T E 1 878 Ei ig b d B
n es u er en au un d E tw i kl g
n c un d e r Sa u g et i e rl un g e .

Z e i t s ch r f . . W i ss Z o o l o gi e
.
,
Bd 30 .
, S pp l m t
u e en .

SU S S DO R F 191 1 D er R pi es i
r a t o n sa pp a ra t (El l enb er g er
’
s H db an u ch d er ver

gl ie c h en d n e m i k ro sk n a t o m e d er. A i H au st er e , i Bd .

T LD
O T 1 888 L h bu
e r c h d e r G ew e e l eh r e b .

W A TE R S 1 8 60 T he an a t o m y
o f t h e h u m a n l un g ( J u st es en an d O pp el ) .
R esu me n por el au tor ,
Chikan o suk e O gawa .

Co ntrib u cio n es a la h i st ol og i a de los espa cios respiratorios de los

pulmo n es de los vertebrados .

Em el prese n te trabaj o se describe el epitelio respiratorio fib ras ,

mu sc u lares fib r as elasti c as y fi b ras retic u lares de los p ul mo n es

, ,

de los anfibi o s ( Di emyc tylus ran a salaman dra gigan te ) reptiles , , ,

( tortu ga gecko c ulebra) aves (paloma gall i n a salvaj e pato

, , , , ,

g orri én) m am if er os ( topo murmel ag o rata co n ej illo d e i n dias

, , , , ,

cabra co n ej o gato perro y hombre ) asi como la membran a pro

, , , ,

pia y los poros alveolares de los p ulmo n es de los m amiferos El .

epitelio respiratorio de los reptiles oc upa un a p o si ci on i n termedia

en tre el de los anfib i os y m am if eros En las aves topo y mu r .
,

ci él ag o el epitelio respiratorio parece faltar En cl emb ri 6n de .

co n ej o esta formado ai m an tes del n acimien to por dos clases de

’

células L a desap araci én de los n ucl eos de las células plan as tiene
.

lugar gradu al men te en cl estado fi n al del desarrollo L a repa .

r aci é n de las cél u las resp i ratorias pla n as parece te n er l u g ar p or

la ex t ensi én de pequ e nas células y desap ari ci én de los n ucl eos .

L as fib ras mu sc ulares faltan en absolu t e en todas las partes de

los co n ductos alveolares del p ul m on del mu rci elago En otros .

m am if er os ex iste mucha difere n cia en lo refere n te a su pres en cia .

L as fib ras elasti cas y retic ul ares tambié n prese n tan difere n te

di sp osi ci én seg un los a n im ales L os poros alveolares ex iste n .

n ormalme n te en m u chos m am if eros .

T ra nsl at i o n by J é F
os N o m d ez
C o r n ell U i v i t y M di
n e rs e c al C ll g
o e e, N Y .
3 34 C H IK A N O S U K E OGAWA

Previo u s histological researches co n cer n i n g the respiratory ,

spaces of the l un g have bee n c on fi ned to a limited n umber of

,

an imals witho ut m u ch co n sideratio n of the whole vertebrate

,

system In thi s paper I have e n deavored to shed fu rther light

.

o n these problems an d to add to the i n formatio n w e already

have .

I wish to exte n d my heartiest than ks to Prof B S u z uki for . .

m u ch val u able advice I am also deeply i n debted to Prof R R

. . . .

Be n sley and D r G L Streeter for their co u rtesy i n placi n g at

. . .

my disposal the facilities of their departme n ts while tran slati n g

this paper .

M ATERIAL AN D M ETH O DS

The materials u sed for the prese n t st u dy are as follows

A mphibia : D i em y ct yl u s pyrrh og ast er R a n a n i g ro m a culat a , ,

Megalobatrach u s j apo n ic u s
R eptilia : Clemmys j apo n ic u s G ecko j apo n ic u s E laphe q u adri
, ,

Virgata .

A ves : Col umba livia G all u s domestica A n as Passer , ,

.

Mammalia : Talpa V esp eru g o E pimys Cavia c ob aya L ep u s

, , , ,

c u m i c u l u s Capra Felis domestica Ca n is familiaris Hom e

, , , , .

In thi s i n vestigatio n the followi n g were u sed for fi x at i on

p urposes :
T en per ce n t formol formol alcohol potassiu m bichromate , , ,

s u blimate Muller formol an d Flemmin g s solu tio n Silver

,
-
,
’
.

n itrate was u sed to i mpreg n ate the i n tercell u lar s u bsta n ce A .

descriptio n of the tech n iqu e is omitted here as it is give n under

each chapter .

RESPIRAT O R Y EPI T HELIU M

1 . R espi ratory epi theli um i n A mp hi bi a

For the st u dy of the respiratory epitheli u m of Amphibia ,

the l u n g s were fi rst i nj ected with a per ce n t sol utio n of

silver n itrate an d the n c ut ope n an d spread o u t Whe n the .

ceme n t s u bstan ce had become dark the pieces were mo unted .

In the case of R a n a a n d Megalobatrach u s the tiss u e which had ,

bee n impreg n ated i n this w ay w as also imbedded i n celloidi n

 R ESPI R AT O RY SP AC E S or THE L U N G S 3 35

an d cut i n to thin sectio n s In ge n eral the respiratory epitheliu m

.

of the amphibian l u n g is more easily impreg n ated than that of

R eptilia an d Mammalia Besides these preparatio n s co ntrols . _

were made by stai n i n g sectio n s an d also some of the u n c u t tiss u e

with ordi n ary dyes s u ch as hematoxyli n an d eosi n
,
.

Di emyctylu s pyrrhogas ter A ccordi n g to El en z the respiratory .

,

epitheli u m of Trito n ( a ge nu s closely related to D i em yctyl u s) ,

co n sists of large fl at cells whose nu clei are also large an d are

, ,

always located i n the i ntercapillary spaces j oi n i n g each other o n ,

the capillaries I w as able to recog n ize a similar co n ditio n i n

.

D i em yc t yl u s ( fi g .

The blood capillaries of the l un g of this an imal are o n ly wide

-

e n o u gh to admit of the passage of o n e blood corp u scle at a time -

.

n some other a n imals which will be me n tio n ed later the c ap il

I , ,

l ari es are so wide that two or three blood corp u scles can pass -

alo n g side by side The i n tercapillary spaces are of varyi n g

.

widths b u t they are u s u ally of a somewhat larger caliber than

,

the capillaries .

For co n ve n ie n ce i n the descriptio n we m ay disti n g u ish two

parts i n the respiratory epithelial cell o n e of which is located ,

i n the i n tercapillary spaces an d holds the ro u n d nu cle u s while ,

the other spreads o ut flat over the capillaries These will be .

called respectively the nu cleated an d flat portio n s

, ,
‘ ’ ‘ ’
.

Ikm s u rface View the epithelial cell prese nts a pe n tago n al or

hexago n al form The fl at portio n is extremely thi n an d c an
.

n o t be see n i n tra n sverse sectio n s of the fi x ed material while ,

the n u cleated portio n which has the same width as the capillary
, ,

reachi n g i nward to the stroma ge n erally has a c u boidal form , ,

tho u gh its thick n ess differs i n differe n t parts of the l un g som e ,

times becomi n g almost flat .

A s a r ule the flat portio n appears more exte n sive i n s u rface

,

View than the nu cleated portio n b u t i n exceptio n al cases the ,

former covers the capillaries o n ly partially an d is co n sequ e n tly

smaller than the latter O ccasio n ally the flat portio n stretches
.

o u t over the capillaries a n d reaches the n eighbori n g i n tercapillary

space In rare cases a s m all i ntercapillary space may have n o

.

nu cleated portio n an d is covered o n ly by the fl at portio n T

he n . “
TH E AM E R I CAN J O U RNA L OF ANA O M Y V OL
T ,
27 , NO 3
1 R pi
es ra t o r y p i t h li um
e e . D i emy c t yl us j p ni
a o c us . S i lv er i mp g n t i
re a on . S f
ur ace vi ew .

2 R pi
es ra t o r y e p i th l i
e um . R an a n i g r om a c u l a t a . S i lv er i mp gn t i
re a on . S f
u r a ce vi ew .

3 R pi
es ra t o r y e p i th li
e um . Me g a l o b a t ra ch u s j p ni
a o cus . S i lv er imp gn t i
re a on . S f
ur a c e

X 350 .

4 R pi
es ra t o r y p i t h l i um
e e . C l emm y s j p i
a o n cus . S i lv er i mp gn t i
re a on . S f
u r ac e vi ew .
 CH IKA N O S U K E OGAWA

2 . R esp i r a tor y ep i theli um i n R ep ti li a .

The method u sed for the impreg n atio n of reptilia n epitheli u m

was ide ntical with that for amphibia n material The res ults .
,

however were somewhat i n co n siste n t

,
.

A ccordi n g to Elen z the respiratory epitheli u m of the tortoise

,

a n d s n ake co n sists of two ki n ds of epithelial cells n amely small , , ,

nu cleated cells an d fl at o n es i n which he was u n able to demo n,

strate any n u clei If this is the case the respiratory epitheli u m

.
,

of R eptilia mu st be q u ite differe n t from that of A mphibia O h .

the othe r han d s u ch authorities as Sch u lze an d O sawa seem to

,

believe that the r espiratory epitheli um of the two phyla has the
same co n str u ctio n .

Compari n g the two Views it is see n that accordi n g to El en z, , ,

the respiratory epitheli u m of R eptilia is n o t esse n tially di ffer

en t from that of M ammalia si n ce the respiratory epitheli u m of ,

the latter which will be described later is also composed of

, ,

two ki n ds of cells o n e of which is n on n u cleated A ccordi n g

,
—
.

to other au thorities the respiratory epitheliu m of R eptilia an d

,

M ammalia m u st be differe nt Some light is throw n o n this .

qu estio n i n the following acco u n t .

Clemmys j ap on i cu s ( tor toi se) A s is well k n ow n the respiratory

.
,

epitheli um of this a n imal co n sists of small cells an d large flat

o n es ( fi g The small o n es which are u s u ally deeply stai n ed
.
,

by the silver sol utio n take a rhomboidal form an d co n tai n

,

n u clei They may appear isolated or i n gro u ps of two or h y e

.

cells an d they are i n close co nn ectio n with the i n tercapillary

spaces The cell borders which are adj ace n t to the large flat
.
,

cells ofte n c u rve i n ward slightly The i ntercapillary spaces

,
.

are said by El en z to be very n arrow ge n erally an d e n tirely fill ed

u p by small cells i n gro u ps which sometimes exte n d over more
,

than the i n tercapillary spaces I also fo un d that the i nter .

capillary spaces are i n ge n eral exceedi n gly n arrow b ut sometimes

wide spaces s u ch as are see n i n amphibia may appear here an d
, ,

there T he i n tercapillary spaces have a ro un d or an elliptic

.

form bei n g sometimes red uced to a mere slit ; i n rare cases the
,

slit becomes so m u ch elo n gated that the le n gth amo u nts to more
 RE SPI R AT O RY SPAC E S OF TH E LU N G S 33 9

tha n t en times the diameter of the blood capillaries The i n ter -

.

capillary spaces are n o t gro uped i n the l un g accordi n g to their

shape and size b u t o n the co n trary they may prese n t vario u s
, , ,

appearan ces eve n i n the same alveol u s .

Whe n the i n tercapillary spaces are n arrow they are as already , ,

me n tio n ed by Elen z fill ed u p e n tirely by small cells while the

, ,

slit like spaces are li n ed o n ly partly by small cells an d for the

-

most part by flat o n es ; occasio n ally the small cells do n ot lie

e n tirely i n the i ntercapillary spaces b ut are somewhat removed ,

from them Moreover the i ntercapillary spaces are sometimes

.
,

n o t fill ed by small cells b u t by large fi at cells alo n e

, , .

In s u rface view the large flat cells show a sq u are to hexago n al

,

form an d their diameters are from fo u r to hy e times as large as

those of the s m all cells A ltogether i n Amphibia both the ceme n t
.

li n es an d the n u clei of the respiratory epitheli um t u r n black by

i m preg n atio n i n the tortoise o nly the ceme n t li n es are impreg
,

n at ed . A s previo u sly me n tio n ed El en z was n ot able to demo n ,

strate nu clei i n the flat cells In order to clear this qu estio n I .

,

stai n ed silver preparatio n s with n u clear dyes and st udied them

i n s u rface view b ut si n ce the n u clei of s u bepithelial cells wer e
,

stai n ed at the same time it w as diffi cul t to j u dge eve n with

, ,

the greatest care whether or n ot the fl at cells co n tai n ed nu clei

, .

S u bsequ e n tly nu mero u s celloidi n sectio n s were made from s u ch

preparatio n s an d the poi n t at which the epithelial cells were c ut
o ff ta n ge n tially from the u n derlyi n g tiss u e w as caref ully ex

am i n ed . Here where the epithelial cells were free their nu clei

, ,

co uld be see n lyi n g withi n blacke n ed ceme n t li n es either ro un d ,

or elliptic and located someti m es i n the ce ntre sometim es

, ,

ecce n trically ( fi g These observatio n s show that Elen z s

.
’

n egative res u lts were d u e to i n adeq u ate tech n iq u e A s a large .

,

flat cell ofte n covers a n u mber of i ntercapillary spaces at the

same time spaces co n tai n i n g n o n u cle u s are n ot as rare as i n
,

D i em yc t yl us .

Elap he qu a dmvi rga ta ( sn a ke) A s far as I k n ow , El en z is the

’

o n ly o n e w h o has give n a detailed stateme n t of the respiratory

epitheli um of the s n ake which may be qu oted as follows :
,
CH IK A N O S U KE OGAWA
3 42 CH I KA N O S U K E OGAWA

with res p ect to the capillary n etworks is that the two layers of
the capillary n etworks of E laphe comm u n icate with each other
he n ce if capillaries are traced i n s urface V iew they si n k to a
, ,

lower level a nd c an be followed thro u gh to the capillary layer

of other side ( fi g .

N ow we c a n disti n g u ish also i n this a n imal t w o ki n ds of epi

t h eli al cells n amely small n u cleated cells a n d large fi at o n es
, , , , .

The small n u cleated cell is n ot rho m boidal as i n the tortoise

, ,

b ut more or less irreg ular i n its shape tho u gh it may be i n some

,

i n sta n ces desig n ated as pe n tago n al or hexago n al ; i n ge n eral it

belo n gs to the c uboidal category a n d is sit u ated mostly i n the
i n tercapillary spaces .

The large flat cell is co n siderably larger than the small cell
,

i n s urface View mo re th an t en times as large i n diameter

—
.

A ltho u gh the border li n es of the cell are n ot always rectili n ear ,

as is the case i n Clem mys it may be described as approximately

,

s q u are or pe n tago n al A s i n Clem m ys the n u cle u s ca nn ot be

.

demo n strated by the silver method Si n ce i n Clemmys rela

.
,

t i v ely thick blood capillaries lie be n eath the flat cells it happe n s ,

that i n sectio n i n g the epithelial cells are separated at certain

poi nts from the s u bj ace n t tiss u e an d co n sequ e ntly the prese n ce
of the n u clei which belo n g to the epithelial cells c an be c o n fi rmed
, ,

by stai n i n g b u t it is n o t so i n E laphe beca u se here the flat cells

, ,

are co n siderably large an d cover both the blood capillaries an d -

i ntercapillary spaces at the sam e time an d sometimes si n k more

or less i nto the spaces which are fairly wide ; he n ce it is di ffi c ult
to c ut o ff the epithelial cells from the s u bj ace n t tiss u e an d to
decide by n u clear ti nctio n whether or n o t the stai n ed n u clei
belo n g to the epitheli um However i n specime n s which have
.
,

bee n impreg n ated with silver an d mo u n ted witho ut cleari n g , ,

i n s u ch a s u bsta n ce as J ap an ese m illet j elly or glyceri n whose ,

refractio n coefficie nt is far less tha n that of Ca n ada balsam a ,

diff u se precipitatio n of silver is sometimes formed on the s urface

of the epithelial cells or i n the cell bodies In each cell however.
, ,

a clear spherical space un to u ched by the precipitate is prese n t

, , ,

i n which a flat n u cleu s c an be observed whe n the light is cu t ,

dow n .
 R ESPIRAT O R Y SP AC E S OF THE L U N G S 3 43

It is to be co n cl uded from the co n ditio n s res ulti ng from

silver impreg n atio n that these n u clei belo n g to the epithelial
cells an d n ot to the s u bj ace nt tiss u e .

The large flat cells cover either the i n tercapillary spaces alo n e
,

or both the capillaries an d the i n tercapillary spaces ; so m etimes

a cell lies over several i n tercapillary spaces at the sam e time .

The arran geme n t of the respiratory epitheli um is n ot u n iform

i n the differe n t parts of the l un g The i n tercapillary spaces .

n ear the ca u dal sac like portio n a n d of the parts o n both sides
-

of the median li n e of the dorsal aspect are li n ed by gro u p s of

nu mero u s small cells It is o n ly i n these regio n s that I can
.

c o nfi rm El en z s stateme n t Most of the epithelial tiss u e is

’
.

differe n t i n that for the most part the small cells are isolated
, ,

an d whe n they do form gro u ps these co n sist of o n ly a few cells , ,

rarely of a large n u mber The flat cells therefore occ u py the

.

greater part of the field an d ofte n cover the i n tercapillary spaces

also It is possible that the differe n ce betwee n El en z s observa
.
’

tio n s and mi n e c an be acco u n ted for by the fact that he st udied

the stru ct u re of o nly a part of the l un g an d believed it to be
represe n tative of the whole .

Gecko j ap on i eu s U p to this time n o acco u n t seems to have

.
,

bee n give n of the respiratory epitheliu m of G ecko .

J u st as i n the tortoise the tiss u e co n sists of two ki n ds of epi

,

t h el i al cells (fi g The i n tercapillary spaces are of the same

.

width as those of the tortoise an d two or three times as wide as

the diameter of the capillaries The two layers of the capillary .

n etwork of each sept u m do n o t directly comm u n icate with each

other .

The small cells are of vario u s shapes sometimes more or less ,

flatte n ed sometimes irregu larly ro u n ded elliptical elo n gated

, , , ,

sq u are or pe n tago n al and approximately twice as large as those

, ,

of Clemmys an d Elaphe The cell borders are n ot straight

.
,

b u t c u rve i nward more or less ; eve n if o n e border is relatively

straight others are ge n erally c u rved The a n gles of the cell
,
.

borders are ofte n obt u se The s m all cells are fo un d either

.

isolated or i n gro ups Whe n they are fo u n d si ngly each on e

.
,

covers either a small i n tercapillary space completely or a wider

3 44 CH IK A N O S U K E OGAWA

one partially O ccasio n ally a part of the cell exte n ds over the
.

capillary an d sometimes the whole cell rests u po n i t In cases .

where a small cell covers a part of an i n tercapillary space it is ,

located n o t i n the middle of the space b ut ecce n trically again st ,

the capillaries an d is ge n erally elo n gated i n its shape the border

, , ,

which is i n co ntact with the capilla ry bei n g n at u rally o u twardly ,

co nvex an d the opposite border c u rvin g i nwardly so that the

, , ,

cell is somewhat cresce n t shaped an d is always slightly n arrow —

at the e n ds ( right above fi g Whe n two small cells are to

,
.

gether their relatio n to the capillary is the same as that of a

,

si n gle cell The bo un dary li n e betwee n these two cells is n early

.

straight Whe n they e n tirely fi ll an i n tercapillary space the

.
,

shape of the two cells is adapted to the S pace bei n g more or less ,

ro un d i n case they occ u py o n ly a part of the space their short

,

edges are i n co n tact with each other an d correspo n di n g the c u rva

t u re of the capillary they are horseshoe shaped or if somewh at
,
-
, ,

n arrowed at the bo un dary li n e they have the form of a n h o u r ,

glass G ro ups of three cells prese n t as a whole a ro un d elo n

.
, , ,

gated or horseshoe like form

,
—
.

The small cells which fil l up an i n tercapillary space vary u s u ally

from on e to three i n rare cases they amo u n t to more than t en
, .

The large cells are also flat their borders n ot bei n g straight , ,

like those of Clemmys are more or less c u rved an d tho u gh the

, ,

form of the cell is irreg ular it mi ght be described at times as ,

pe n tago n al or hexago n al A s the diameter of the large cells .

are fo u r or hy e times as lo n g as those of the small o n es the ,

differe n ce i n their sizes is co n siderably less than is the case i n

E laphe The large cell un like the small cell n ever prese n t s an
.
, ,

elo n gated form a n d i n vests either a whole i ntercapillary space

or both the space and the capillary i n which case the cell exte n ds ,

over the capillary to the edge of a n eighbori n g i ntercapillary

space an d there meets the cell of that space ; that is to say the ,

cell is bo un ded by the co n to u r of the capillary ; b ut i n some

i n stan ces the cell exte n ds over i n to an adj ace n t space .

In Clemmys an d Elaphe the nu clei of the flat cells as above , ,

me n tio n ed co uld n o t be demo n strated by mean s of the u s u al

,

impreg n atio n with silver b ut those of the small cells were easily
,
3 46 CH IK A N O S U K E OGAWA

nu clei i n the cells of the i n tercapillary space lie sup erfi ci al l v

a n d are n o t S pherical A s already n oted they are ro u n d or
.
,

elliptical i n s urface View b ut i n side View appear somewhat

,

flat the lower parts of the n u clei followi n g the co n to u r of the

,

capillaries o n both sides an d si n ki n g o n ly slightly i n to the i nter

capillary spaces ( fi g T hu s it is obvio u s that the n u clei are
.

sit u ated sup erfi c i ally In attempti n g to fi n d out whether or n ot

.

the cell bodies themselves are also sup erfi c i al I fo u n d that the ,

hematoxyli n eo si n method was u seless as eosi n does n ot differ

—
,

en t i at e the cell body from the s u rro un di n g tiss u e The n for .

,

the p u rpose of differe n tiati n g u n derlyi n g tiss u e from the epi

theli um I u sed Mallory s v an G ieso n s an d Bielschowsky s
,
’
,
’
,
’

methods They also proved u n satisfactory However with

. .
,

Heide n hai n s iro n hematoxyli n the preparatio n s were obtai n ed

’
—
,

which showed disti n ct differe n tiatio n an d proved that the cell

body dips o n ly slightly i n to the i n tercapillary space n ot reachi n g ,

as far as the lower level of the capillaries ( fig It is therefore .

evide n t that the large ce ll s of the tortoise are q u ite fl at while ,

the respiratory epitheliu m of Am phibia i n clu din g n u cleated as ,

well as flat portio n s is n ot q u ite flat This differe n ce is more

,
.

si g n i fi can t tha n may appear at fi r st as will be show n below ,

.

The di ffere n ce becomes more obvio u s whe n w e co n sider the

s n ake an d gecko for here the i n tercapillary spaces are very wide
,

a n d the large cells so fl at eve n i n the spaces that they c an

, ,

hardly be recog n ized i n cross sectio n s At times they are co n—

.

fi n ed e ntirely to the i n tercapillary spaces ; co n sequ e n tly it is

here impossible to disti n g u ish two portio n s tho u gh it is possible ,

i n A mphibia The respiratory epithelium of Mammalia co n sists

.

of small n u cleated an d large n o n nucleated cells an d di ffers from

, ,
-

R eptilia i n that the flat cell of the latter has n o nu cle us The .

phyloge n etic developme n t of respiratory epitheliu m may thu s be

s ummarized as follows : The respiratory epithelium which c on ,

sists of o n ly o n e ki n d of cell i n Amphibia is divided i nto two ,

ki n ds of cells i n R eptilia n amely n u cleated large flat cell an d

, , , ,

c u boidal cell the former bei n g very flat compared with the cells
,

of A mphibia In Mammalia the epitheliu m co n sists of two ki n ds

.

of cells while the nu cle u s is lost i n the flat cell The develop
,
.

me n t of the large fl at cell will be clear fr om fig ures 9 1 0 an d 1 1

, , ,
.
 R ES P IRAT O R Y SP AC E S OF TH E LUN GS 347

4 R esp i ra tory
. ep i theli u m i n A ves

In the respiratory apparat u s of birds the bro n chi w hich are , ,

always located n ear t h e su rface of the lun g give rise to the ,

p ar a b r o n c hi which bra n ch sideways i n to the i n terior of the l u n g .

The p arab ro n chi divide i nto the l un g flu t es whose walls are -

,

fi l l ed with small cavities These cavities have hitherto bee n called

.

bro n chioli b u t it seems to me more fitt i n g to call them flu t e

,

holes From the bottom of each fl u te hole exte n d several small

.
-

ca n als which are tr u e respiratory spaces an d were erro n eo u sly

called alveoli by some au thorities I will desig n ate the ca n als .

as respiratory can alic u li .

While the fl u te holes are li n ed by co n ti nu o u s paveme n t epi

-

theliu m the wall of the respiratory ca n alic u li co n sists of close

,

blood capillaries whose spaces have n o coati n g an d admit free

—
,

passage of i n spired air resu lti n g i n the commu n icatio n of all

,

the respiratory ca n alic u li with each other Betwee n two n eigh .

bori n g respiratory ca n alic u li there are u su ally capillary n etworks

of several layers so that the spaces are highly complicated an d
,

diffi cul t to st u dy microscopically The i ntercapillary spaces have

.

abo u t the same diameter as the blood capillaries —

.

The respiratory epitheliu m of birds has bee n stu died by vario u s

i n vestigators Baer stu died the lu n g of the pigeo n an d came
.

to the co n clu sio n that the i n tercapillary spaces are coated by

delicate fi at Belag while the capillaries themselves are n aked
,
‘
,
’

i n every case A ccordi n g to E berth the blood capillaries of

.
,
-

th e fi n est air passages ( respiratory ca n alic u li ) are especially

-

u n covered with flat cells attached here an d there

,
I thi n k that .

these co n clu sio n s are do u btful for the methods u sed were i n ,

effi c i en t .M ost au thors as Elen z O ppel Sch u lze are of the

, , , ,

opi n io n that the respiratory epithelium of birds thou gh n o t ,

actu ally demo n strated mu st be prese n t ,

.

If the respiratory spaces of birds were bare it wo uld be the ,

o n ly case i n higher a n imals i n which either an exter n al or an

i n ter n al s u rface of the body has n o coati n g of epitheli um .

In st u dyi n g thi s q u estio n several methods were u sed In .

j e c t i o n of the l u n gs of the d u ck pigeo n a n d sparrow w ith

, ,
3 48 CH I K A N O S U K E OGAWA

to 1 per ce n t silver n itrate was u n s u ccessful S u ch reage n ts as .

ammo n ia silver protei n silver protargol gelatin silver an d

, , , ,

osmiu m silver were u sed O n ly the osmi u m silver showed some

.

i n tercell ular ceme n t li n es alo n g the blood capillaries of respira -

tory can alic uli I fo u n d that solu tio n s of silver n itrate above
.

3 per ce n t also gave resu lts of this ki n d If the respiratory .

epitheliu m were impreg n ated by this the cemen t li n es which , ,

are see n o n the capillaries mu st be co n ti nu o u s with those of the

,

epithelial cells of the flu te holes ; b u t o n the co ntrary I fou n d

‘

-
, ,

that the ceme n t li n es do n o t pass over to the epitheliu m of the

fl u te holes b u t to the e n dotheliu m of the capillaries be n eath
-
,

the epi theliu m of the fl ut e holes Th u s it was bro u ght ou t that

—
.

eve n by i n j ectio n of silver sol utio n i n to the trachea itself the

e n dothelium of the capillaries are impreg n ated In order to .

make my resu lts more certai n I i n j ected the silver sol utio n i n,

the blood vessel system an d fo u n d that images obtai n ed were

-

j u st like those resulti n g from i n j ectio n i nto the trache a .

O n ly the embryol o gical evide n ce remai n s to be disc u ssed i n

relatio n to this problem J u illet claims to have s u cceeded i n
.

impreg n ati ng the respiratory epitheliu m of adva n ced chick

embryos with silver sol u tio n s He n ce I also st u died the embryos
.

of the chicke n du ck a n d goose at several stages of i n c u batio n

, ,

by mea n s of silver impreg n atio n as well as ordi n ary stai n i n g ,

a nd was co n vi n ced that tho u gh the fl u te holes are li n ed by o n e

,
—

layer of c u boidal or flat cell s n o epitheliu m is Visible i n the respira

,

tory ca n alic u li themselves ; at least impreg n atio n of the respira ,

tory epithelium n ever takes place I am i n cli n ed to thi n k that .

J u ill et mistook the epitheliu m of fl u te holes for respiratory -

epitheli u m .

With this the prese n ce of respiratory epitheliu m i n birds

,

became exceedi n gly do u btfu l The fact that n o a n alog ou s case

.

has bee n fo u n d i n which the epithelial coati ng of a free su rface

is lacki n g has led O ppel an d other au thorities to co n cl u de that
the l un g of the bird m u st have res pi ratory epi theliu m i n spite
of the abse n ce of act u al c o n firm at i o n If it were o nly the u s u al .

silver method which failed here s uch a n i n fere n ce might be ,

warra nted b ut it is d i ffi cul t to comprehe n d why n o respiratory

,
 C H IK A N O S U K E O G A \ VA

In the l un gs of these two mammals j u st as i n birds n o impreg , ,

n atio n of ceme n t li n e occ u rs with the ordi n ary silver method ,

b ut it c an be bro u ght abo u t by u si n g co n ce n trated sol utio n s ,

or osmi um silver I exami n ed these ceme nt li n es i n vario u s

—
.

ways an d it became clear that the li n es belo n g to the e n dotheli u m

of the blood capillaries for the followi n g reaso n s :
—

1 Followi n g the blood capillaries to the blood vessels I w as

.
- —
,

able to show that the ceme n t li n es o n the blood capillaries are -

sometimes co n ti n u o u s with those of the blood vessels -

.

2 A s already me n tio n ed the i n tercapillary spaces are pores

.
, ,

b u t i n the places where they come i n to co n tact with other tiss u e ,

s u ch as ple u ra they cann ot form any pores If the ceme n t

,
.

li n es which appear on the blood capillaries by the silver method

,
-
,

belo n g to the respiratory epitheli um they o u ght to co n ti n u e to ,

the spaces at this place In order to decide this qu estio n I cut

.

the silver preparatio n s tan ge n tially to the s u rface of the ple ura
an d fo u n d that u n der the microscope some of the i n tercapillary

spaces i n the bases of the alveoli which adj oi n ed the ple ura , ,

co n tai n ed o ne or two ro un d nu cleated cells while i n other , ,

similar spaces there was n o coati n g of cells at all This c an .

also be see n i n stai n ed sectio n Eve n with the most careful .

exami n atio n I co u ld n o t follow the ceme n t li n es of the blood

capillaries to the spaces .

From this it mu st be co n cl uded that by i n j ectio n of the air

passages with silver sol u tio n capillary e n dothelial cells are the
,

o n ly o n es which are impreg n ated except for the cells at the bases ,

of the alveoli adj ace n t to ple u ra which are also impreg n ated ,
.

The embryos collected for the st u dy of developme n t of respira

tory epitheliu m i n the mole were of too earlier stages b ut i n the ,

yo u n g mole I perceived freq u e n tly ro un d n u cleated cells i n the

i n tercapillary spaces b ut n o trace of epithelial cells o n the
,

blood capillaries
—
.

F r om these s tu di es it s ee ms most p roba ble to con c lu de tha t i n the

mole an d ba t, as m the bi r d, there i s n o c oa ti n g o f resp i ra tory ep i

theli um .

B R esp i r a tory ep i theli u m i n

. It is ge n erally a du l t mamma ls .

believed at prese nt that the respiratory epitheli u m of ad ult

 RE SPI R AT O R Y SP AC E S OF THE L U N G S 3 51

mam m als co n sists of flat n o n n u cleated cells an d s m all nu cleated

,
-

cells A s the respiratory epitheli um is esse n tially the sam e i n

.

the mammals which have bee n st udied the followi n g descriptio n ‘

will apply to them all ( fi g .

T he small nu cleated cells prese nt i n s urface View a ro un d or

,

elo n gated form from 2 u to 1 5 1 i n breadth an d averagi n g 5 u i n

,
1

height It has bee n described som etimes as flat so m etimes as

.
,

c u boi d al A s a matter of fact it is i n termediate betwee n the t w o

.
,

types an d might properly be called either flat or c u boidal b u t ,

i n order to avoid co n f u si n g it with the n o n nu cleated plate it —

,

wil l be co n ven ie n t to desig n ate the n u cleated type as c uboidal .

These cells are distrib u ted n ot always un iformly an d the size

of the alveoli is also variable so that the n u mber of nu cleated ,

cells i n differe nt alveoli differs very m u ch amo un ti n g from o n e ,

to t en or more It is obvi ous that i n the g u i n ea pig rat and

.
—
, ,

others which have small alveoli the n u cleated cells i n an alveol u s

, ,

are fewer Tho u gh these cells u s u ally appear scattered they

.
,

sometimes form gro ups of two or fi v e cells A ltho u gh they border .

the n on nu cleated cells sometimes each cell appears si n gly i n

-
,

the m idst of the n o n n u cleated cells -

.

The large flat n on nu cleated cells are approximately pe n ta

, ,
-

go n al or hexago n al i n shape tho ugh their o u tli n es are n ot straight

,
.

Their diameters are u su ally at least t en times as lo n g as those

of the small cells Cells of i n termediate size betwee n the t w o
.

are almost n ever see n The n o n n u cleated cells i n o n e alveol u s

.
—

n u m ber from three to t en or more A s there is n o co n siderable .

differe n ce i n the size of the epithelial cells of differe n t an imals ,

they are fewer i n the an im als with s m aller alveoli .

A ccordi n g to K é ll i k er the edge of the alveol u s is al m ost en

,

t i rel y covered with the n o n n u cleated an d o n ly seldom with the —

n u cleated cells I agree with this statemen t The n on n u cleated

. .
-

cells are be n t over the edges of the alveolar septa so that they ,

cover two sides of the wall at the same time .

For st u dyi n g the relatio n betwee n the capillaries of the l un g

an d the respiratory epitheli um it is best to select either the ,

preparatio n s i n which the capillaries as well as the epithelium

are impreg n ated or the o n es which are mo un ted i n glyceri n .

TH E AM E R I CAN J O U RNA L OF ANA O M Y V OL

T , . 27 , NO 3
 C H IK A N O S U K E O G A VV A

Fi g . 12 R pi
es rat o r y p i t h l i um C t S i l v i m p g
e e . a . er re i
nat o n . X 3 50 .

Fig . 13 R pi
es ra t o r y p i t h l i um R bb i t m b
e e .
(11
a e ry o S i lv i mp g
er re i
n at o n ,

i i g X 350
st a n n . .

F ig 1 4 R p i t y p i t h l i um R bb i t m b y ( 1 1
. es ra o r e e . a e r o H e m at o xy li n -e os ni . X 600
f m t h i nj t d m t i l b t f m t h m t i l
ro e ec e a er a ,k d in fi
u ro e a er a so a e x i ng i
so lut o n ) .

F i g 1 5 R p i t y p t h l um R b b i t m b y f f u l l t
. es ra or e 1 e l . a e r o o er m . H e m :1 t o x y l i n —
c os i li . X
354 CH IK A N O S U K E OGAWA

fi rst also appeared u n satisfactory becau se the large flat cells , ,

are exceedi n gly thi n an d it is i m possible to decide whether the

n u clei which are stai n ed belo n g to the s ubepithelial cells or
, ,

to the epithelial cells themselves But after f urther attempts .

I happe n ed to obtai n a preparatio n which showed a large flat ,

cell co n tai n i n g n o n u cle u s withi n its limiti n g ceme n t li n es owi n g

to lack of s u bepithelial cells here This image proves that the .

large flat c ell h as n o n u cle u s I applied the third method at

,
.
.

the mo uth of the alveolar d u cts Here the several ki n ds of fi b er .

b un dles run circ ular and accordi n gly the s u bepithelial nu clei are
poor so that I affirm ed several times the fact that the epithelial
,

cells are here n on nu cleated -

.

L an ge has i nvestigated the respiratory epithelial cells i n an

experime n tal way b ut it seems to me that his work is ope n
,

to criticism from several poin ts of view H e bled rabbits ex .

,

tracted the l un gs i nj ected them with isoto n ic salt sol utio n an d

, ,

collected the liqu id r un n i n g off from the tracheal st ump an d that

obtai n ed by slitti n g the s u rface of the l un g H e the n ce ntri .

f u g ed the liq u id a n d st u died the resid u e microscopically The .

followi n g are my arg ume n ts agai n st his work :

1 In the ri n sed liqu id he fo un d alveolar ( respiratory ) epi
.

t h eli al cells e n dothelial cells from blood vessels polym orph e

,
-
,

n u clear le u cocytes a n d red blood corp u scles I sho u ld thin k -

.

that epithelial cells of the bro n chioles an d mo n o n u clear leu co

cytes o u ght to have bee n prese n t also Whe n these cells are .

i n n ormal positio n they may be easily disti n g u ished b u t whe n

, ,

isolated it is impossible to j udge from what part of the l un g some

,

of them came I have i nj ected the trachea with several macera

.

tio n liqu ids c ut the s u rface of the l un g after some ho u rs an d

, ,

exami n ed the escapi n g fl uid b ut I co u ld see o n ly o n e ki n d of

‘

ro un d n u cleated cells N at urally it is impossible to distin gu ish

.

them tho ugh there m u st be di ff ere n t ki n ds

, .

2 The alveolar epithelial cells which were described an d

.
,

fi g u r ed by L a n ge are all nu cleated ,

H e did n o t seem to recog .

n ize the fact that there are two ki n ds of respiratory epithelial

cells It therefore seems that L an ge started his work with some

.

fun dame n tal mistake which re n ders his fu rther experime nts
,

un reliable .
 RE SPI R AT O R Y SP AC E S OF THE L U N G S 3 55

It will perhaps n o t be o u t of place here to disc u ss the methods

u sed by pathologists i n st u dyi n g respiratory epitheli um tho u gh ,

they have already bee n co n sidered by M uller The pathological .

chan ges i n respiratory epitheli um have bee n determi n ed by the

st u dy of thi n sectio n s of the l un g stai n ed i n the ordi n ary way , ,

a n d this method is exceedi n gly i n adeq u ate beca u se it does n ot

reveal the n on n u cleated cells which make up the greater part

—
,

of the coati n g of the respiratory spaces The s m all n u cleated .

cells may at fi r st seem easy to recog n ize i n these preparation s ,

b ut i n fact this is n o t the case becau se owi n g to the thi nn ess , ,

of n on n u cleated cells it is n ecessary to make a caref ul examin a

-
,

tio n before decidi n g whether a n u cle u s which appears withi n ,

the alveolar wall belo n gs to the small n on n u cleated cells or

, ,
-

to the s ubepithelial cells .

C Cri ti ci sm of the op i n i on tha t the r esp i ra tory ep i theli um i s m ade

u p of on e ki n d of cell A ccordi n g to O ppel , the large n on n u cle
.
-

ated cells of the respiratory epitheli u m are n ot separate cells ,

b u t parts of small n u cleated cells the impreg n ated ceme n t lin es ,

bei n g the borders between these portio n s an d the blood capi l —

l ari es. S cym o n o w i c z an d O sawa agree with this statemen t .

My reaso n s for obj ecti n g to this opi n io n are as follows

1 If the n o n n u cleated cell is co n sidered as a part of a nu cle
.
-

ated cell the n i n what way can the border li n es which appear
, ,

with impregn atio n be explai n ed ? In Am phibia the border

,

betwee n the fl at portio n an d the nu cleated portio n un do u btedly

follows the co n to u r of blood capillaries ; b ut this is n ot the case
-

i n mammals beca u se it ca n be eas i ly show n i n silver preparatio n s

, ,

which are mo un ted i n somethi n g like glyceri n that the ceme n t ,

li n es of the epithelial cells do n ot correspo n d to the edges of the

blood capillaries O ppel believes that the prese n ce of border
-
.

li n es betwee n the two parts of the cell s may also be explai n ed

i n some other w ay b u t he does n o t himself explai n i t
,
.

2 The large n o n nu cleated cells are u s u ally adj ace n t to the

.
,
-

small nu cleated cells b ut someti m es the former do n ot to u ch

, ,

the latter an d are e n closed e n tirely by n o n n u cleated cells —

.

( For example n on nu cleated cells i n the middle of fi g ure

,
-

In s u ch cases the n o n n u cleated cells c an hardly be co n sidered

-

as parts of nu cleated cells .

356 CH I K A N O S U K E OGAWA

3 . A ccordi n g
to m y observatio n i n regard to the form atio n
of the respiratory epitheli um n o n n u cleated cells arise from,
-

n u cleated cells by the loss of their n u clei the details of which ,

process will follow i n the n ext sectio n ( D ) .

In short fro m the above arg ume n t it seems that the n o n

, ,

n u cleated cells ca nn ot be co n sidered as parts of the n u cleated

cells .

D . R esp i ra tor y ep i theli u m in em br yon i c an d n ew- bor n ma m

m a ls . No
ag reeme n t has hitherto bee n reached i n the disc u s
sio n as to the form of this tiss u e i n foetal life an d its chan ge
at birth K utt n er makes seemi n gly co ntrary s t ateme nts i n his
.

work o n the o n e han d describi n g the respiratory epitheli um of

,

the embryo as grad u ally fl att eni n g ou t u p to the time of birth ,

a n d o n the other ha n d speaki n g of the c u boidal cells of the

embryo n ic alveol u s as chan gi n g to the flat cells with the fi rst

respiratio n S t ii h r seems to be of this same opi n io n co n cern i n g
.

the s u dde n cha n ge of the fl at cell at the first breath Stieda .

showed that i n the l un g of a 2 50 mm sheep embryo an d of a -

.

large ox embryo the alveolar d u cts are coated with a simple flat
epitheli um Kti lli k er also fi n d s a homoge n eo u s li n i n g of flat
.

epitheli u m i n the alveoli of mat u re e m bryos an d accordi n g to ,

him the respiratory epitheli um is formed i n s u ch a w ay that the

epithelial cells which cover the blood capillaries are s u bj ect to -

great press ure by the mechan ical expan sio n of the alveol u s at
the first breath an d co n seq u e ntly are stretched co n siderably .

Sch ulze describes the alveolar epitheliu m of fully developed

embryos as flat an d he claims that the formatio n of n o n n u cleated
,
-

cells is du e to the expan sio n of the blood capillaries of the alveol u s -

a n d that the f u sio n of adj ace n t cells also participates i n this

formatio n Eb n er believes that the large flat cells arise from

.
,

the f usio n of embryo n ic flat cells A ccordi n g to Croix the .

,

alveolar epitheli u m of the h u ma n embryo before respiratio n has

take n place is all flat an d two ki n ds of epitheliu m are formed i n
the same way that Sch ulze describes o n ly witho ut the process ,

of cell fu sio n Moreover Croix asserts the loss of the n u cle u s

-
.
,

from the large cells is ascribed to their fl at t en i ng All above .

me n tio n ed au thors co n sider that the respiratory epitheli um

3 58 C H IK A N O S U K E OGAWA

theli um others are coated by epitheli um which is di ffer

an d

en t i at ed i n to cells of two di ffere n t shapes ( fi g A lso i n .

s urface View the n u clei of the epithelial cells are see n separated
from each other an d between them the red blood corp u scles are —

ofte n appare n t implyi n g the prese n ce of blood vessels i n these

,
-

S paces Whe n looki n g at the sectio n ed s u rface of the alveolar

.

wall it is ofte n impossible to disti n gu ish any epithelial cells on

,

the blood capillaries which shows that very thi n portio n s of

-
,

epitheli um exi st here .

I also removed the embryos from the mother a n imals j u st

before birth an d treated the l un gs i n the same w ay I fo u n d .

here also that the epithelial cells were di ffere n tiated i nto two
ki n ds The flat cells are partly n on n u cleated an d partly nu cle
.
—

ated Whether or n o t a cell co n tai n s a nu cle u s requ ires very

.

caref u l co n sideratio n tho u gh it is less diffi cu l t here becau se the

, ,

alveolar walls of embryo n ic l un gs are somewhat thicker than

those of the ad ult an imals Cells i n which n o nu clei co uld be . .

impreg n ated by silver an d also disti ng u ished by exami n i n g i n

a less refractive mediu m like glyceri n under regu latio n of light
were co n sidered to have n o nu clei Whe n a nu cle u s was n ot .

impregn ated b ut appeared withi n the border li n e of a cell by

,

light adj u stme nt it w as decided by carefu l foc u sin g whether it

,

belo n ged to an epithelial or s ubepithelial cell But eve n then .

it w as sometim es impossible to decide I co n cl u ded that an .

epithelial cell co n tai n ed a nu cle u s whe n an impregn ated nu cleu s ,

appeared withi n the border li n es of the cell I the n proceeded .

to fi x an d stai n the embryo n ic l u n g of the same litter an d fo un d

the microscopical image di ffere n t from that of an 1 1 cm embryo -
. .

The blood capillaries i n the walls of the alveoli were fill ed by

-

blood corp u scles tho u gh respiratio n had n ot occ u rred an d ao

-
, ,

c o rd i n g ly co uld be traced witho u t a ny i n j ectio n See n i n s u rface .

view ( fig 1 5) an d at the c u t plan e the n u clei rarely appeared

.
,

on the blood capillaries In the i n tercapillary spaces there

-
.

existed c u boidal n u cleated cells which were nu cleated epi

, ,

t h eli al cells T he border li n es of the flat cells had disappeared

.

i n these stai n ed preparatio n s I n ext tried silver impreg n atio n

.

a n d ordi n ary stai n i n g o n the l u n gs of n ew bor n rabbits j u st after -

 RE SPI R AT O R Y SP AC E S OF TH E LUN G S 3 59

the birth The silver preparatio n s of these n ew born a n imals

.
-

appear almost the same as those of adu lt an imals an d n o nu clei

are V isible i n the flat cells (fi g The flat cells were exami n ed
.

i n the w ay me n tio n ed above a n d tho u gh I co u ld fi n d places i n

,

which the abse n ce of n u clei w as perfectly certai n it was tech ,

n i c al l y impossible to show that all of the flat cells i n the fi el d

were n o n n u cleated The stai n ed preparatio n s also prese n ted

-
.

the sam e appearan ce as those of t h e fu ll term embryos ( fig -

.

It is clear from these descriptio n s that the respiratory ep i t h e

liu m of the embryo i n early stages co n sists of a si n gle ki n d of
c u boidal cell an d as developme n t proceeds an d comes n earer to
,

the fin al stage some of them become flatter This di ffere n tia

,
.

tio n does n ot occ u r to the same exte n t i n all alveoli some alveoli ,

bei n g covered by c u boidal an d flat epithelial cells an d others by

c u boidal cells alo n e In the fi n al stage the respiratory epi th e
.

liu m of all the alveoli co n sists of a mixt u re of the two ki n ds of

cells Therefore the prevaili n g opi n io n that respiratory ep i th e
.
,

lial cells are n ot di ffere n tiated i n to two ki n ds u nt il the first respi

ratio n is n ot tru e Bu t it is beyo n d doubt that the fl at cells
.
,

which have differe n tiated before the begi nn i n g of respiratio n ,

become fl at t er i n co n sequ e n ce of respiratio n .

D isappeara n ce of the n u clei of fl at cells takes place al so i n the

fi n al embryo n ic stages an d occ u rs n ot s u dde n ly b u t gradu ally
, , .

It c an be i nferred from preparatio n s i n silver an d ordi n ary stai n s

that the greater part of the nu clei probably has disappeared before
part uritio n .

N ow I will describe the process of disappeara n ce of n u clei .

A s can be see n i n fi g ur e 1 4 the nu clei of some of the flat cells are

,

stai n ed more deeply than those of other adj ace n t cells an d are
smaller i n size an d irregu lar i n shape n ot bei n g ro un d Some
,
.

times chromati n s u bsta n ces are assembled at on e side of the n u clei

a n d the free edges of the n u clei are irregu lar M oreover they .
,

are ofte n stai n ed by eosi n These differe n t n u clear co n ditio n s

.
,

which cann ot be co n sidered as n ormal correspo nd to the phe

,

n o m en a of pyk n osis a n d karyorrhexis an d may be i n terpreted

as bei n g i n the process of disappearan ce In the n ew bor n .

—

foet u s w e sometimes fi n d the same s u ggestio n of nu clear di sap

p ear a n c e i n alveolar walls .
 C H IK A N O S U K E O G A IV A

Fig . 16 R pi
es ra t o r y e p i t h e l i um. St i ll b
-
o rn r a bb i t em b ry o S i l v e r i mp re g n at i o n X 3 50
. . .

Fig . 17 R pi
es r a t o r y e p i t he l i u m
. S t i ll b
-
o r n r a bb i t e m b r y o
. H em at o x y l i n eo s i n X 600
-
. .

Fi g . 18 R pi
es r at o r y e p i t h e l i u m . R at . Tw o d ay s a f t e r w a t e r i n j e c t i o n
. S i l v er i m p reg
i
n at o n . X 3 50 .
3 62 C H IK A N O S U K E OGAWA

flatte n ed eve n i n the prese n ce of n u clei This is clearly show n i n .

fi g u r e 1 3 which represe n ts the preparatio n s made from a n 1 1

,

cm rabbit embryo expa n ded by silver sol utio n It c an be see n

.
,
.

i n a view of sectio n ed al veolar walls that the n u clei are fl at t en ed

i n accorda n ce with the fl at t en i n g of the cells .

A ccordi n g to Eb n er an d Sch u lze the fu sio n of the epithelial ,

cells is partly respo n sible for the formatio n of flat cells Bu t .

hitherto n o positive proof of the fu sio n of embryo n ic epithelial

cells has bee n give n an d as fi gu re 1 3 shows the flat cells c an be
‘

, , ,

formed witho u t f u sio n so that it see m s su p erflu ou s to look for

,

the cau se there .

In co n cl u sio n of this topic a brief descriptio n of the blood ,

capillaries of the alveolar walls of embryo n ic l u n gs may be give n .

It is already k n ow n that i n the foetal circ u lati n g system most of

the blood from the right ve n tricle fl ow s i n to the aorta an d o n ly
a small amou n t of it e n ters the l u n g Therefore I imagi n e at .

fi rst that the blood capillaries of the alveolar walls wo u ld be i n a

-

somewhat collapsed state Bu t o n the co n trary the micro .

, ,

scopic exami n atio n shows that the capillaries are fil l ed with

blood corp u scles Therefore it ca nn ot be tr u e that the collapsed
—
.
,

blood capillaries are exte n ded an d produ ce n ew l umi n a at the

- -

time of the fi rst respiratio n b u t they i n crease i n their le n gth ,

a n d caliber o n ly i n accorda n ce with the expa n sio n of the alveoli ,

a n d i n co n seq u e n ce of i n crease of capacity the blood starts to flow

i n to the l un g from the ge n eral circ ulatio n .

E R ep ar a ti on of r esp i r a tory ep i theli um

. There mu st be a .

limit to the duratio n of the respiratory epitheliu m It wo u ld be .

a n i n teresti n g problem to i nvestigate how the dege n erate epi

t h eli al cells especially n o n nu cleated cells are repaired b u t u p

,
—
, ,

to this time there has bee n n o report made alo n g this li n e .

There is a disti n ct di ffere n ce betwee n the small n u cleated ,

cells an d large n o n nu cleated cells i n ordi n ary m icroscopical

,
-

preparatio n s an d n o i n termediate forms are fo un d The proc

,
.

esses of reparatio n or rege n eratio n cann ot be followed i n s uch

material In order to bri n g abou t rege n eratio n i n an ex p eri
.

me ntal way I tried several methods A t first I i n j ected the

, .

vario u s ki nds of macerati n g sol u tio n s s u ch as alcohol chromic , ,

 RE S P IRAT O R Y SP AC E S OF THE L U N G S 3 63

acid an d so on i n to the l un g thro u gh the thoracic wall of the

, ,

rabbit with a hypodermic syri n ge an d after variou s i n tervals ,

of time I killed the an imals and impreg n ated the lun gs Bu t n o .

impreg n atio n of the i n j ected areas was obtai n ed becau se of the

i n fl amm at i o n an d exu datio n which occ u rred there precl u di n g ,

impreg n atio n I n j ectio n of macerati n g fl u ids i n to the trachea

.

was also u n su ccessful I the n tried i n j ecti n g the l un gs of the

.

rats with cc of distilled water thro u gh the trachea an d ex am

.

i n ed the impreg n ated lu n gs after vario u s i n tervals of time In .

some of the preparatio n s areas were discovered i n which n o

exu datio n had occ u rred an d which showed certain variatio n s i n
the respiratory epitheliu m Besides the n ormal n u cleated cells
.

there occ u rred remarkably small an d remarkably large nu cleated

cells the latter bei n g i n some cases as large as the n on nu cleated
, , ,
-

cells (fi g . Bri efl y I discovered n u cleated cells of variou s

,

gradatio n s of size How c an this best be explai n ed ? It seemed

.

possible to i n terpret the small cells as the res u lts of cell divisio n ,

si n ce they co uld n ot very well have become small as resu lts of

irritatio n du e to i n j ectio n of water an d ofte n occ u rred i n pairs,
.

The presen ce of the nu cleated cells which are larger than n ormal
may be i n terpreted either as the res u lts of swelli n g cau sed by
slight i rritatio n or as the result of chan ge which the nu cleated
cells u n dergo i n order to make up for the loss of n on nu cleated -

cells If the swelli n g be supposed to be the explan atio n the n

.
,

the cells wo u ld h ave i n creased n ot o n ly flat b ut i n all dir ectio n s ,

.

I fo u n d however that the n u cleated cells of vario u s sizes seem

, ,

to ex te n d i n on e pla n e o n ly so that I am i n cli n ed to believe

,

this i n crease i n the size of the n u cleated cells to be du e to some

thi n g else tha n mere swellin g from i n j ectio n of water ; an d it
seems qu ite probable that the nu cleated cells are able to cha n ge
i n to large flat cells I wo uld the n s u ggest the followi n g sequ e n ce
, .

of eve n ts A s a co n sequ e n ce of water i n j ectio n of the l u n g a

.
,

slight i nfl amm at i o n takes place which leads to desqu amatio n of

the epithelial cells after which the small n u cleated cells cha n ge
,

i n to large flat o n es i n order to make up for the loss of the epi

,

t h el i al cells especially flat n o n nu cleated o n es a n d at the same

,
—
,
3 64 CH I K A N O S U K E OGAWA

time they m u ltiply by cell divisio n T o my regret I co u ld n o t .

,

c o n fi rm this by mitotic fi g u r es i n the alveolar walls of the l u n gs

which were i n j ected with water a n d stai n ed with ordi n ary dyes .

M US C LE FIBERS
1 . M u sc le fi bers i n A mp hi bi a

D i emyctylu s pyrr hoga s ter M uller dem on strated i n the l u n g

.

of T rito n a m u scle layer i n which the whole l un g w as almost

e n tirely e n closed These mu scle fi b ers are well developed i n the
.

Trito n cristat u s b u t mu ch less so i n the Trito n t aen i at us A c

,
.

cordi n g to his stateme n t the fib ers ru n circ u larly crossi n g some

, ,

what here a n d there an d are located betwee n the capillaries an d

weak co nn ective tissu e which cover the l u n g O ppel observed .

that the mu scle layer of Trito n alpestris co n sists of circ u larly

arra n ged mu scle cells which are ofte n grou ped In D i em yc .

tylu s p yrrh o g ast er the mu scle fib ers are for the most part ci r cu '

lar ; occasio n ally lo n git ud i n al fi b ers are i ntermi n gled w ith them .

The m u scle fi b ers i n D i em yct ylu s do n o t appear i n gro ups which ,

is co n trary to O pp el s statem e n t They have i n terwove n with

’
.

their s u bstan ce elastic an d collage n o u s fi b ers .

R a n a n tgroma cu la ta Co n cer n i n g the l u n g of R an a G aupp has

.
,

already made detailed stateme n t which is as follows : ,

The
smooth m u scle tiss u e which represe n ts esse n tial fou n datio n of
,

the l ung wall an d septa form fi n er an d coarser b un dles which

, ,

make a firm framework co nn ecti n g with each other The

,
.

stro n gest thickest of these m u scle b u n dles lie i n the free thick
, ,

e n ed edges of the chief septa the mu scle b u n dles i n the septa of

,

the seco n dary order are correspo n di n gly thi nn er From these .

pri n cipal b u n dles of the edges of the sep t at hi nn er fib er s origi n ate ,

which g o dow n i n the septa a n d co n n ect with the fi n er m u scle

b u n dles of the o uter l u n g wall A ccordi n g to my fin di n g i n
.

R a n a n i g ro m acul at a I c an disti n g u ish two systems of m u scle

,

fi b ers i n the o u ter l u n g wall O n e forms relatively stro n g b u n

.

dles the other co n sists so m e irreg ularly arran ged fi b ers In the
,
.

septa the mu scle fi b ers appear i n b u n dles an d i n n o i n sta n ce do ,

they seem to appear si ngly The m u scle b un dles i n the edges

.
 CH I K A N O S U K E O G A IV A

2 . M u scle fi bers i n R epti li a

Clemmys j a p ontcn s The arra n geme n t of the m u scle fi b ers

.

in the septu m an d the o u ter l u n g wall is the sam e as i n R an a an d

Megalobatrach u s ; that is to say the m u scle b u n dles of vario u s
,

stre n gths i nterweave with each other an d amo n g these the ,

i n divi du al fi b ers may be see n The free edges of septa come

.

together from three or fo u r directio n s There are two ways i n .

which the mu scle fib ers i nterweave i n those portio n s where three

free edges come together as will be see n i n fi g u r e 24 A ll three
, .

free edges exchan ge their m uscle fi b ers as a b c or o n ly two , , ,

adj ace n t edges excha n ge their fi b ers as a b a c i n which n o —

,
-
,

fi b ers co n n ect 5 a n d c In s u ch a case b a n d 0 form u s u ally a

.

sharper an gle Where fo u r free edges are merged the m u scle

.
,

fi b ers are n o t o n ly excha n ged betwee n adj ace n t b u t also with ,

those free edges directly opposite Si n ce the free edge i n the .

tortoise is mu ch thicker than that of R a n a the mu scle fi b ers ,

are of correspo n di n gly greater stre n gth i n the former tha n i n the
latter The mu scle fi b ers of the free edges of the primary septa
.

are n ot flat s u ch as i n M egalobatrachu s b u t somewhat cir

, ,

c u l ar i n sectio n This is tru e of the l ower orders of septa as

.

well .

Gecko j ap on i cu s The m u scle b u n dles i n the free edges of the

.

septa i n terweave each other j u st i n the same man n er as i n the

tortoise The mu scle b u n dles which start from the free edges
.
,

of septa an d ru n therei n are weakest compared to the a n imals

,

before me n tio n ed O bserved i n s urface view the b u n dles i n

.
,

the o u ter l u n g wall are also weak b u t the fi b ers which are at
,

right an gles to the septa i n the o u ter wall are somewhat stro n ger .

They mu st be co nsidered as i n comp l etely developed septa b e

cau se of their co urse an d becau se they form small promi n en ce
o n the wall .

Ela p he qua drtvtrga ta The m u scle b u n dles i n the edges of the

.

primary septa are fairly well developed Y et the b u ndles i n .

septa themselves are n o t developed to a correspo n di n g exte n t ,

sometimes eve n co n sisti n g of b u t a few fibers From septa of .

the lower orders there are formed septa which ru n parallel to

 RESPIRAT O R Y SPAC E S OF TH E LU N G S 3 67

24
F ig 1 9 t 2 3 D i g m
s . o a ra of m usc le b un d l es in f r ee edg es o f se pt a in t he lun g
of M g l b t
e a oh j p i
a ra c us a o n cu s .

F ig 24 D i g m f m
. a ra o u s cl e b un d l es in f re e e dg es of se pt a i n t he l un g of

C l emm y s .

TH E AM E R I CA N JoU RN x L O F A N ATO \ 1Y, V OL 27 , NO 3

 C H IK A N O S U K E OGAWA

the o uter l ung; w all accordi n gly i n them the m u scle fi b er s take
,

t h e co u rse par a llel to the o u ter wall In the o u ter l u n g wall I

.

n o t iced weak m u scle b u n dles similar to those i n G ecko .

3 . M u scle fi ber s i n A ves

It is already k n ow n that mu scle tissu e is n ot fo u n d i n the

respiratory ca n alic u li bu t they form circ u larly arra n ged m u scle
,

fi b er s i n fl u te holes
— M y i nvestigatio n s h ave n o t revealed a n y
.

fu rther di ffere n ces A ccordi n g to Piso Borme s stateme n t birds

.
-
’
,

capable of lo n g flights have large nu mb ers of mu scle fi b ers aro un d

the fl ute holes bu t this stateme n t seems to me rather u n fo u n ded
-
, .

4 M n sc le fi bers i n M amma li a
.

Whether the m u scle tiss u es are prese n t i n the alveolar system

of mammalia n l u n gs has bee n a mu ch debated qu estio n But n o —
.

c o n c l u s1 o n has yet bee n draw n from a large l i t er at u r e u O ne

reaso n for the lack of agreeme n t i n the reports of variou s observers
is the difficul t y i n disti n gu ishi n g the mu scle tissu e from the other
tiss u e of the lu n g especially from the other co n n ective tiss u e
,
.

Where a n u mber of mu scle fi b ers co n stit u ti n g a b u n dle an d

havi n g a regu lar arran geme n t are prese n t they are disti n g u ished ,

from the other tiss u es b u t i n i n sta n ces where they are distrib u ted
,

si ngly i n other tiss u es it is diffi cult to disti ng u ish them by dif

fere n ce i n stai n i n g The form of n u cle u s of the mu scle cell alo n e
.

may be u sed as a criterio n Hirsch m a nn says that the m u scle

.

c ells c a n be disti n g u ished by the form of the n u cle u s with cert a i n ty .

E berth holds that the form of the n u cle u s of the m u scle cell c an
n o t be u sed as a mark of disti n ctio n I thi n k both stateme n ts
.

g o t o the extreme . If the form of the n u cleu s an d the co n ditio n

o f i ts prese n ce are take n i n to co n sideratio n the n it is n o t i mp o s ,

sible to j u dge the existe n ce of the m u scle cells i n other tissu es .

With the prese n t microscopic tech n iqu e we have n o better w ay

of determi n i n g their prese n ce .

The thick sectio n s are n ecessary for st u dy of the alveolar

'

s ystem b u t o n the other ha n d the u se of them makes it d i ffi c u lt

,

t o discover the m u scle cells A n other reaso n for the disagree

.
3 70 CH IK A N O S U K E OGAWA

2 . M u sc le fibers i n the a l veolar mou ths Ko ll i k er a n d V E b n er

. .

(i n m an ) S u ssd o rf a n d M uller ( i n domestic mammals ) demo n

, ,

s t r a t ed the mu scle fi b ers i n the alveolar mo u ths These res u lts .

were co n trary to Sch u lze s descriptio n I co u ld n o t fi n d any

’
.

m u scle fi b ers aro u n d the alveolar mo uths i n the lu n gs of ( the bat ) ,

mole rat g u i n ea pig rabbit a n d goat while i n the l u n gs of the

, ,
—
, , ,

dog an d m an m u scle fib er s were demo n strated i n some of the

alveolar mou ths an d i n the lu n g of the cat the greater n umber of ,

alveolar mo u t hs showed fairly stro n g fi b ers In the case of the .

cat the m u scle fi b ers which depart from the mu scle ri n gs aro u n d
, ,

the mo u ths of the alveolar du cts ru n i n the wall of the alveolar

,

d u cts i n vario u s directio n s an d participate at the same time i n

formatio n of the mu scle ri n gs aro u n d the alveolar mo u ths There .

are also prese n t circ u lar fibers which are c o n fin ed to the mo u ths
alo n e In the l u n gs of the dog an d m an o n e may recog n ize
.

o n ly o n e or two m u scle fi b ers aro u n d the alveolar mo u ths There .

fore if there are o n ly a few fi b ers aro u n d the mou ths of the peri
,

p h er a l parts of the alveolar d u cts as well as aro u n

,
d the alveolar
mo u ths it is hard to decide whether or n o t s u ch fi b ers form closed
,

ri n gs .

3 M u s c le fi bers i n the a l veolar wa lls

. E xiste n ce of t h e m u s cl e
.

fi b er s i n alveolar walls has bee n c o n fi rm ed by Piso Borme ( i h —

several domestic mammals an d m an ) an d M uller (i n sheep an d

oxe n ) bu t n o t by Kii lli k er an d V Eb n er ( i h m an ) as well as by
,
.

Schu lze an d Carado nn a (i n mammals ) Tho u gh I co u ld demo n .

strate occasio n al m uscle fib ers which origi n ate from the m u scle
ri n gs of the mo u ths of the alveolar d u cts i n alveolar walls of the
l u n gs of the cat dog ( fi g ,
a n d m an. I co u ld fi nd n o s u ch ,

m uscle fi b ers i n the alveolar walls of the l u n gs of the other mam

mals before me n tio n ed .

E LASTI C FIBERS
Tho u gh w e have several detailed reports co n cer n i n g the elastic
fi b ers of the h u ma n l u n g as yet n o o n e seems to have u n dertake n
,

an i n vestigatio n of the whole vertebrate phyl u m for the p u rpose

of compariso n The method u sed w as observatio n of the l u n g

.

wall an d alveolar walls i n s urface View an d i n sectio n They were .

stai n ed mai nly i n resorci n fu chsi n -

.
 RE SPIR AT O R Y SP AC E S or THE LU N G S 37 1

27

Fig . 25 M us c l e fi b er s o f t h e m o ut h o f t h e a l v eo l a r d u ct M o l e H em at o x y l i n e os i n X
. .
-
.

Fig . 26 M u s c l e fib e rs o f t h e m o u t h s o f t h e a l v e ol ar d u c t s a n d a l v e o l i D o g H e m at o x y . .

X
Fig . 27 El as t ci fi b ers . T he l un g of t he b at . R i f i X 350
es o r c n — u c h s n . .

Fi g . 28 El as ti c fib er s . T he l un g of t he ra t . R in fu h i
e so r c -
c X 350
s n . . E ll l p t i c ar

c e n t er i s t h e m o ut h of the al v e ol ar duct .
37 2 C H IK A N O S U K E OGAWA

1 . Ela sti c fi bers i n A mp hi bi a

Di emyctylu s p yrr hogaster The elastic fib er s are fi n e an d form
.

thick n ets i n s u bmu co u s layer of the l u n g wall These fi b er s .

are see n i n two layers : the i n n er which is circ u lar an d rich i n

,

elastic fib ers the o u ter which is i n the mai n lo n gitu di n al an d less

,

rich i n fi b ers In both layers the fi b ers are n o t wavy b ut ru n

.
,

straight formi n g the n ets by ram ific at i on s an d a n astomoses

, .

The thick n ess of the n et is u niform thro u gho u t the l un g except

for a slight i n crease i n stran ds aro un d the blood vessels other -

tha n capillaries A ccordi n g to O ppel the elastic fib ers are more

.
,

stro n gly developed i n the m u scle b u n dles of the lun g of Trito n

alpestris while i n the lu n g of Di em yc t ylu s the mu scle fib er s do
,

n o t prese n t b u n dles an d these fi b er s do n o t co n tai n elastic tiss u e .

O ppel moreover states that i n his Trito n the o u ter layer

, ,

( which he calls s u bsero u s ) is stro n ger than the i nn er while i n ,

D i em yc t ylu s the reverse is the case .

R an a n tgr omacu la ta The elastic fi b ers of this a n imal are

.

see n i n the co nn ective tiss u e of the alveoli as well as the wall of

the lu n g Thu s i n the septa the fib ers are sit u ated betwee n the
.

two capillary n ets Here w e see that differi n g from D i em yc t y

.
,

lu s elastic fibers take two distrib u tio n s o n e forms b u n dles which

, ,

m ix with m u scle fi b ers while the other has n o relatio n to m u scle

,

tiss u e
.

In ge n eral the elastic fi b ers i n R a n a form looser n ets tha n i n

D i em yc t yl u s . Si n ce the m u scle fi b ers i n the free edges of the
septa are well developed the elastic fib ers are also well developed
, ,

especi ally j u st aro u n d the m u scle b u n dles The i n terweavi n g .

n at u re of the fi b ers which accompa n y the m u scle b u n dles is the

same as described i n the chapter deali n g with m u scle fi b ers .

M ega loba trachus j a p oni cus The elastic fi b ers see n i n the
.

co nn ective tissu e i n the o u ter part of the l u n g are ab u n dan t an d

are coarser than i n D i em yct ylu s : A lthou gh n ot as evide n t as
i n D i em y c t yl u s there are two layers the o u ter co n sisti n g of
, ,

lo n gitu di n al fi b ers the i nn er of fi b ers of irregu l ar co u rs e In

, ,

ge n eral the fi b ers di ffer from those i n D i em yc t ylu s by ramifyi n g

a n d a n astomosi n g more freq u e n tly a n d resemble them by bei n g

prese n t i n mu scle fi b ers .

374 C H IK A N O S U K E OGAWA

p a ra ffi n sectio n s w ere u sed The p a r a f fin sectio n s were n ot

.

a ffi x e d b u t treated the same as the froze n sectio n s after the

,

removal of the paraffin The sectio n s were cu t to the same

.

thick n ess me ntio n ed i n the chapter o n m u scle fib ers .

A E las ti c fi ber s of a du l t ma mma ls

. Ta lp a ( m ote) In the . .

m ole there are few m u scle fi b er s these are paralleled by few ,

elastic fi b ers These fib er s form ri n gs w hich s u rro u n d the

.

mo u ths of the alveolar d u cts In the ri n gs i n some places the .

elastic fib ers form a si n gle sheaf while i n others they prese n t a ,

mu ch looser appeara n ce The large m o u ths of the alveolar

.

d u cts are sometimes s u rro u n ded by ri n gs co n sisti n g of several

sheafs In the mo u th of periphery of the alveolar d u cts the
.

i n dividu al fib ers i n the ri n gs are fi n er The fi b er ri ngs se n d off .

fibers toward the alveolar m o u ths an d the alveolar walls In .

the fi rst part of the alveolar du cts the bra n chi n g fi b ers which
ru n alo n g i n di ffere n t directio n s the alveolar d u cts are stro n g a n d

participate i n the formatio n of ri n gs aro u n d the alveolar mo u ths ,

while we have also the fib er ri n gs which c o n fin e the mo u ths alo n e .

In parts other tha n this there are n o bra n chi n g fi b ers to

stre n gthe n the alveolar mou th ri n gs which co n sist of on e or
more fi b ers The fi b er ri n gs aro u n d the alveolar mo u ths i n t u r n
.

se n d ou t several fi n e fib ers i n to the alveolar walls .

In this a n imal the i n tercapillary spaces are alveolar pores ,

th u s the elastic fi b er s ru n alo n g the capillaries In their co urse .

,

the fi b er s do n o t follow a n i n dividu al capillary b u t p u rs u e as ,

straight a path as the capillary area allows which res u lts i n a ,

slightly c u rved li n e The fi b ers goi n g i n to the alveolar walls

.

a n astomose with each oth er .

Vesp er ug o ( ba t) This a n imal resembles the mole i n t hat the

.

fiber ri n gs are prese n t aro u n d the mo u th of d u cts a n d alveoli ,

b u t di ffers i n that the fi b ers are coarser an d u s u ally loosely

u n ited A s i n the m ole there are fi b er ri n gs aro u n d the mo u ths
.
,

of the alveolar d u cts which also give off bran ches r u nn i ng alo n g
alveolar d u cts an d s u pplem en t the ri n gs aro un d the alveolar
mo u ths .

In the bat however this co n ditio n is see n thro u gho u t the

, ,

alveolar du ct system i n stead of bei n g merely i n the fi rst part

-
 RE S P I R AT O R Y SP AC E S OF TH E LU N G S 37 5

of the d u cts as i n the mole A lso i n the bat the alveolar wall
,
.

fi b ers are a little coarser tha n i n the mole ( fi g .

E pi mys (r a t) There are a great ma ny more elastic fi b er s

.

thro u gho u t the rat tha n i n either the mole or bat The fi b er .

ri n gs arou n d the mo u ths of the alveolar d u cts are very marked

a n d co n sist of loose b u n d l es co m posed of coarse a n d fi n e fi b ers .

The fi b er ri n gs aro u n d th e alveolar mo uths also co n sist of ma ny

coarse fi b ers which whe n they cross each other spread an d
, , ,

i n terlace form i n g a co nfu sed n et ( fi g

,
.

This same pict u re w as see n i n the bat altho u gh it was n o t ,

e v ide n t e n o u gh to deserve descriptio n .

Ca vta ( gu i n ea p i g ) N o remarkable di ffere n ce from the rat

.

c a n be c o n fi rm ed .

L ep us ( r a bbi t) The fibers are remarkably rich an d prese n t

.

t h e same arra n geme n t aro u n d the mo u ths of the alveolar d u cts

"

a n d alveoli as i n the precedi n g mammals b u t the fi b er s i n the ,

alveolar walls differ slightly Here w e have ofte n on e or two .

thick fib ers i n additio n to the fi n e fi b er s These thick fib ers .

p u rsu e a co u rse t hro u gh the alveolar wall so that they u n ite with
either the fi b er ri n g where they started or an adj oi n i n g o n e .

D u ri n g this co u rse the fi b ers have differe n t appeara n ces n amely , ,

sometimes it divides itself i n several less thick fi b ers sometimes ,

n o t i n either case it gives o ff fi n er fi b er s to the alveolar walls

,
.

The thick fib ers show n o wavy wi n di n g A t depart u re from t he .

fib er ri n g the thick fi b er is made by the u n io n of ma n y fi n e fi b er s ,

altho u gh seldom i t begin s at the ri n g i n a si n gle fi b er

. .

F elts ( ca t)
. The arran gemen t a n d thick n ess of the fi b ers are
abo u t the same as i n the rabbit The elastic fi b ers i n the alveo.

lar walls ofte n cross on e a n other an d sometimes there is a m u t u al

excha n ge of fi b ers In the above me n tio n ed ma m mals the elas
.
—

tic fi b ers of the blood vessels give some fi b ers to the alveolar
-

walls bu t this process is especially evide n t i n the lu n g of the

,

cat with the resu lt that both coarse an d fin e fib ers bra n ch from
,

the walls of the blood vessels i nto the mou ths of the alveolar
—

du cts as well as the alveoli an d i n to the adj ace n t alveolar walls

, ,
.

Ca p ra ( goa t) .The differe n ce from the cat seems to be that i n

this mammal the elastic fib ers are a little richer .
37 6 CH I K A N O S U K E OGAWA

Ca m s ( dog) The dog di ffers from the above described mam

'

-
.

mals i n that the fib ers form closer n ets .

H omo ( ma n ) . the elastic fi b ers are abo u t as close as

In m a n
i n the dog a n d they are f u ll of coarse fi b ers There is n o pri n cipal .

differe n ce betwee n the arr an g m en t of the elastic fib ers i n m an

a n d i n the cat dog etc We have deta i led descriptio n s by
“
.
, ,

L i n ser M iller a n d O rsos

, ,
L i n ser states as follows : Some
.

times the alveoli are s u rrou n ded by a broad l ayer of elastic fi b ers ,

sometimes o n ly by o n e or several h u e fib er s an d sometimes the

fi b ri ll ary gro u n d s u bsta n ce alo n e exists u n der the epitheliu m
” ,

A ccordi n g to my fi n di n g s there are always thick fi b ers passi n g,

thro u gh the alveolar walls i n additio n to the fi ne fib ers If L i n

“
.

ser s stateme nt that fi b ri llary gro un d su bstan ce alo n e exists

’
,

u n der the epitheli u m mea n s the abse n ce of elastic fi b ers i n

,

the alveolar walls the n ou r res u lts dis agree O r sé s disti n g u ished
,
.

two systems i n the elastic fib ers of the alveolar walls n amely , ,

the respiratory an d the i n tercapillary explai n i n g as follows

“The former system bra n ches from the elastic fiber ri n gs of the
,

alveolar mou ths spreads i n t h e alveolar walls an d co n sists of

, ,

stro n g fib ers The latter starts from the lo n gitu di n al elastic

.

fi b er s of the larger blood vessels especially the arteries an d goes

-
, ,

to the alveolar wall Its co u rse is partly alo n g the ramifyi n g

.

vessels a n d partly i n depe n de n t an d forms fi n e fi b er n ets close ,

to the capillaries Both systems are differe n t i n origi n stru e

.
,

t u re a n d fu n ctio n
,
My work c o n firm s the bran chi n g of fi b ers
.

from the mo u ths of the alveolar du cts an d the blood vessel walls -
,

b ut I ca n n ot c o n firm his stateme n t that the fibers from the

mo u ths of the alveolar d u cts are thick an d that the fibers from
the blood vessels are fi ne Besides it is difficul t to disti n g uish
—
.

two disti n ct systems si n ce greater nu mbers of the alveoli are n ot

,

i n co n tact with larger blood vessels so that the elastic fi b ers -

,

of these alveoli are bra n ches of the fi b er ri n gs of the mo u ths of

the alveolar d u cts an d alveoli .

B Ela sti c fi ber s of embryon i c a n d n ew born mamma ls

. In this —
.

st u dy the rabbit dog an d m an were observed L i n ser believes

, ,
.

that the elastic fib ers i n the lu n g of the h u man embryo thereby ,

s upposi n g the same co n ditio n i n other mammals are i n an ,

378 C H IK A N O S U K E OGAWA

led to c u tti n g p araffin sectio n s 1 0 u thick an d stai n i n g It w as .

impossible becau se of the str u ct u re of the l un g to u se froze n

sectio n s i n Amphibia a n d R eptilia except the s n ake The ,
.

sectio n s i n the mammalia n l un gs requ ired a thick n ess of 40 u

to 50 for observatio n of these fi b ers eve n i n this thick n ess the
u
, ,

res u lts were satisfactory becau se of the i n here n t or n at u ral thi n

n ess of the alveolar walls Froze n sectio n s were u sed i n these .

thick mam m alia n specime n s R u ssak off states that the elastic .

fi b ers are als o stai n ed by this method res u lti n g i n a co n f u se d pic ,

t u re I fo un d that i n my case n o elastic fib ers were stai n ed an d

. ,

i n ge n eral these are easily disti n g u ished by their morphological

characteristics .

1 . R eti cu lar fi bers i n A mphi bi a , R ep ti li a , an d A ves

Di emyctylu s . M an y attempts to get

impreg n atio n su ffi c i en t

were made b ut the res u lts allow o n ly the followi n g obser vatio n s
, .

The s u rface view shows fi n e somewhat parallel retic u lar fibers ,

aro u n d the s u bepithelial blood capillaries The fi b ers are wavy -

.

i n ge n eral b u t sometimes are straight

,
It may be that these .

fi b ers are rather more characteristic of the capillaries tha n the

l un g itself The stroma shows n o retic u lar fib ers at least by
.
,

Bielschowsky s method ’
.

R an a n tgr omacu la ta M ega loba tr achu s j a p oni cu s ,

N o res u lts .

were obtai n ed .

Clemmys j aponi cu s Fi n e r etic ular fi b ers are see n aro u n d the

.

blood capillaries A lso there are extremely fi n e fi b ers which

—
.

form a n astomosi n g ri n gs aro u n d the m u scle fi b ers an d fi b er

b un dles The retic ular fi b ers are n o t Visible i n the collage n o u s
.

gro un d s ubsta n ce .

Gecko j ap oni cus The tech n iq u e proved so e ffective i n this

.

a n imal that a detailed descriptio n is possible Two syste m s of .

retic ular fib ers are see n the o n es aro un d the blood capillaries ,
—

a n d the others i n the s u bepithelial co nn ective tiss u e The fibers .

appear to twi n e themselves closely aro un d the capillaries an d

are marked by an especial fi n e n ess these p u rs u e a circ ular co u rse ,

a n d are n o t wavy ( fi g In the space of 1 0 u there are from

.
 RE SPI R AT O R Y SP AC E S O F TH E LU N G S 37 9

seve n to t en fi b ers It w as co n cl u ded that these fib ers exte n d

.

all the w ay aro un d the capillary altho ugh this co n cl u sio n co uld ,

n o t be absol u tely v eri fi ed beca u se the stroma stai n ed very deeply .

These fi b ers very seldom an astomose with each other These same .

fi b ers are see n also i n the larger capillaries an d appear to form

a co n ti nu o u s syste m i n the l un g The fibers i n the s u bepithelial
.

co nn ective tissu e are somewhat thick an d spread o ut flat ly

(fi g . A ltho u gh a careful search w as made n o fib ers co u ld ,

be see n passin g over capillaries also n o circ u lar fi b ers were see n
,

aro un d the m u scle fibers as i n the tortoise ,

.

Elap he qu a dri vtrga ta A h appeara n ce similar to that i n G ecko

.

w as see n except that the s u bepithelial fib ers are somewhat fi n er

,
.

A ves The domestic fowl an d d u ck were u sed i n this st u dy

. .

There are b u n dles of retic u lar fib ers slightly wavy aro u n d the , ,

flu te holes In the respiratory can alic u li the retic u lar fib ers
—
.

are exceedi n gly fi n e an d twin e themselves aro u n d the capillaries ,

b u t they do n o t e n circle the capillaries as they do i n the gecko

a n d s n ake ( fi g .

2 . R eti cu lar fi bers in M amma li a

A . R eti cu lar fi bers of the a du lt m amma ls The retic u lar fi b ers

.

in the ad u lt m ammals are in ge n eral similar i n arran geme n t to

the elastic fi b ers .

Ta lp a (mole) ,
The retic u lar fi b ers of the
Vesp er ugo ( ba t) .

walls of the bro n chioli co n ti nu e i n to the walls of the alveolar

d u cts where they e n circle the mo uths of the alveoli (fi g .

T he fi b ers also form ri n gs aro u n d the mo uths of the alveolar

d u cts A maj ority of these ri n gs are composed of a si n gle fi b er
.
,

b u t some are made u p of several fi b ers These ri n gs i n t u r n .

se n d fi b ers i n to the walls of the alveolar d u cts where they form

ri n gs aro un d the alveolar mo uths The alveolar mo u ths i n the .

fi rst part of the alveolar d u cts sometimes co n sist of several fib ers ,

b u t f u rther dow n the d u cts the mo u ths have o n ly a si n gle fi b er

i n the ri n g In both the mo u ths of the alveolar d u cts a n d the
.

alveoli the fi b ers of the ri n gs show a reg u lar wavy or spiral wi n d

ing . The fib ers of two adj ace n t ri n gs arou n d alveolar d u ct
38 0 C H IK A N O S U K E OGAWA

mo u ths or larger alveolar mo uths do n o t j oi n b u t the ri n gs of ,

other adj ace n t alveolar mo uths j oi n together The fib er ri n gs .

of the alveolar m o u ths give off fi n e fib ers to the alveolar walls .

These fibers travel alo n g the capillaries ramifyi n g a n astomosi n g , , ,

a n d formi n g loose n ets T he retic ular fi b er s of the blood vessels

.
-

a n d ple u ra give off fi b e rs which a n astomose with the fi b ers i n

the alveolar d u cts an d alveoli .

E p i mys ( ra t) The arra n geme n t of the fi b er s i n the l un g of

.

this an imal is similar to that i n Talpa an d V esp erug o The .

fi b er s however are a little coarser ( fi g

, , A s i n the previo u s
.

a n imals each fi b er ri n g aro u n d the mo uths of the alveolar d u cts

,

a n d alveoli co n sists of o n e fi b er b u t i n this case the fi b er some

,

times splits an d rej oi n s itself Si n ce the fi b er ri n gs of adj ace n t

.

alveolar mo u ths are c on flu en t i n their commo n edge trian g ular ,

areas are sometimes formed betwee n alveolar mo u ths In this .

formatio n the ri n gs se n d o ff fi n er ram i fi c at i on s to each other so

that the ri n gs here a n astomose together The fi b er ri n g s of the .

alveolar mo uths agai n se n d off bran ches i n to the alveolar walls

-
.

The bran ches either leave the rin g perpe n dic u larly or i n an
obliq u e directio n It will be remembered that at the depart u re
.

of elastic fi b ers from the ri n g there were several fib ers which

becam e c o n flu ent i n a commo n fib er trun k b ut the retic u lar ,

fib er ri n gs give o ff i n divid u al fibers These vary i n size ; they

.

are someti m es as thick as the fib er ri n g someti m es bran ch ,

immediately at depart u re an d sometimes they are exceedi n gly

,

fi ne . In the alveolar walls the co u rse of the fi b ers is more ir

reg ular tha n i n the ri n gs a n d the fib er s form close ram i fi cat i o n s
,

a n d m a n y a n astomoses .

Ca vi a ( gu i n ea T he retic u lar fi b ers i n this a n imal appear

to be coarser tha n those i n the rat The ri n gs aro u n d the mo uths
.

of the alveolar d u cts an d alveoli co n sist of several fi b ers o r i n , ,

other words they have m a n y splitti n gs Whe n a fib er bra n ches

,
.

from the ri n g it occasio n ally starts as several fib ers which later

,

form a thick fi b er A s i n the rat the bra n ches goi n g i n to the

.
,

alveolar wall are of vario u s thick n esses b ut here well marked ,

—

thick fi b ers are formed which give off fi n e bra n ches an d which
exte n d to the opposite side of the ri n g or reach i n to the adj ace n t
 RE SPI R AT O R Y SPAC E S on THE LU N G S 83

alveolar walls The fi b ers i n the alveol ar wall do n o t always

.

ru n o n the same side b u t sometimes pass thro u gh the i n ter

,

capillary spaces an d app ean i n the opposite side of the wall .

( In the mole bat a n d rat the fi b ers might also pass thro u gh t h e
, ,

i n tercapillary spaces b u t t h is co uld n ot be c o n fi rm ed beca u s e

,

of the i n su fficie n tly i m preg n ated specime n s ) The fi n est retic u lar .

fib ers i n the alveolar walls are so fi n e th a t the oil i m m ersio n is

n ecessary

L ep u s (r a bbi t) The sit u atio n here is similar to that i n the

.

g u i n ea pig except that the fib er ri n gs are coarser an d the fi b er

—
,

n ets i n th e alveolar n ets are closer ( fi g .

F elts ( ca t) a n d Ca m s ( dog) T he n ets of retic u lar fi b ers ar e

'

i n ge n eral very de n se an d the fi b er ri n gs of the mo u ths of the

alveolar d u cts an d alveoli are exceedin gly thick (fi g The .

arran geme n t of the fib ers is the same as i n the above described -

a n imals The specime n s obtai n ed showed the fi b ers i n relatio n

.

to the capillaries very clearly The retic ular fib ers i n the alveolar .

walls pass over the capillaries su p erfi ci ally b u t may sin k slightly ,

in to the alveolar spaces The thick fib ers either pass over the .

capillaries u n chan ged or form the n several fi n e fib ers ; this will

be descri bed u n der the retic u lar fi b ers i n m an The fi b ers i n .

the i ntercapillary spaces sometimes run very close alo n g t h e

co n to u r of the capillaries i n which case they sometimes form ,

clos ed ri n gs an d someti m es they are slightly apart from t h e

,

c apillary an d p u rs u e a co u rse parallel to them ( fi g This .

co n ditio n is probably the same i n the g u i n ea pig an d rabbit -

,

altho u gh I was u n able to show i t .

H omo (ma n ) There is n o si g n ifi c an t di ffere n ce i n fi b ers b e

.

twee n m an an d the cat or dog ( fi g R u ssak o ff however .

, ,

poi n ts o ut that the retic u lar fib ers of the alveolar walls of the
h u man l un gs seldom pass over the blood capillaries Co n trary —
.

Fi g . 33 R ti
e c ul a r fi b e rs . T he l u n g of t he ra t Bi el s c h o w s ky ’

s i i g
st a n n .

X 1 50 .

Fig . 34 R ti
e c ul a r fib e r s . T h e l un g oi t he ra bb i t . Bi el s c h o w s ky '

i i g
s st a n n .

X 1 50 .

F ig . 35 R ti
e c ul a r fi b er s . T h e l un g of the de g . Bi el s c h o w s ky ’
s i i g
st a n n .

X 1 50 .
38 4 C H IKA N O S U K E O GA V VA

to this I h n d that the fi b ers ofte n pass over the capillaries

, .

Besides the fi b er system which I have me n tio n ed i n the precedi n g

a n imals there is accordi n g to R u ssak o ff a fi n e circ ular fi b er
, , ,

system aro u n d the alveolar capillaries I recog n ize the same cir .

c ul ar fi b ers i n the l u n gs of A mphibia a n d R eptilia b u t I fi n d i n ,

Mam malia an d the h u ma n there is a di ffere nt co n ditio n Here .

,

whe n the somewhat thick fi b ers pass from on e i n tercapillary

space to a n other the fiber divides someti mes i n to fi n e fi b ers which
,

either co n tinu e as s u ch i nto the other i ntercapillary space or are

agai n gathered i nto a si n gle thick fi b er ( fi g In this bra n ch .

i n g the fib ers may pass over o n e side of the capillary o n ly or

,

they may split an d pass over both sides b u t i n all cases their ,

co urse is o n e approximately at right a n gles to the axis of the

capillary In short t h e fib ers of the capillaries are co nn ected
.

with the fib ers i n the i n tercapillary spaces an d th u s belo n g to

the same system Perhaps R u ssak off saw o n ly these fi n e fib ers
.

i n relatio n to the capillaries a n d co n cl u ded that they formed a

separate system I was u n able to detect the fi n e retic ular fi b ers
.

o n the capillaries i n the l u n gs of the mole a n d bat b u t i n the ,

rat these were i n dicat ed altho u gh the tech n iq u e did n ot allow

,

c o n fi rm a t i o n .

B The . r eti cu l ar b
fi s
er f
o the embryon i c mamma ls . R u ssak o ff
observed i n the 3 2 cm n ew bor n that the retic u lar fi b ers arise
-
.
-

from cells i n the alveolar walls p u rsu e a slightly tort u o u s co u rse , ,

a n d a n astomose with each other H e also states that these .

fi b er s are i n co n n ectio n with fi n e fib ers which form close n ets

be n eath the epithelial cells I w as un able to see any speci a l n ets.

o f s u bepithelial fib ers i n di ffere n t stages of the rabbit embryo In .

a dditio n it was see n t hat i n the e m bryo the fibers i n the alveolar

walls are less coarse less tort u o u s an d form looser n ets than i n
, ,

the ad ult ( fi g I n the embryo as i n ad ult an imals the fi b er

.
, ,

rin gs of the alveolar d ucts an d alveo l i are thicker than those of

t h e alveolar walls .

The degree of stai n i n g of the retic ular fi b ers does n ot vary with
a g e as it does i n the case of elastic fi b ers .

C . The r ela ti on between ela sti c fi bers , r eti cu lar fi ber s , an d mu s

c le fi bers i n regar d to p os i ti on . R u ssak o ff describes the relatio n
3 86 C H IKA N O S U K E OGAWA

betwee n these three kin ds of fi b ers m a fi ve to six mo n th child -

a s follows :
“ The retic u lar fi b ers i n the mo uths of the alveolar
d u cts an d alveoli e n close t h e elastic fi b ers spirally a n d hold them
tightly together The elastic fi b ers i n t u r n almost always a c
.

c ompa n y the m u scle fibers Several of the spiral retic u lar fi b ers
.

a ccompa n y the elastic fib er s which exte n d from the elastic ri n gs

of the alveolar d u cts i n to the alveolar walls .

N ow sectio n s from t h e above me n tio n ed a n imals were stai n ed

,
-

fi rst by Bi el sbh o w sk y s method an d the n for o n ly a few mi n u tes

’

w ith resorci n f u chsi n becau se this stai n is deleterio u s to the

-
.

s ilver precipitatio n It was fo un d that altho u gh the retic ular

.

fi b er s always accompa n y the elastic fi b er s at the mo u ths of the

alveolar d u cts an d alveoli there was a relatio n co n trary to that
,

described by R u ssak off The spiral retic u lar fi b ers are i n cl u ded
.

withi n a sheath of elastic fi b ers an d where there are few elastic

fi b ers they e n close o n ly the side of the retic ular fi b ers n earest
the l um e n .

The thicker elastic fi b ers an d the thicker retic ular fi b ers which
bran ch from the fi b er ri n gs i n to the alveolar wall always aecom
pa ny o n e a n other w ith the elastic fi b ers co n stan tly sup erfici al
,

to the retic u lar fi b ers This does n o t hold tr ue for the fi n er

.

bra n ches .

R u ssak o ff states that elastic fi b er s always accompa n y m u scle

fi b ers b ut he does n o t describe them i n detail
,
I i n vestigated .

t h e preparatio n s i n which elastic fi b er s an d nu clei of m u scle

fi b ers were stai n ed a n d fo u n d that the elastic fi b ers are likewise
,

s it u ated s u p e rfi c i al to the m u scle fi b ers .

The re m ai n der of the article is devoted to the relatio n of

r etic u lar an d m u scle fi b ers In Bielschowsky s method the ’
.

retic ular fibers are stai n ed so deeply that the n u clei of the mu scle
fi b ers are co n cealed i n the mo uths of the alveolar d u cts an d
alveoli By v a n G ieso n s p i c rofu ch si n method both the n u clei
.
’

a n d fi b e r ri n gs are see n altho u gh the fi n e retic u lar fi b er s are n o t

,

s tai n ed It is clear by this m ethod that the nu clei of the m u scle

.

cells are u s u ally located more deeply tha n the retic ular fi b ers
a n d sometimes they are sit u ated side by side .
 RE SPI R AT O R Y SP AC E S OF TH E LU N G S 38 7

In the fi rst part of the alveolar d u cts i n the l u n gs of the

h uman dog cat etc the fi b er rin gs are rich i n the three ki n ds
, , ,
.
,

of fi b ers an d they do n ot co n stan tly adhere to the described

relatio n ship In the alveolar walls of mam mals the mu scle
.

fi b ers are u s u ally abse n t while the thicker elastic an d retic u lar
,

fi b ers accompany o n e a n other The m u scle fib ers are eve n

.

abse n t i n the mo uths of the alveolar d u cts an d alveoli i n s u ch

an imals as the mole an d gu in ea pig while the bat has n o mu s eu
-
,

l at ure at all i n the respiratory spaces In short it is see n i n .

,

mammals that the thicker elastic an d retic ular fi b ers always

accompan y o n e an other b u t mu scl e fib ers are n o t n ecessarily
,

presen t .

M E M BRANA PR O PRIA AND A LV EO L A R P O R E S

The elastic an d retic u lar fi b ers are the pri n cipal tissu es i n t he
str u ct ure of the alveolar walls of mammalian l un gs These .

fi b ers are e n closed by a thi n membran e which is called membra n a

propria This membran e is u su ally co n sidered as stru ct ureless
.
,

b u t Sch u lze an d others state that it is slightly striped S u ssdorf .

an d M il l l er believe that membra n a propria ( which they call

alveolar membran e ) is an elastic membran e A ll of the proofs .

of this poin t give n by M uller hold more tru e for the elastic fi b ers
i n the tran sverse sectio n of the alveolar wall ; he may have mis
take n these fi b ers for an elastic membran e I agree that the .

membran a propria is stru ct ureless The stripin g seen by Schu lz e

.

w as probably cau sed by the prese n ce of the fi b ers e n closed i n the

membran e .

It has already bee n men tio n ed that there is almost n o mem

bran e m the intercapilla ry spaces i n the l un gs of the mole and
bat b ut ded u ci n g from the fact that variou s fib ers run alo n g the
,

capillaries there mu st be a membran e to e n close these fi b ers

, ,

altho ugh it is impossible to i n spect it directly becau se of its

exceedin g thin n ess .

The s ubj ect of alveolar pores has bee n on e of mu ch disc u ssio n .

U p to the prese n t it has n ot bee n decided whether or n o t alveolar

pores are prese nt i n the alveolar walls of n ormal mammalian
l un gs The detailed literat u re on this problem may be foun d
.
3 88 CH IK A N O S U K E OGAWA

in Schulze Marchan d M u ller etc so that I will merely classify

, , ,
.
,

it as follows :
1 The so called alveolar pores are artefacts
.
—
.

2 The alveolar pores are n ot existe n t i n yo u n g a n imals

. .

3 The alveolar pores are existe n t both i n yo u n g an imals an d

.

i n ad ults .

In the microscopic i n vestigatio n of alveolar pores it is o n ly

n ecessary to have thick sectio n s i n w hich alveolar walls are

darkly stain ed Thu s the sectio n s st u died were selected from

.

preparatio n s with ordi n ary stai n with Bielschowsky stai n or , ,

with silver impregn atio n .

A s it w a s referred to i n the work of Sch u lze an d Marcha n d ,

we observed that the i n tercapillary spaces i n the alveolar walls

of the l u n gs of the mole an d bat lacked membran es These .

spaces form pores I agree with Marchan d that there are seldom
.

membran es i n t h e i n tercapillary spaces These membran es .

,

whe n fo un d are perforated by several pores aro un d which cells

,

approximate closely It w as me n tio n ed above that the i n ter

.

capil lary spaces i n the alveolar walls i n the l un g of the yo un g

mole were n o t perforated b u t were covered with epithelial cells ;
,

from this I co n cl u de that the frequ e n t alveolar pores i n the

ad u lt mole l un g appear as the an imal grows Materials were .

take n moreover from the rat g u i n ea pig rabbit cat dog an d

, , ,
-
, , , ,

h uman They showed i n the greater part of the specime n s the

.

existe n ce of smooth edged alveolar pores The nu mber an d the

—
.

si z e of the pores varied eve n i n the same a n imal sometimes ,

o n ly several pores co uld be see n i n a sectio n an d sometimes

every alveol u s showed some pores Miller s argume n t that .
’

the alveolar pores are artefacts seems to sta n d o n an erro n eo u s

fo u n datio n A ccordi ng to V E b n er it is di ffi cul t to decide
. . .

whether or n ot the pores are artefacts I am able to perceive i n .

the s u rface View of the alveolar walls that there are breaches
res ulti n g from the treatme n t i n preparatio n These pores can .

be disti n gu ished from the tru e alveolar pores becau se the latter
di ffer markedly i n bei n g smooth edged an d ro u n d or elliptically
-

shaped By the reg ularity of shape it is impossible to deem them

.

artefacts In ad ult a nimals a few isolated cases showed n o

.
390 CH I KA N O S U K E OGAWA

5 . The cleated cells of mammalia n respiratory epi

n o n- n u

theli um ca nn ot be co n sidered as parts of the nu cleated cells ,

tho ugh som e a uthors are i n cli n ed to believe s o .

6 The respiratory epitheli u m of the rabbit embryo i n early

.

stages co n sists of a si n gle ki n d of c uboidal cell an d as develop ,

me n t proceeds an d comes n earer to the fi n al stage some of them ,

become flatter In the fi n al stage the respiratory epitheli u m of

.

all the alveoli co n sists of a mixt u re of th e two ki n ds of cells

witho ut respiratio n The fl at cells become flatter at the begi n
.

n i n g of respiratio n D isappeara n ce of the n u clei of flat cells

.

takes place i n the fin al embryo n ic stages an d occ urs n ot s u dde nly , ,

b u t grad u ally by processes of pyk n osis karyorrhexis etc The , , .

greater part of the n u clei probably has disappeared befor e

part uritio n .

7 R eparatio n an d rege n eratio n of n o n n u cleated fl at cells

.
-

seem to be bro u ght abo u t by exte n sio n of small n u cleated cell s

a nd disappeara n ce of n u clei .

8 The m u scle fi b ers of the l un g appear isolated i n D i emyc t yl us

.

j apo nic u s an d gro uped i n Megalobatrachu s j apo ni c u s whil e i n ,

R a n a ni g rom ac u lat a a n d Clemm ys j apo ni c u s there are both ki n ds

of appeara n ce The m u scle fi b ers of Elaphe qu adri v i rg at a and
.

G ecko j apo n ic u s form mostly b un dles .

9 The m u scle fi b ers are e n tirely abse n t i n all parts of the

.

alveolar d u cts i n the bat l un g There are existe nt some circ u larly
.

arran ged mu scle fi b ers at the mo u ths of the alveolar d u cts i n

the mole rat gu i n ea pig rabbit goat cat dog an d m an In
, ,
—
, , , , ,
.

the mole and gu i n ea pig the mu scle ri n gs are sometimes abse nt

-

at the mo u ths of the periphery of the d u cts There are n o m u scle .

fi b ers aro u n d the alveolar mo u ths i n the l un gs of the mole rat .

,

rabbit an d goat while i n the l un gs of the do g an d m an m us c le

, ,

fi b ers are demo n strated i n some of the alveol ar mo u ths an d i n

t h e l un g of the cat the greater n u mber of alveolar mo u ths show
fairly stro n g fib ers M u scle fibers i n the alveolar walls are very
.

seldom demo n strated i n t he cat dog an d m an b ut they are , , ,

n ot fo u n d i n the other mammals before me n tio n ed .

1 0 The elastic fi b ers of the mole an d bat form a weak rin g

.

aro un d the m o uths of the alveolar d u cts and alveoli From .

 RESPIRAT O RY SP AC E S OF THE L U N G S 39 1

these fiber ri n gs a few fi b ers are se n t off to the alveolar walls .

In the bat the fi b er ri n gs aro u n d the alveolar mo u ths are s upple

m en t ed also by the fi b ers which bra n ch off from the mo u ths of
the alveolar du cts an d ru n alo n g the alveolar d ucts In the l un g .

of the bat an d g ui n ea pig the elastic fib er rin gs co n sist of loose

—

b un dles of several fibers an d the fib ers i n the alveolar walls b e

come rich The elastic fi b ers i n the rabbit l un g are richer
.

an d the thick mai n fibers appear i n the alveolar walls In the .

goat cat dog an d m an the fi b ers i n crease i n closen ess Fin din gs
, , , .

by L i n ser and O rsos co n cer n i ng the elastic fi b ers i n the h uman

l un g are sometimes co ntrary to min e .

1 1 R etic ular fibers i n the l u n g of G ecko an d E laph e are

.

divided i n to two systems the fi n e circ u lar fi b ers aroun d the blood
,

capillaries an d fl atly spreadin g fib ers i n the s u bepithelial con

hective tiss u e In the respiratory can alic uli of t h e domestic fowl
.

an d d u ck the retic u lar fi b ers twi n e themselves aro un d t h e

capillaries .

1 2 In the l un g of th e mole an d bat a maj ority of the retic u lar

.

fibe r ri n gs aro un d the alveolar d u cts an d alveoli are composed of

a si n gle fiber From the fib er ri n gs of the alveolar mo uths several
.

fi ne fi b ers are give n off to the alveolar walls In the rat and .

g u in ea pig the fi b ers are coarser an d closer ; the fi b er rin gs co n sist

-

of several fi b ers In the gui n ea pig the thick main fi b ers appear
.
-

i n the alveolar walls In the rabbit cat do g an d m an t h e

.
, , ,

fib ers are very close Circ ular fib ers aro un d the blood capillaries
.
-

cann ot be c on firm ed R etic ular fi b ers of the embryo ( ripe

.

rabbit embryo ) are a little fi n er less tort u o u s an d form looser

, ,

n ets.

1 3 The elastic fi b er s of the mo u ths of the alveolar d u cts

.

an d alveoli are always located sup erfi c i al l y to t h e retic u lar fi b ers

a nd m u scle fi b ers .

1 4 Membran a propria of the alveolar walls is n ot elastic

.

membran e .

1 5 The alveolar pores are n ormally fo u n d i n man y mammals

.

an d o n ly seldom ca n n ot be see n .
3 92 CH I K A N O S U K E OGAWA
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A or T 27 , NO 4
E P E M E R 1 920
,

S T B ,
R es um e n por a utor Alexan der S Begg
el , . .

E sc u ela Médica Harvard Bosto n , .

A u se n cia de ve n a cava i n ferior en u h emb rl o n de cerdo de 12 mm

asociada c o n el d re n aj e del sistema de la porta '

en el sistema cardi n al .

L os casos de a usen cia de la ve n a cava i n ferior descritos hasta

el presen te lo han sido en ad ultos 0 en embrio n es avan zados en ,

los c uales esta co ndici on era d uradera mien tras q ue el prese n te ,

caso se prese n ta en u m emb ri é n j ove n ( 1 2 eu un estado c r i

tico de la f ormaci on de los vasos san g u n eos E xisten las dos ’

ve n as cardin ales posteriores y los can ales sub cardi nal es n o son ,

gran des L os sinu soides hepat i co s y las ve n as sub cardi n al es n o

.

se han un id o t o davi a aun c u an do n ormalmen te esta un i on tie n e

'

l ugar en u n periodo mas tempran o El caracter de la c u bierta .

del h i gado es de tal clase qu e parece poco probable qu e la an as

tomosis t en ga l u gar mas tarde El au tor con sidera importan te .

a este p un to porq u e si la un i on n o se lleva a cabo habra a u se n cia

, ,

de la ve n a cava i n ferior aun c u an do las otras porcion es existan .

L a seg un da an omal i a es mas Visible L as con exio n es de las .

ve n as mesent éri cas c on el h i gado son m uy delicadas y un nu evo

can al est a formando se por medio de la ve n a esp léni c a y el sistema
c ardin al posterior El au tor hace refere n cias de ciertos casos
.

importa n tes de ambas an omal i as previame n te descritos eu la

literat u ra .

T ra n sl a t io n by J osé F N o m d ez
C o r nell Me di c a l C ll g
o e e. N ew Y o rk
3 96 AL E XAN DE R s . B E GG

O R TA

A PUL M

V S PL
. .

V M ES S U
. .

R EN S IN. .

Fig . 1 T he he a r t, l i ver ,
a n d a ss o c a t e i dv i e ns o f a p ig s ix w ee k f t bi
s a er rt h .

F m
ro a di i
s s ec t o n of an d p im
i nj R d u d pp x i m t l y
ect e s ec en . e ce a ro a e A .

p u lm , . p lm yu o n ar ar e r t y ; V h mi h m i z yg
. e ( l ft zyg ) v i ; V
az .
,
e a os or e a os e n .
p or t .
,

p o rt a l v i ; V
e n . r en . in l ft
s ,
en l v i ; V pt
re pl ni v in
a e n . s , s e c e .
 A B S E NC E or THE VENA CA V A IN F E R I O R 397

fe i r
r e ?
” Dwight believed that the vei n w hich ex te n ds thro u gh
the diaphragm to empty i n to the lower part of the atrium is the
i nferior cava regardless of the arran gemen t of its abdomi n al
,

trib u tar ies If the vei n as i n certai n ab n ormal c a ses receives

.
, ,

o n ly the bran ches from the liver an d if this vei n is called the ,

commo n hepatic vei n ( followi n g Kaest n er then co n sist ,

e n cy requ ires that i n n ormal c a ses the cava be desc r ibed as emp
tyi n g i n to the commo n hepatic vei n an d n o t i n to the heart .

The work of L ewis however o n rabbits h a s fu lly established the

, ,

i m po rtan ce of the su bcardi n al vei n s an d the esse n tial n atu re of

their a n astomosis with the si nu soids of the liver i n the formatio n
of the i n ferior ve n a cava In f a ct this seems to be the cr u x of
.
,

the whole matter If this an ast omosis t akes pl a ce a cava is

.
,

presen t ; if it does n o t the n a cava is wan ti n g even tho u gh the

, ,

o t her par ts be presen t i n proper sequ en ce from heart to pelvis .

In this sen se o n ly the embryo abo u t to be de scribed is c o n sid

,

er ed a case of abse n ce of the ve n a cava .

The s ubcar di n al vei n s may play a very importa n t p art i n the

produ ctio n of a n om alies as show n by J oh n son
,
while c on
si d er ab l e i m port an ce is att a ched by H u n ti n gto n a n d M c Cl u r e to

the su pracar din al vei n s i n the pr o du ctio n of var io u s a n omalies

i n the cat There seem to b e therefore a nu mber of chan n els
.
, ,

thro u gh which blo o d may dr ai n fr o m the pelvic limbs an d ab

d o m i n al parietes i n order to reach the heart n amely the cardi n al , ,

vei n s the s u bcardi n al a n asto m o sis with hepatic si nu soids an d

, ,

the su pracardi n al vei n s to which mu st be added their derivatives

, ,

the azygos vei n s ( Parker an d Tozier Sabi n ) ,

.

To facilitate the i n terpretatio n of co n ditio n s i n the embryo

the azygos system has been stu died i n a series of you n g pigs after
birth a n d i n older embryos by mean s of i n j ectio n s an d dis
sectio n s Figu re 1 show s on e su ch preparatio n draw n as seen
.

from the left side The ve n a hemiazygos is seen to lie somewhat

.

to the left an d behi n d the aorta throu ghou t the greater part of
its co u rse receivi n g the later al w a ll vei n s from both sides The
,
.

co n n ectio n with the ve n a cava at the lower en d of the hemiazygos

(X i n fi gu r e) is n o t u su al b u t does occ u r A t the u pper end the
,
.

vei n tu rn s sharply fo rw a r d p assi n g to t h e left c f the aorta an d

,

over the root of the left lu n g w her e it t u r n s dow n ward i n close

3 98 AL E XAN D E R s . BE GG

relatio n to the left atri u m A t the atriove n tric u lar groove the
.

vei n t u r n s to the right a n d after receivi n g the cardiac vei n s

’

empties i n to the right atri u m The sple nic vei n a n d the hemi
.

azygos are i n rather close proximity bei n g separated by the cru s

,

of the diaphragm a n d while n o co n n ecti n g vessel was observed

, ,

capillary co n n ectio n wo u ld n ot seem improbable .

Heretofore the a n omalou s abse n ce of the i nferior ve n a cava

has bee n fo u n d either after birth or i n older embryos at a time ,

whe n the ab n ormal co n ditio n has bee n lo n g established The .

specime n here to be described is a pig embryo of 1 2 mm a stage .

,

n o t m u ch later tha n that i n which the a n astomosis betwee n the

s ubcardi n als an d the hepatic si nu soids fi rst occ u rs The ar .

ra n geme n t of vessels so closely resembles the co n ditio n n ormally

f e nn d at an earlier period that o n e qu estio n s at first exami natio n , ,

if i n deed a n ythi n g is amiss b u t a stu dy of other embryos i n the

,

Harvard Collectio n shows that the formatio n of a ve n a cava i n

f eri o r has beg u n i n 6 mm specime n s a n d is well adva n ced i n
-
.

those of mm A s is well k n ow n a large vessel sho u ld be

.
,

prese n t at 1 2 mm The developme nt of the V ariou s orga n s cor

‘

respo n ds with that i n other 1 2 mm embryos an d the prese n ce

—
.
,

of a n associated an d u n qu estio n ed a n omaly i n the portal system

of ve i n s c o n firm s the correct n ess of the recorded measureme n ts .

In additio n to these co n sideratio n s the de n sity of the mese n

,

ch y m a i n vesti n g the liver as well as that withi n the caval mese n

,

t er y leads to the co n cl u sio n that this specime n wo u ld n ever

,

have possessed a ve n a cava i nferior .

The cardi n al system of vei n s with adj ace n t ve n o u s cha nn els

has bee n modeled i n the ab n ormal specime n an d also i n a n ormal
e mbryo of like size for compariso n These models were made i n
.

w ax a n d plated with copper followi n g Walli n s method While ’

.
,

the cardi n al an d the vei n s of the W olfli an bodies of both sides

w ere modeled o n ly the right side is show n i n the drawi n g
,
.

The appeara n ce of the vei ns as show n by the model of the n or

m al embryo is that made familiar by the graphic reco n stru ctio n s
p ublished by L ewis a n d by the cleared i n j ected embryos

exhibited by Professor Sabi n at the Philadelphia meeti n g of the

(

A merica n A ssociatio n of A n atomists i n 1 9 1 3 ( afterward pu b

li sh ed ,
The ve n a cava i nferior is see n to have reached a
400 AL E XAN D E R s . B E GG

predict the perma n e n t form which it wo uld assu me i n this case .

It serves as a commo n starti n g poi nt for several types of the ab

—

se n ce of the ve n a cava i nferior This wo u ld of cou rse be tru e of

.

a n ormal embryo at a still earlier stage a n d as the variatio n is ,

o n e of arrested developme n t the striki n g resemblan ce to a

,

yo u n ger embryo is to be expected .

A co n spic u o u s a n d more u nu su al a n omaly is i n process of de

v el op m en t i n co n n ectio n with the portal sy stem The commo n .

V itelli n e vei n an d the s u perior mese n teric vei n are see n to be

‘

well formed b u t the u s u al large portal vei n is represe n ted o n ly

,

by small capillary co n n ectio n s with t he hepatic si nu soids The .

m ai n drai n age from the portal system appears to be by way of

the sple n ic vei n i n to the s u bcardi n al a n astomosis Si n ce this .

an astomosis n orm ally forms part of the left re n al vei n i n the

ad ult it follows that this co n ditio n wo u ld lead to the produ ctio n

,

of a tr u nk passi n g from the sple n ic vei n to the left re n al A rare .

case of this sort has bee n reported i n the hu ma n adult by Pe n sa ‘

w h o cites a few other cases from the literat u re In order .

to be co m pat i ble with co n ti nu ed developme n t however the por

‘

, ,

tal co n n ectio n with the liver sho u ld be of some size Cases i n .

which the portal vei n empties directly i nto the systemic circ u la
tio n have however bee n reported for example by Aber n athy
, , , ,

( 1 7 93 ) i n a child of t en mo n ths by L awre n ce ( 1 8 1 4,) i n a perso n

o f some years a n d by Hyrtl

,
i n two a n e n cephalic mo n sters .

The an om aly of the portal system is of a radically di ffere n t

n at u re from that of the abse n t ve n a cava ; i n the latter case large ,

cha n n els n ormally prese n t i n yo u n ger embryos have persisted i n

their earlier proportio n s whereas i n the portal an omaly o n the
, ,

co n trary a large cha n n el always fo u n d i n yo u n ger embryos

,

which sho u ld persist has atrophied The place of this cha nn el

,
.

has bee n take n by the e n largeme n t of a n ormal b u t i n si g ni fic an t

c o n n ectio n which however has n o t escaped the atte n tio n of
, , ,

e mbryologists A mo n g others D avis has partic u larly called at

.
,

te ntio n to capillary co n n ectio n s betwee n the sple n ic a n d su b c ar

d i n al vei n s i n pig embryos These n ormally remain mi nu te or
.

disappear b ut occasio n ally may e nlarge an d prod u ce the extra e r

,

di n ar y co n ditio n fo u n d i n this embryo an d i n the rare cases ,

cited i n the adu lt It is si g ni fi c an t i n i n terpreti n g the co n di

,
.
,
 A B S E NC E OF THE V E NA CA V A IN FE RIO R .
‘

40 1

tio n s fo un d i n the pig embryo that the ve n a cava i nferior was ,

said to be abse n t i n both A ber n athy s an d Hyrtl s cases Al ’ ’

.

tho u gh Aber n athy s case w a s descri b ed man y years ago his care ’

f ul acco u n t of it is ill u strated satisfactorily a n d his specime n has

l o n g bee n preserved i n the mu seu m of S t Bartholomew s Hos
’
.

p ital L o n do n
,
It has more rece n tly bee n see n an d described by .

M cWh i nni e A ltho u gh Hyrtl s cases represe n t a slighter ’

d ep art u re from the co n ditio n s fo un d i n the pig embryo i n asmu ch ,

as both cardi n als persisted it is qu ite possible that o n e of them ,

s ho u ld disappear i n later developme n t an d the co n ditio n s fo u n d

b y A ber n athy wo u ld the n be realized M orphologically there .

,

f ore this embryo helps to explai n two importa n t types of ve n ou s

,

a n omaly which have n o t previo u sly bee n see n i n act u al process

o f developme n t .

LI TERATURE C ITED
A BE R N A HY T ,
J 1 7 93 P h il oso ph T . ran s . R y o al S i ty L d
oc e ,
on on pt . 1 , pp .

59— 63 .

D A R R A CH , W . 1 907 A n at . R ec .
,
v ol . 1, p . 30 .

D AV IS, D M . . 19 10 Am . J ou r . A n at .
, v ol . 1 0 , p p 461 — 47 1 . .

D WI GH T, T . 1 90 1 Jour . A n at . and Phy s , v o l 3 5, p 7 . . .

H A LL ,
E S . . 1 890 P r o c ee di n g Z s oo l . Soc . L d o n o n , p t 3 , p 57 7 . . .

H U N T I NG T O N G S A N D MCC L UR E C F , . .
, ,
. . 1 90 7 A n at . R ec .
, v ol 1 , p 33
. . .

H YRT L J 1 8 3 9 M ed J ah rb u c h e r d es
, . O est e rre i c h S t aa t es , . Bd 27 , S . . 3 13—
.

Wi en .

NS O N F B
J OH , . 1 9 1 2— 1 3 J ou r . A n at . an d Phy s , v ol 47 , p 2 35
. . .

KA S N R S
E T E , . 1 9 00 A rc h iv fur A n at . u .Phy A s ,
n a t A b t , S 2 7 1 — 2 80
. . . .

KE ITH ,
A R H UR 1 895 P T ro c . A n at . Soc . Gt B it i . r a n an d I re l an d , v ol . 30 , p . II,
N v mb o e er .

K E R CK R I N G I U S T 1 67 0 i c i l egium A n at om i c i u m A m st el o d am
, . Sp .

V Ko no sr K
. 1 91 0 rc h f En tw
,
m ec h d O rg Bd 2 9 H 1 S 1 50
. A . . .
-
. . .
, .
, .
, . .

L A W RENCE Med C h i ru r g T r an s T 5 p 1 7 4 1814 . .

, .
, . .

L E W I S F T 1 902 a Am J o u r A n a t v o l 1 pp 2 29 244
,
. . . . .
,
.
, .
—
.

1 90 3 b A m J o u r A n at v o l 2 n o 2 pp 2 1 1 22 5 . . .
,
.
,
.
—
.

M CW HI NN I E 1 8 39 40 L o n d o n M e d G a z M a r c h 2 7 v o l 2 N ew S er i e s p 3 1 —
. .
, , .
, ,
. .

M A R T I N 1 862 M o n at s f u r G e b u r t sk u n d e Bd 20 S 1 7 2 .
,
.
, . .

P A RK E R G H A N D T O Z I E R C H 1 8 98 B u l l Mus C om p Z o o l M ar c h
, . .
, ,
. . . . . .
,
.

P E N S A A 1 908 B o ll d el l a S o c M ed C h i ru r g di P av i a Jun e 26
,
. . . . .
, .

P ETS C J Z HE 1 73 6 al l e r s
,
n at . . H ’
A Sl . e ec t , v ol . 6, sec t . 7 6, p 7 8 1 . . H al l e .

P H I S A L I X, C 1 8 9 8 M em r s C o m . . p R d . en u Soc . Bi ol .
,
X se r es , T 5, p i . . 1 52 .

S A BI N ,
F . 1 9 14 a A n at . R ec .
,
vol . 8, no . 2 .

1915 b C on t r ib u t i on s to em b ry o l o g y . C arn eg i e I n st itut i on of W ash

i ng t on , vol . 3, no . 7 .

W A LLI N ,
I E . . 1913 A n at . R ec .
,
vo l 7 , p 2 51 . . .

Z U M S TE I N , J . 1 8 97 A n at . H ft e e, A b t I , H 2 5, Bd 8 , S . . . . 1 65— 1 8 8 .
 40 2 AL E XAN D E R s . EGG
B

Vc a rd a n t
.
.

V: c a r d . eo ma e mt .

V c a r d p ost .

V u m bi l d ex t

V ca ua a n f

V anes .

V t ten .

F ig . 2 M o d el sh o w n i g n o r m al a rra n g e m en t of p i ip l v i
r nc a e ns i n t he r i gh t
ha l f of a p ig em b ry o of 1 2 mm M a g n i fi c a t i o n 3 3 33 V
.
, . . c a r d co . m d ex t
. .
, V . c ar d .

co m . si n .
,
r i ght an d l ef t c a r di n a l v e i n ( d u c t o f C u v i er ) ; S in . ven .
,
i
s nus v en o u s ;
V . c a rd . an t .
, V . ca rd .
p os t ,
i
a n t e r o r an d p i
o st er o r c ar di n al v i e ns ; V . c a va . i nf .

i f i
n er or v en a c a v a; V . hep . co m .
, c o mm o n he p ti v i
a c e n ; V . l i en .
,
s pl i v i
en c e n ;
 R es ume n porau tor Hayato A rai el , .

I n stit u to Wistar de An atomia y Biologia .

Sobre el desarrollo post n atal del ovario ( rata albi n a ) c on especial

m en c i on del n umero de ov u l e s .

El peso d e los ovarios a ume n ta co n ti n u ame n te c o n la edad ,

pero ge n era me n te el ovario derecho es mas ligero y pesa el 90

l

p e r cie n to d e s peso del izq u ierdo A l n acer ambos ovarios

. c o n ,

tie n e n un os ov ul e s c u yo n umero dismi n u ye a medida q u e

,

a ume n ta l a edad A los vein te y tres dias hay é v ul os y ,

este n umero se ma n tie n e co n stan te casi hasta la p ubertad sig u i ,

e n do un a di sm i n u c i é n a ov ul e s a los sete n ta dias D esp u é s .

hay un a di sm i n u ci en bastan te regu lar hasta q u e el n umero d e

é v ul os e s de a los trei n ta y u n m eses En ge n eral la dis .

m i nu c i é n del n umero de é v ul os se debe a la d eg en eraci é n de l os

é v ul o s primitivos pero los ov u l e s d efin i t i v o s dege n eran tambié n
,
.

L a n u eva form aci é n de é v ul o s a expe n sas del epitelio germin ativo

co n ti n ua desp u és del n acimie n to form an d o se de este modo los ,

é v ul os d efi n i t i v o s D u ran te la p u bertad esta n u eva f orm ac i on se

.

hace me n os activa pero p u ede co n ti nu ar u n ano desp u és del

,

n acimie n to L a o v ul ac i é n p u ede ten er l u gar espo n t an eame n t e

.

( si n la i n fl uen c i a del sexo m acho ) L os c u erpos amari llos se en

.

c u en t ran primerame n te en las ratas q u e mide n de 1 4 8 a 1 50 mm .

de lo n git u d y aparece n en ambos ovarios L os q u e se forman en .

ratas q ue h an sido fec un dadas son un poco mas gran des q u e los
q u e aparece n en ratas si n fec u n dar Hemos e n co n trado eh ambos .

ovarios hasta v ei nt i um f ol i cul o s p rox i m o s a mad u rar .

T r a n sl a t i by J é F
on os N o m d ez
C Y
.

o r n ell Me di l C l l g
ca o e e, N ew o rk
 ’
AUTHO R S A RAC
BS T H I P AP ER I U E D
T or T S SS

BY TH E I L I O GR A P H I C E R V I C E
B B S 12 , J UL Y

TH E P O ST N A T A L DEVEL O P M EN T O F TH E O VA R Y
( A L BI N O R A T ) WITH E SPE CI A L R E F E R E N C E
,

T O TH E NUM B E R O F O V A
HA Y AT O ARAI
The W i s tar I n s ti tu te o f A n a tomy

F O UR CH A R TS

I NTR O DU C TI O N
N u mero u s data on the post n atal developme n t of the ovaries
of lower mammals are to be fou n d bu t so far as I am aware , , ,

there are n o i n vestigat i o n s o n the nu mber of ova i n these an i

mal s For the hu man ovaries however there are fi ve observa
.
, ,

tio n s on the nu mber of ova tho u gh the tech n iqu es u sed by several ,

au thors are ope n to some criticism .

Stra n ge to say despite the great emphasis rece n tly placed o n

,

the st u dy of reprod u ctive phe n ome n a especially i n relatio n to ,

the qu estio n s of i n herita n ce sex differe n ce or of physiology of , ,

the i n ter n al secretio n s n o o n e has attempted to obtai n fun da

,

me n tal data o n the nu mber of ova m the ovaries altho u gh s u ch ,

data may throw some d efi ni t e light o n the vario u s problems i n

the physiology of reprod u ctio n .

I have take n this st u dy of the total nu mber of ova i n the rat

du ri n g the e n tire spa n of life at the s u ggestio n of Prof H H . . .

D o n aldso n to whom I am happy to ack n owledge my i n debted

,

n ess n o t o n ly for his deep i n terest show n d u ri n g the progress of

the work b u t also for n u mero u s s u ggestio n s while prepari n g

,

this paper .

I shall fi rst prese n t the data o n the n u mber of ova i n hu ma n

ovaries as reported by the fi v e i n vestigators .

He n le estimated the nu m ber of the follicles i n an ovary

of an eightee n year old woma n a n d stated that it co ntai n ed
- —
,

abo u t follicles or n o t less tha n ,

i n both ovaries .

The method by which he estimated this n u mber is as follows :

40 5
40 6 HAYATO ARAI
Ich a z hlte
ei e
in v ri ei e
n m S ag i tt al s c h n i t t e a u s d em O a u m n s 18
wel her etw
j ahri g en M ad ch en s , c e h te heil
a d en s c s e i herie
n T d er P r p
f t lcher
um a ss , 2 0 so ze eri herie
Bl asch en ; l aln g s d er g an n P p
’

d es F ro n t al
h itte w r e e e l o etw
sc n s u d n d r n a s l eri herie ei e
a 1 2 0 , an g s d er P p n s , d em
r h e er
l ang st en D u c m ss v ri r llele
des O a um p a a rch h itt Viellei ht
n Du sc n s c
z eh e ei h Z hl
300 an u n m n s n , u n d son a c w ii rde d i e a ei e
d er Bl asch en i n n m
v ri etw
O a um a bei e i ht viel we i er
in d n n c betr e
n g al s ag n

Th u s He n le co u n ted the n u mber of the follicles co n tai n ed i n

o n e six th of o n e sectio n a n d the n obta i n ed his res u lt by m u lti
—

plyi n g this nu mber of ova by the total nu mber of sectio n s .

Heyse co un ted more exactly the nu mber of follicles i n

the ovary of a wo m a n seve n tee n years old The method which .

he adopted w as as follows :
I ch z ahlte i n je de m d e 4 2 S chnitte di e F o llikel u n d m aass u n term
r

Mi krosk o p di e Fl achen ausdeh nun g j ede s ei n zel n en S chn itte s au s .

D i e S umm e aller F ollikel betrug 1 1 65 di e v o n d er S umm e d er S ch n itte

,

ei ng en omm en e Fl ache um fasste 1 99 6 qmm D a di e Di cke d er ei n .

z el n en S ch n itte m m betr ug so erg i eb t si ch ei n R aumi nh alt v on

. .

1 0 0 cb mm auf wel che n al so di e Z ahl v o n 1 1 65 Fo llikel n k o mm t

. N a ch .

A n g abe n v on G e g enb au r u n d Vi ero rdt n ahm i c h d en I nh alt d es g aze n

'

O v ari um s u 3600 c b mm an d an a ch Wur d en i m g an ze n O v ari um

z .
,

ziem li ch F ollikel u z ahlen sei n

z I n de ssen besteht n o ch ei ne
.

H au ptfehler q u elle . W e n n v on d en durch g ez ahl t en S ch nitte n zwei

u nm ittelb ar auf ei n an d er g eh 6re n so wer d e n di e ei n
,
zel nen durch ,

s c hn i tt en en F ollikel i n je d e m d er bei d e n S chn it t e g ez ahlt wer de n al so ,

doppelt ; Graaf sche F ollikel wii rden so sog ar vielf ach i n R e chnung
g e setzt wer de n . A ber au ch we nn di e ei n zel n en S chn itte d ur ch d a z
Wi schen lieg en d e S ub st an z g etrenn t sin d wer den i m mer dem ei n zeln en
,

S ch nitte ei ne A n z ahl F ollikel zu g ere chn et di e mi t ihrem H au pt t hei l e

,

ihm g ar n i cht ang eho r en . D aher werde n al so all e F ollikel di e n i cht ,

auf ei ne n S ch n itte be s chr an kt si n d i m pli cite do ppelt g ez ahlt S o

,
.

habe i ch daher an 1 0 m ein er S chnitte n o ch fe stg e stellt wieviel von den ,

fi b erh au pt i n ih n e n z ahlb are n F o llikel n m i t ihre m C e ntr u m o der bei ,

d en g 6sseren m i t d er Eizelle i n den S chn itte n l ag e n

r ,
Ich f an d dass .
,

die se pr e c e n t d er G esam m t m en g e betr ug e n V o n d en obe n .

erh altenen Follikel n nahm i ch d aher au ch pr e c e n t u n d

es erg ab si ch al s wirkli che A n z ahl der i n d em n or m ale n O v ari um
,

v orh an de n e n F ollikel ( ab g erun d et )

By the method j u st stated Heyse determi n ed the number of

,

follic l es i n both ovaries to be Heyse also obtai n ed his

n u mber by estimatio n .

This method of Heyse is certai nly more exact that Hen le s ’

,

but it is n ot suffi c i ent ly so becau se accordi n g to Vi ero r d t s table

’
,
408 HAYATO ARA I
N umber f
o o va in on e ( 2) ova r y of ma n at d i fi cr en t ag es ( 0 H a n sema nn )
.

A ge N u mber of O va

mon t hs
1 y ea r 2 m o n t h s

While there is still some do u bt whether the n umbers give n by

V. Ha n sema nn are for o n e or for both ovaries yet they show ,

c l early what the previo u s data s u ggest ; n amely that there are ,

man y more ov a presen t d uri n g the earliest years of life tha n

at p uberty an d that even after p u berty the n u mbers show a
,

si g n i fi c a n t decrease .

To obtai n some acc u rate i n formatio n regardi n g the relatio n

between the n u mber of ova an d the age of the a n imal I have ,

u tilized the ovaries of albi n o rats .

M ATERIAL AND TE C HN I Q UE
The material u sed i n this st u dy was all s u pplied from the rat
colo n y at The Wistar I n stitu te The n u mber of rats ex a m m ed .

for the sta n dard table w a s thirty n i n e from o n e day after - —

birth to 947 days of age an d the material was collected duri n g

—

seve n mo n ths from A pril to N ovember 1 9 1 8

—

,
.

In each i n sta n ce the body weigh t body le n gth the weights of , ,

the ovaries a n d the appeara n ce of corpora lu tea were recorded

,
.

The removal of the ovary especially whe n the rats are very small
, ,

requ ires some practice E ach ovary was qu ickly weighed an d

.

the n fix ed i n Bou i n s sol u tio n for from six to eight ho urs The
’
.

material w as washed i n r u n n i n g water for twe n ty to thirty m i n

u tes sometimes for o n e ho u r r u n thro u gh the alcohols cleared
, , ,

i n xylol a n d fi n ally imbedded i n p araffin

,
E ach e n tire ovary .

was sectio n ed i n series at 1 0 u an d the sectio n s were stai n ed with

,

hematoxyli n an d eosi n .

In order to obtai n approximate l y the tr u e n u mber of ova i n

the ovaries I have co u n ted u n der the microscope the nu cleu s of
,

every ovu m i n the series of sectio n s obtai n ed from each e n tire

 NUM BE R OF O VA : AL BIN O RAT 409

ovary . The n u clei which were co un ted were those most dis
t i n c t l y stai n ed by hematoxyli n The diameter of the n u clei of .

the ova ra n ge from 8 to 1 2 1 both i n the primitive ova an d i n the 1

d efin i t i v e ova By this method we might make a do u ble co un t

.

i n g occasio n ally becau se it is based o n the co u n ti n g of the dis

,

t i n c t ly stai n ed n u clei O n the other ha n d there is n o fear of

.
,

missi n g ov a becau se the sectio n s were 1 0 u thick while the

, ,

smallest nu cle u s has a diameter of 8 u at least .

For coun ti n g the nu mber of ova i n yo un ger ovaries especially ,

those before twe n ty days of age the u se of the n et micrometer ,

-

is n ecessary For the measureme n t of the diameter of the larger

.

ova a Zeiss c omp en s oc u lar n o 6 an d obj ect 4 was u sed an d

. . .

the micrometer eyepiece was so adj u sted that each divisio n

equ alled 4 u.
,
I have divided all the ova accordin g to their ,

diameters i n to fo u r gro u ps :
,

Gr ou p

We fin d ova especially i n the matu re ovaries which show sev

, ,

eral stages of degen eratio n These stages I have divided i n to .

fou r gro u ps .

A The follicles are poorly developed an d have o n e to three

.

layers of cells In the cen tru m the ovu m is n ot fo u n d b u t i n

.
,

stead there is a homoge n eo u s hyalin mass which stai n s red with

eosin .

B The follicles which belo n g to this grou p show merely the

.

o u tli n e of the ova i n follicles b u t the ovu m is witho u t a n u cleu s

,
.

S u ch ov a stai n red with eosin b u t fain tly .

C In this gro u p the follicles are well developed an d some

.
,

times are almost like the mat u re follicles havi n g man y layers of ,

follicle cells b u t these cells are either already dege n erated or

,

are abo u t to dege n erate In these follicles the large cavity u su

.

ally co n tain s fl u id or sometimes a colloidal mass which stai n s a

deep red with eosin However n o ova c an be fo u n d The
.
,
.

shape of the follicles is either circ ular or oval .

TH E AM E R I CAN J O U RNA L OF ANA O M YT , VOL 27 , NO . 4

41 0 HAYATO A R AI

t D In . this gro u p the follicles show an appearan ce similar to

those i n G ro u p C b u t the o u tli n e of the ova shows an irreg u lar
,

co n to u r an d the nu cleu s cann ot be fo u n d Sometimes we fi n d .

n u mero u s cell co n to u rs devoid of n u clei which are stai n ed more

, ,

plai n ly by eo sm than n ormal ova Whether these n u mero u s .

c ell co n to u rs were prod u ced from ma n y ova or from dege n er

ated follicle cells which have falle n 1 n t o the cavity is n o t yet ,

determin ed .

In the ov ary after abo u t sixty to seve n ty days of age corpora

l u tea are pr esen t These we divided i n to two gro ups accordi n g
.

to their characters while each of these may be f u rther su b

,

d ivided i n to two s ubgro u ps .

A L arge corp ora lu tea

. 1 The corp u s l u te u m which belo n gs
. .

to this su bgro u p is large i n size an d its l u tei n cells show n o de

ge n erative processes It co n tai n s some blood i n its ce n tru m
.
,

an d the l u tei n cells are f u ll size .

2 Those belo n gi n g to the s u bgro u p ( 2 ) show the l u tei n cells

.

less fresh an d sig n s of dege n eratio n are prese n t Whether or

,
.

n o t these two s u bgro u ps of large corpora l u tea are prod u ced as

the resu lt of preg n an cy is n o t clear

,
.

B S ma l l corp or a lu tea
. In this seco n d gro u p the corpora
.

l u tea are sm all an d the l u tei n cells i n process of dege n eratio n

'
.

A m o n g them we c an disti n g u ish two forms : B ( 1 ) i n which the

corp u s l u te u m is rich i n blood capillaries a n d its l u tei n cells are
fresh i n appeara n ce while i n
B ( 2 ) the l u tei n cells appear to be resorbed a n d i n their place
the co n n ective tiss u e appears b u t this form c an hardly be dis
,

t i n g u i sh ed from the se called corpora l u tea atresia

‘ ’
-
.

We have co un ted separately the e n tire n u mber of n ormal ova ,

maki n g fo u r gro ups accordi n g to diameter the dege n erate ova ,

an d the corpora l u tea i n each ovary right an d left side an d

— —

fi n all y the total n umber of ova has bee n obtai n ed by addi n g the
n u mbers of ova fo u n d i n both ovaries .
412 HAYATO ARAI

less complete atrophy In passi n g w e may n ote that R iddle s

.
’

observatio n s o n the testes of pigeo n s an d doves show that i n

healthy birds the right testis is larger than the left i n most cases ,

so that i n bir ds asymmetry of the go n ads appears i n both sexes ,

b u t i n opposite se n ses .

Si n ce as w e shall see the weight of the ovaries is i n flu en c ed

, ,

mai n ly by the ab u n dan ce of well developed follicles an d of cor

—

pora lu tea it appears to me probable that the larger ovary

,

sho u ld be bet ter developed i n this respect than the smaller .

The n u mbers of ova fo u n d i n the two ovaries show the fol

lowi n g ratios ( table The nu mbers of ova fo u n d i n the right
ovary were compared with those fo u n d i n the left taki n g the ,

left as the stan dard O ut of the thirty n i n e cases seve n tee n

.
—
,

show the ratios smaller than o n e two show their ratios as

while i n the remain in g twe n ty cases the ratios are larger than
on e .

The averaged ratios i n the two lots ran ge from to

for the right ovary The average of all the thirty n i n e cases
'

—
.

gives the ratio of for the right ovary thu s showi n g that the
,

differen ce i n the n u mber of ova i n the two ovaries is very slight

From the above it is clear that the right ovary co n t ai n s ap
proxim ately the s ame nu mber of ov a as the left or a few more ,

despite the f act that it weighs somewhat less .

The app aren t co n tr a dictio n thu s reve aled m ay be du e to the

presen ce i n the left ovary of a gre a ter nu mber of the l arve o v a ,

as well as of cert ai n st ages of the degen erati n g follicle an d may ,

be i nflu en ced also by a slight i n equ ality of the corpora l u tea o n

the two sides .

If with these su ggestio n s i n min d we co n sult table 1 it appears,

that the combin ed nu mbers for the variou s classes of larger ova
( more tha n 2 0 u i n diameter ) are appro x imately the same i n both
ovaries an d by co n sequ en ce the nu mbers of the smaller sized
,
-

ova are also approximately equ al It has n ot been deemed

.

n ecessary to p u t i n the n u mbers for the ova u n der 2 0 u i n diame,

ter as these may be obtain ed by subtractio n From the fact that

,
.

while the total number of ova con tain ed i n both the right an d
the left ovary is approx imately the same while n evertheless the
,
Gi vi n g wi th ag es the n u m ber of ova i n the l ef t a n d the r i g ht ova r y— tog et her w i th thei r
r es p ec t i ve wei g hts A l s o t he r el a ti ve wei g hts of the r i g ht ova r y a n d the r el a ti ve
.

n u m ber o f ova in i t . The n u mber s f o r the cor p o r a l u tea ar e a l s o g i ven .

LE FT OVARY R I GH OVA RY
T RA IO T S

A GE

mg m

Av e ra g b f
es e o re a pp e a r an c e o f c o r p o ra lutea .

Av er a g ft
es a er a pp e ara n c e o f c o r p o ra l u t ea .

P gre n an t .
414 HAYATO ARAI

right ovary is smaller it seems reaso n able to i n fer that the right
,

ov ary m ay be sligh t ly ret arded a n d co n seq u e n tly is smaller an d

,

also co n tai n s more ova than the left as the n u mber of ova ,

decreases with i n creasi n g age .

This co n cl u sio n seems to be s u pported by the fact tha t previ

o u s to the fi r st appeara n ce of the corpora l u tea at sixty fo u r days -

3 50 0 0

3 00 0 0

2 50 0 0

1 50 0 0

50 0 0

2 00 300 50 0 60 0 700 8 00

C h a rt 1 Sh wi g th t t
o n e o al n u mb er of ov a in b ot h o v i
a r es o f t he al b in o rat

at di ff er en t ag (i d t il)
es n e a .

the average relative weight of the right ovary is while after

this age it is ( bottom of table

O n the n um ber o f ova in r ela ti on to a ge

table 2 the data o n the thirty n i n e rats are give n

In -
.

A s will be see n from both table 2 an d chart 1 the total n u m ,

ber of ova co un ted i n both ovaries o n e day after birth was

This nu mber decreases rapidly to at twe n ty days after ,

which there is a very slow decrease to abo u t at sixty fo u r -

 41 6 HAYATO ARAI

days A t this age the corpora l u tea may appear that is at abo ut
.

sixty fo u r days ovulatio n occ urs From sixty fo u r days o n t h e

-
.
-

total n u mber of ova decreases slowly b u t steadily to abo u t

at 94 7 day s This was the oldest rat available for st u dy U si n g
.
.

the ratio of the span of life 1 to 3 0 which we commo n ly employ , ,

whe n compari n g the rat with m an 947 days for the rat is equ i v a ,

le n t to seve n ty eight years for m an

—
.

3000 0

2 50 0 0

1 50 0 0

50 0 0

50 0 60 0 800 900 10 0 0
Ag e — d a ys

C h a rt 2 Sh wi g o n t he t o t al nu m b er o f o v a , as w e ll as t he num b er o f ov a of

di ff e r en t s iz es i n t he al b ino r at at diff e r en t a g es ( c o n d en se d ) .

Ou acco un t of the high i n dividu al variatio n I have averaged ,

the data i n table 2 an d give n the averages i n table 3 an d chart 2

with the hope that the data th u s arran ged might reveal more
clearly the real te n de n cy to the chan ges i n the n u mber of ova
accordi n g to the age of the rat .

From birth up to 1 1 0 days the n u mber of ova i n the ovaries

of the rats were gro u ped together withi n t en day i n tervals an d -
,

after 1 1 0 days at i n tervals of abo u t fift y days The res ults of .

this gro upi n g are show n i n table 3 a n d chart 2 We n otice that .

from the averages of fo u r to twe n ty three days the mean total —

n u mber decreases very rapidly from to ova an d , ,

 NUMBE R OF O VA : ALBIN O RA T 417

from twen ty three to sixty three days the n u mber of ova with
- -

slight V ariatio n s dimi n ishes from ( twe n ty three days ) to -

( sixty three days )

-
From sixty three days on it de
.
-

creases rapidly at fi rst then rather steadily to 559 days droppi n g

, ,

markedly at the last e n try for 947 days .

T ABL E 3
The between the ag e a n d the n u m ber of ova i n both ova r i es ; the a ver ag e
r el a ti on

n u mbers Thi s i s the f un d amen ta l ta bl e u sed f or the d i s c u s s i on

.

NUM E R B O F O VA

Chart 2 shows two sharp drops the fir st drop exten ds from fo u r

to twen ty three day s an d the seco n d exte n ds from sixty three to
- —

seven ty days with a period of relative co n stan cy between twe n ty

,

three an d sixty three days The si g n i fi c an c e of these phases

-
.

will be taken up later (p .

Si n ce the total n u mber of ova i n cl u des o v a of v arl ou s sizes I ,

have attempted to an alyze the ge n eral graph chart 2 i n to its — —

418 HAYATO ARAI

compo n e n ts i n order to throw some light on the growth of the

several gro ups .

The graph for the n u mber of ova with diameters less than 2 0 u
wo u ld be n early the same as that show n by the graph for the total
n u mber of ova an d has therefore n o t bee n draw n
, .

The form of the graph for the n u mber of ov a with diameters

of 2 0 to 40 u ho wever shows a slight differe n ce The n umber
, , , .

decreases rapidly from fo u r days u n til t we n ty three days an d -

,

then con tinu es to decrease gradu ally till 947 days the flu c tu at i o n s ,

i n n u mber bei n g appare n tly du e to i n dividu al variatio n .

The graph for the number of o v a with diameters of 40 to 60 u

i n creases rapidly from twelve days an d reaches its maximum at
twe n ty three days A fter twen ty three days it decreases agai n
—
.
-

at first rapidly the n more grad u ally

,
.

The graph for the nu mber of ova with diameters more than
60 12 shows a rapid decrease from twe n ty three days to forty -

three days A fter forty three days it shows b ut a very slight

.
-

fall It is to be n oted that as the diameters of the ova i n crease

.
,

the age of their appearan ce advan ces .

We c an obtain more clearly the relative growth of these gro ups

of ov a from table 4 i n which the total nu mber of ova is take n
,

as the stan dard an d the n u mbers of ova of vario us sizes are rep
,

rese n ted as the perce n tages of this total The percen tage valu es .

of the n u mber of ov a with diameters less than 2 0 Mran ge from 1 00

per cen t to 88 per ce n t of the total thro u gho u t the en tire Span of
life Withi n these two limits there are nu merou s flu ct u at i o n s
.
,

the reaso n s for which will be disc u ssed later .

Taki n g the e n tire series from twe n ty days whe n the largest ,

follicles appear to 559 days the approximate age of the me n o

, ,

pau se the average perce n tage val u es for the several classes of
,

o v a are

U n der 2 0 u per ce n t,

2 0 to 4 0 u per ce n t
,

40 to 60 y per ce n t
O ver 60 u per ce n t
1 I m medi ate ly after bir t h both the absolu te n u mber of sm all
.

ova as w ell as their perce n tage val u es decrease rapidly reachi n g

, , ,
42 0 HAYAT O ARAI

a mi n i m u m at abo u t twe n ty days This striki n g phe n ome n a is .

associated with the rapid i n crease of the larger ova d uri n g this
period From twe n ty days u p to abo u t thirty six days the
.
-
,

perce n tage val u e i n creases agai n thou gh the absol u te n umber ,

of the small ova is decreased owi n g to the appearan ce of the ,

larger ova ( table From thirty six to sixty fo u r days the - -

percen tage val u es for the n u mber of small ov a are approximately

co n stan t This is associated with slight nu merical chan ges i n
.

the n u mber of ova of all sizes .

2 Betwee n 64 a n d 1 1 0 days i n those ovaries i n which cor

.
,

pora l u tea have n o t yet appeared the percen tage val u es of the ,

small ova ran ge from per cen t to 9 7 per cen t In the ovaries .

i n which the corpora lu tea have appeared b ut exclu di n g the preg ,

n a n t an imals the n u mber of small ova ran ges from

,
per ce n t
to per ce n t showi n g n o chan ge i n the nu mber of these cells
,

i n relatio n to the appearan ce of the corpora lu tea .

3 From 1 40 to 9 4 7 days the corpora l u tea are always prese n t

.

a n d the perce n tage val u es fl u c t u at e abo u t a mean val u e of 9 4

per ce n t the very oldest rat givi n g 8 9 per ce n t Therefore

,
.
,

ge n erally speaki n g the perce n tage val u es of the small ova ( less
,

than 2 0 u diameters ) remai n co n stan t i n the older rats

,
.

4 In the preg n an t rats fo u r cases w e h n d the perce n tage

.
— -

valu es 95 to 9 7 per ce n t i n three cases an d 8 8 per ce n t i n o n e ,

which is the y ou n gest For this un u su al val u e w e have n o ex

.

plan atio n .

From the foregoi n g it seems that the perce n tage valu es for the
ova u n der 2 0 u decrease rapidly from o n e day after birth till
,

twe n ty days This is followed by an i n crease of 4 per ce n t u p

.

to thirty si x days From thirty six days o n to the en d of the

-
.
-

series the perce n tage valu es remai n n early co n stan t ( the fo u r

preg n an t cases are n o t i n cl u ded i n this gen eral stateme n t ) .

The n u mber of o v a 2 0 to 40 u i n diameter show perce n t age ,

.

valu es which i n crease rapidl y from three days up to seve n days ,

at which age they reach a maximu m of per ce n t This i n i .

tial i n crease is followed by lower val u es up to sixty fo u r days -

,

after which there is a slight te n de n cy to higher perce n tages

with advan c i n g age .
 NUM BER OF O VA : AL BIN O RAT 42 1

The perce n tage valu es for the nu mber of ova with diameters
from 40 to 60 H i n crease rapidly from t en days to twen ty days ,

at which age they reach an absolu te maximu m This i n tu r n is .

followed by a rapid d ecrease t i ll forty o n e days ‘

-
.

From forty on e days o n to 9 47 days the perce n tage valu es r e

-

mai n less than 2 per ce n t ex cept i n three cases Th ere seems to

,
.

b e however a slight te n de n cy for the perce n tage valu es i n this

, ,

gro u p to i n crease i n the older an imals .

The ov a more than 60 u i n diameter o n ly o n ce represe n t more

than 2 per ce n t of the total an d u su ally less than 1 per ce n t so ,

that n o attempt is made to correlate the variatio n s i n their

ab u n dan ce with other chan ges .

In this table are the determin atio n s for fo u r preg n an t rats In .

the ovaries from a rat preg n an t at eighty days the perce n tage
val u es of all ova with diameters more than 2 0 y are co n siderably
higher than i n the ovaries of n on preg n an t rats b ut i n the r e —
,

mai n in g three c ases this pec u liarity does n ot appear Wh ether .

this is a si g n ifi can t differe n ce cann ot at the m ome n t be deter

min ed .

From table 4 it may be seen that i n the ovaries i n which cor

pora lu tea are absen t the mean nu mber of ov a per mgm of .

ovary weight ten ds to run i n versely to the percen tage valu e for
the n u mber of ov a havin g a diameter of 60 u or more an d a ,

similar relatio n holds after the corpora l u tea appear i n th e ovar

ies It therefore follows that i n the former cases the nu mber
.

of largest ova is i n large measu re respo n sible for the greater

weight of ovary while i n the latter cases both the largest ova an d
,

the corpora lu tea may be take n as the respo n sible factors .

It shou ld be stated that i n yo un g ovaries there are also man y

larger ova with diameters 40 to 60 u or more thou gh it is qu estio n ,

able whether these are matu re becau se the layer of follicle cells
is abou t three to fo u r cells thick an d moreover the cavity is n ot
,

yet formed I am rather i n cli n ed to believe that these ova are

.

i n the fi rst stage of dege n eratio n rather than i n a stage of de

v el op m en t A fter thirty days we fin d i n the ovaries man y
.

well developed follicles which n ot o n ly have a cavity b u t man y

-
, ,

layers of follicle cells an d as i n the mature follicles the c u mulu s

,
42 2 HAYAT O ARAI

ovigeru s is w ell developed These follicles co n tai n ripe ova .

,

whose diameter is u su ally 60 to 66 u Sometimes we observe a .

very large ovum with a diameter of 7 6 H or m ore bu t s u ch an ,

ovu m may be i n an early stage of dege n eratio n becau se both the ,

n u cle u s an d cell body are n o t well stai n ed as i n the case i n the ,

n ormally ripe n ed ovu m .

The n umbers of the matu re follicles an d of largest ova i n both

ovaries are give n i n table 5 .

A s is show n i n table 5 the n u mber of the matu re follicles is

,

n o t strictly proportio n al to the n u mber of the largest ova with

T ABL E 5

The n u m ber s of ma tu r e f oll i c l es and o f l a rg es t ova i n bo th ovar i es

NUM E R NUM ER
L A RG E O VA L A RG E O V A
B or B or
A GE A GE
ST ST

1
Sh w p
o s r esen c e o f c o r p o ra l u t ea .

2
St d f p g
an s or re n an c y .

a diameter of 60 u an d more N evertheless if we co n sider the .

,

data from forty six to on e h un dred days the n u mber of largest

—

ova are a little more than twice as n u mero u s as the matu re fol
li c l es while i n the remai n der of the table they are a little less
,

than twice Th u s there is a te n de n cy for the proportio n of

.

largest ova to slightly dimi n ish with age .

We n otice also that i n the rat from sixty fo u r to ei g hty days the -

n u mber of mat u re follicles ra n ges from thirty o n e to thirty n i n e - -

,

an d this large n u mber of follicles may be related to the attai n

me n t of sexu al matu rity si n ce after eighty days the n u mber of

,

follicles te n ds to decrease The mea n nu mber of these matu re .

follicles i n all eightee n cases is twe n ty on e -

.
424 HAYATO ARAI

three days grams If n ow chart 3 is exami n ed we see

, .
, , ,

that betwee n these body weight limits the n u mber of ova shows
-

little chan ge j u st as it did betwee n the correspo n di n g age limits

—

i n chart 2 The fall i n n u mber after the body weight of

.

grams is however less marked i n chart 3 than after sixty three

, ,
—

days i n chart 2 This slight di ffere n ce is du e to the fact that

.

3 50 0 0

2 50 0 0

1 50 0 0

10 0 0 0

50 0 0

Bo d y w e ug h t g ms
—

C h ar t 3 G r ap h s h ow n i g t he i
r el a t o n b et w e en t he b dy w i g h t
o e an d t h e t o t al
n um b er o f o v a in b ot h o v i
a r es , to g e t h er i
w t h t he numb of t h
er e cor p ora lu t ea
an d o f t h e t h r ee g r ou p s of ov a m o re t h an 2011 i n di a m e t er .

some of the older rats were ill n o u rished an d small for their
age This is show n by table 7 i n which the observed body
.
,

weights for give n ages are compared with the expected ages as
recorded by D o n aldso n
A s these fi g ures show the coi n ciden ce between the two series
,

of ages is fair ex cept for the last two body weight gro u ps The -
.

an imals i n these gro ups were light for their age .

 NU MBE R ”
OF O V A : AL BIN O R AT 42 5

4 O h the
. n u mber o f o va a n d the body len g th

I have arran ged the nu mber of ova accordi n g to the body

le n gth of the rats an d the resu lts are give n i n table 8 G raph s
, .

TA B E 6 L
The r el a ti on between the bod y wei g hts and to ta l n u mber o f o va i n bo th ova r i es ; a verag e

n u m ber . F r om thi s ta bl e the p r eg n a n t r a ts wer e o mi tted

NU M E R NU M E R
C O R PO R A L U E A
B OF
B OF O VA
T

N UM
I N E R VA L B ODY
B O DY W E I GH W E I GH
BE R T OF
OF T T
RA TS

R at s i
w t hou t cor p o ra lu t e a

g ra m s gm ms mg m .

R at s i
w th cor p or a l u t ea

9 7 8— 1 0 6 3 1 02 O 3 2 0 1 65 71 52 12 15

w
33 8 2 60 131
u
hr 1 1 3 5 1 2 9 5 1 22 8
—
51 10 14 24
1 42 l 46 9 1 50 83 30 54 29 83
1 55 0 1 61 1 53 6 2 18 94 31 39 20 59
2 3 8 O 65 2 1 28 61 21 90 18 1 08

have bee n made for these data also b u t they show relatio n s which ,

i n all respects are so similar to those obtai n ed whe n the data are
plotte d o n body weight that they are n ow omitted It is to be .

n oted that whe n rats reach a body le n gth of 1 4 8 mm ovu latio n .

occ u rs almost i nvariably as show n i n tables 2 an d 8 an d I shall

, ,

take u p this poi n t i n the disc u ssio n .

TH E AM E R I CAN J OU R N A L OF ANA O M Y V OL
T , . 27 , NO . 4
42 6 HAYATO A R AI
T ABL E 7
D a ta on bod y wei g ht a c co r d i ng to ag e

B O DY WEIGH A A GE RA ( AN D A R D A G E I N D A Y S
C O RR E P O ND I N G B O D Y
T OF B TH E S T

( A VER A G E D I N
TS
A V E R XG E D
I N D A YS ) ( O E R V E D )
TH E OF TS
G rcA M S )
O E RV E D ) W E I G H ( D O NA L D O N 1 5)
S r o
'

BS '
( BS T S ,

T ABL E 8
The r el a ti on between the body l en g th an d the n u mber o f ova i n both ovar i es ; a ver ag e

n um ber s

M ER OVA NU M E R
C O R PO R A L U E A
B OF
NU B OF

IN E R WEIGH
T
NUM
B O DY B
T T
BE R
B O DY L ENG H
. VAL O F OF O TH

L ENG H RIE
OF T O VA
RA TS T S

R at s i
w t hou t cor p ora lu t ea

mm . mm mg m

R at s w thi c or p ora lut ea

1 53 1 56
—
1 55 21 1 111 92 21 10
1 62— 1 68 1 68 2 54 1 21 56 8 , 869 7 15 22
17 1 17 1
—
17 1 45 3 1 93 1 19 30 26 25 51
1 8 2 1 85
—
1 84 54 7 1 79 94 51 42 22 64
1 8 9— 200 1 94 56 6 1 54 74 31 43 20 63
215 1 28 61 21 90 18
 428 HAYAT O A R AI

followed by a period of approximate co n stan cy up to an ovary

weight of mgm the en d of the period precedi n g the ap
.
,

p earan c e of the corpora l u tea This stage marked by a break

.
,

i n the graph passes i n to the stage i n which the corpora l u tea

,

appear an d after the appeara n ce of the corpora l u tea the graph

,

3 50 0 0

2 50 0 0

20 00 0

1 50 0 0

10 0 0 0

50 60 70 80
W e i g h t o f o va rne s m ml lhg ra m s

C h a rt 4 G rap h sh ow i n g t he rel a t i o n b etw een t he w e ig h t o f b o t h o v ar i es an d

t h e t o t a l n u m b e r o f o v a t o g e t h e r w i t h t h o se o f t h e t h r ee g r o u p s m o re t h an 20 1
, 2

i n di am e t er . T h e g r a p h s a r e b r o k en a f t e r p u b e r t y
.

for the total n u mber of ova falls very slowl y to the en d of the
record The several graphs for the nu mber of ova more tha n
.

2 0 M i n diameter are very similar i n form to the correspo n di n g

graphs i n chart 3 an d are s u bj ect to a like i n terpretatio n .

It may be appropriate however to call atte n tio n here to the

, ,

relatio n s of the vario u s gro ups of larger ova In the first place .
,

all the larger ova are derived from small ova M oreover the .
,
 NU M BER OF O VA "AL BIN O R AT 42 9

arran geme n t i n gro ups is for co n ve n ie n ce merely an d they rep ,

rese n t i n reality a co n ti n u o u s series It follows from this that .

the grou ps of smallest diameter sho uld be the fir st to appear an d

T ABL E 10

The r el a ti on between the wei g ht f bo th o va r i es an d the n u m ber f mber

'

o o ova ; a verag e n u

NUM E R NU M E R
C O R PO R A L U E A
B OF
B OF O VA
W EI GH T

I N E RV A L
T

O V A RY OH
T OF

W E I GH
OF B T

RI E
T OVA
S

R at s i
w t ho u t cor p o ra lu t ea

mg m . g r a ms

R at s w thi c or p o ra lu t ea

2 20 1 — 22 2 2 1 2 111 4 1 55 64 46 9 9 18
3 31 1 32 5 1 27 2 2 60 1 35 48 31 23 54
3 36 5 4 1 6 3 8 5
—
1 43 1 241 1 22 73 40 16 56
3 43 8 —
49 2 47 3 1 24 8 1 89 94 47 27 23 50
2 54 8 —
58 1 56 4 1 65 6 1 59 99 25 33 27 60
2 69 4 1 99 0 1 29 85 66 61 16 77
73 5 82 6 78 O
—
1 80 1 1 64 65 19 26 18

O v y w i ght
ar e s w e re t a k en f ro m T h e R a t f o r t he
‘ ’
o b se r v d b dy w i gh t
e o e s .

the other gro ups follow i n time of appearan ce i n the ord er of

, ,

their size .

Table 1 0 an d chart 4 show that this is what occ u rs Fu rther .

,

at the time of fi rst appearan ce an y gro up is represen ted by o n ly

a small n u mber an d the appearan ce of the n ex t grou p coi n cides
with the maximu m n u mber i n the gro up j u st precedi n g The .
430 HAYAT O A R AI

gro u p with the smallest diameter ( 2 0 to 40 u) is most s u bj ect to ,

dege n eratio n for the n u mbers i n it fall most rapidly an d c o n

,

t i nu o u sly
. The gro u p with the largest diameter (more than 60 p )
is most co n stan t i n n u mber This relatio n does n o t n ecessarily
.

mean that these largest ova persist for any lo n g period b u t ,

merely that the balan ce betwee n their formatio n an d dege n era

tio n is rather eve nly mai n tai n ed .

The first phase of the graph for the e n tire nu mber of ova may
be i n terpreted as du e to the fact that duri ng this phase man y
primitive germ cells are growi n g rapidly yet at the same time an ,

excess of cells is u n dergoin g dege n eratio n so that the nu mber ,

falls rapidly .

From 8 up to mgm i n ovary weight the e nlargeme n t of

.

n ewly formed d efi n i t i v e ova is mai n ly respo n sible for the i n crease

i n the weight of the ovaries co n tai n i n g a n early co n stan t n u mber

,

of ov a A ll the groups of ova more than 2 0 M i n diameter b e

.

have i n mu ch the same mann er A fter su ch ova are first recog

.

n i z ed there is a short period i n which they i n crease i n nu mber ,

followed i n each grou p by a more or less pro n ou n ced decrease to

the en d of the series O n the whole the n the total n u mber of
.
, ,

ova decreases accordi n g as the ovaries grow i n weight an d i n the ,

fi r st phase this decrease is very rapid b u t from a weight of ,

mgm it becomes mu ch slower

. .

The nu mber of ov a less than 2 0 u i n diameter is n ot plotted on

this chart b u t as table 1 0 shows it wou ld give a c u rve p r a c t i
, , ,

cally iden tical with that for the total n umber .

The compariso n of my o w n data on the weights of the ovaries

with those give n by D o n aldso n shows the followi n g rela
tio n s accordi n g to age ( table
S o far as the critical periods for the data are co n cer n ed there is ,

g ood agreemen t bewee n the two determi n atio n s i n table 1 1 b u t ,

the last e n try i n dicates that my an imals were somewhat retarded .

The data show that the weights of the ovaries hold an i n verse
relatio n to the nu mber of ova that is the heavier the ovaries
, ,

the fewer are the ov a i n them This in verse relatio n may be

.

du e i n part to the formatio n of the i n terstitial tissu e b u t de ,

pe n ds mainly on several other factors su ch as the corpora lu tea, ,

the number of well developed follicles dege n er ate follicles etc

-
, ,
.
43 2 HAYAT O ARAI

nu mbers of ova te n d to be similar i n both ovaries eve n i n the ,

cases where the total nu mber is markedly l o w ( see 1 98 day rat

i n table 1 as the extreme i n sta n ce ) A ppro x imate equ ality is ,

a lso fo u n d for the n u mbers of ova o n the right an d left sides if

the mean s of the valu es for the low records at 8 0 1 00 1 1 0 1 9 8 , , , ,

a n d 2 62 days are compared These are : right ovary 222 3 an d

.

left ovary 2 357 .

The gr owth f
o ovary i n wei ght a cc or di n g to a ge

The growth of the ovary of the albin o rat after birth has bee n
stu died by J ackso n an d H at ai an d it shows sev

eral phases J ackso n who co n stru cted a chart showi ng the rela
.
,

tive w eight of ovaries o n body weight poi n ted ou t two of these ; ,

the first phase begi nn i n g at birth showi n g an i n creas e to a maxi

m um at a body weight of 1 0 to 1 5 grams followed by a decrease ,

up to a body weight of 60 grams A t 60 grams a seco n d period

.

of acceleratio n which correspo n ds to the adven t of p uberty

, ,

begin s an d du ri n g this phase the ovaries i n crease to a seco n d

,

max imu m at 1 1 0 to 1 2 0 grams i n body weight A fter this the .

relative weight steadily decreases To make a compariso n with .

J ac kso n s res u lts I have prepared table 1 2

’
.

This shows that accordin g to my data the relative weight of

o varies i n creases to the fi r st max imu m at a body weight of 22

to 3 3 grams an d the n decreases gradu ally u n til a body weight

,

o f 95 grams A fter 95 grams the c u rve rises rapidly at 1 0 2 grams

.

i n body weight A bo u t this time corpora l u tea appear i n the

.

ovaries an d the maximu m relative weight is reached at a body

,

weight of 1 42 to 1 61 grams after which it decreases In ge n eral

,
.

the two sets of observatio n s agree .

The disagreeme n t i n the exact periods of the max 1 ma betwee n

J ac k so n s data an d my o w n may be cau sed by the smaller n u m
’

ber of rats ( thirty n in e ) u sed by me The explan atio n as to the

-
.

tr ue si g n ifi can c e of the two maxima is d i fficul t especially the first ,

max imum owi n g to the complexity of the factors which e n ter

,

i n to the growth of ovaries s u ch as the e n largemen t of follicles

, ,

the i n crease of stroma tissu e etc The cause for the seco n d,
.

max imu m may be i n terpreted as follows :

 NUM BER OF O VA : AL BIN O R AT 43 3

A ccordi n g to D o n aldso n the correspo n di n g ages for the

body weight of 1 1 0 to 1 2 0 grams i n the female albi n o rat is abou t
seve n ty five to eighty days an d as ovulatio n has occ u rred at this
—
,

age fresh corpora l u tea are al ready prese n t A ccordi n g to J ack .

s o n ovu latio n that is the age of p u berty takes place at seve n ty

, , ,

T ABLE 12

The r el a ti o n between the body wei g ht an d the wei g ht o f both o va r i es ; a ver ag e n u mber

WEIGH
BO H P E RC E N A G E
T or
nm
WEI GH
T o va s T

NUM E R I N E R VA L IN
AGE O VA R I E
TS
B
RA
OF
BODY WEIGH
T OF
M I LLI GR AM
A CC O R D I N G O BO D Y W E I GH
S OF S ON
TS T

B O D Y WE I GH D
T
OB
T

( D O NA L D O N )
T S ERVE
S

R at s i
w t h ou t c or p o ra lu t e a

g r a ms gr a ms mam .

R at s w thi c or p o ra l u t ea

97 8 —
1 06 3 85
1 1 3 5 1 29 5
—
1 10
l 38 7 . 305
1 55 0— 1 67 1 . 27 1
9 47

F ro m th i s b
t a le t he p g re n an t r a t s w ere om it t de .

days In my rats however the age of seve n ty days is r epre

.
, ,

se n ted by a rapid i n crease i n the weight of the ovaries b u t they ,

have n o t yet attai n ed their maximu m Co n sequ e n tly the sec .

o n d maxim u m of J ackso n or the period of rapid i n crease wo u ld

, ,

be cau sed pri n cipally by the formatio n of a n u mber of corpora

l utea i n the ovaries .
43 4 HAYAT O A R AI

We have see n that i n ge n eral the weight of ovaries i n which the

corpora l u tea are n o t prese n t i n creases with the growth of the
rat either i n body weight or body le n gth With the appear
,
.

an ce of the corpora lu tea the weight of the ovaries becomes

more fl u ctu at i n g owi n g perhaps to i n dividu al variatio n s i n the
,

additio n of the corpora lu tea .

T hu s these flu ct u at i on s j u st n oted may have n o mean i n g ex

cept as showi n g a greater variatio n i n the growth rate of the ovary
at this period E ven when the weights of the ovaries fo u n d by
.

myself an d those give n by D o n aldso n are arran ged accordi n g to

the i n creasi n g body len gth the resu lts show a high degree of sim
,

i l ari t y to those which were arran ged accordi n g to i n creasi n g

body weight ( table M y observatio n s show that with the
appearan ce of corpora lu tea there is a su dden i n crease i n the
weight of the ovaries an d we may i n fer therefore that the i n
, , ,

crease i n the weight of ovaries with i n creasi n g body weight as ,

n oted i n The R at chart 2 1 is also du e to the same cau se

‘ ’
.
, ,

In woman the ovaries are said to atrophy after the me n o

pau se so the weight of ovaries might show a decrease after this
,

eve n t S u ch weight alteratio n s as well as the period at which

.
,

the max imu m weight is attai n ed are n o t recorded i n the litera ,

tu re It wo u ld therefore be of i n terest to ex ten d these observa

.

tio n s to h uman ovaries .

We may n ow su mmarize the vario u s factors which are r esp on

sible for the i n crease i n the weight of the ovaries In additio n to .

the formatio n an d en largemen t of the in dividu al ova these are ,

the growth of i n terstitial tissue the nu mb er of the mat u re fol ,

l i cl es ; of dege n erated follicles small or large ; of corpora l u tea

, ,

fresh or ol d as well as the co n ten t of blood The exact amou n t

, .

of blood however has n ot been determi n ed yet it is worth while

, , ,

to n ote here that either a hyperaemic or an aemic state is recog

n i z abl e accordi n g to the di ffere n t physiolo g ical states of the

ovary i n relatio n to heat .

436 HA YATO ARAI

to disc uss Ki ng ery s stateme n t co n cern i n g the proliferatio n of the

’

germ cells from the germi n al epitheliu m i n foetal life However .

,

i n the ovaries of albi n o rats after birth the man n er of the p rol if er
atio n of germ cells appears to be similar to that described by
Ki n gery In my case the oocytes from the g erm m al epitheli u m
.

begi n to form at abo u t t en to fift een days after birth The n u .

clei are large relatively to the cell body n early filli n g the cell, ,

which begi n s to grow an d e n large i n sit u i n the germi n al epi

theliu m The shape of these cells is at first more or less spherical
.
,

an d they are larger tha n the other epithelial cells A s they en .

large the adj ace n t epithelial cells are crowded to either side an d
, ,

when the developme n t of the egg cells proceeds fu rther they are ,

e n closed by the fl at t en ed epithelial cells th u s formi n g t h e pri

,

mary follicle .

A t this time the layer of t u n ica alb u gi n ea appears to be si n gle

or do uble b ut as the developme n t proceeds the follicles pass
,

thro u gh the tu n ica alb u gi n ea i nto the stroma This n ew forma .

tio n co n ti nu es with the growth of the ovaries an d the nu mber of ,

the follicles thu s formed may reach its maximu m at the period
of p uberty .

A fter p u berty this process of n ew formatio n may yet c on

t i nu e b u t is n o t as active as before p u berty
,
Withi n the first t en
.

days after birth this proliferatio n of n ew egg cells cann ot u su ally

be see n .

A ccordi n g to Ki n gery i n the mo u se the cavity i n the larger

,

follicles begi n s to appear betwee n fift een an d eightee n days after

birth an d a dege n eratio n of the egg cells sets i n at abo ut t h e
same time In the albi n o rat the cavity i n the large sized fol
.
-

li cl es begi n s to appear at abo u t twe n ty days after birth an d at

-
,

abou t twen ty six days t he middle sized dege n eratin g follicles

- —
,

accordi n g to my cl assi fi cat i o n are to be see n,

.

Ki n gery stated that the formatio n of egg cells from epitheliu m

is most rapid from three to twe n ty fi v e days after birth an d that
-

it goes o n practically up to sexu al mat u rity altho u gh more slowly ,

i n the later part of this period It is completed at forty or forty

.

hy e days after birth at which age as a r u le female mice are

, , ,

sexu ally mat ure .

 NUMBE R or O VA : AL BIN O R AT 43 7

My ow n observatio n s o n the rat n atu rally show some slight

di ffere n ce i n the age periods from those give n by Ki n gery In .

the rat the fir st appearan ce of the n ewly formed follicles is at

abou t t en days after birth an d their n u mber i n creases u n til abo u t
,

six ty to seve n ty days at which age as a rule ovu latio n occ u rs ;

,

b u t this process co n ti nu es at least to the age of o n e year tho u gh ,

n o t as rapidly as before .

The problem as to the origi n of the d efin i t i v e ova is n o t yet

d efi n i t el y settled Some au thors J en ki n son
. Kirkham
,
’
d H ol l an d er an d S o n n en b r o d t stated that the
d efin i ti v e ova are formed from the pr1 m ordi al germ cells thou gh ,

some observers Du sti n Karchakew i t sch A lle n an d Skrob an sky

“
,

( cited from O ogen esis i n the white mo u se

,
” Wi n iwarter
,

an d S ai n m o n t co n sider that the primordial germ cells

mai n ly dege n erate an d thu s as the r u le the defin i t i v e ova c an
, , ,

n o t be prod u ced from these Ki n gery holds the opin io n that the
.

d efi ni t i v e ova are developed from the primary follicles formed

after birth bu t he did n o t state clearly the relatio n betwee n the
,

primordial germ cells of other au thors an d the defin i t i v e ova of

his own desig n atio n .

There is n o t mu ch literat u re o n the n ew formatio n of the egg

cells after birth V an Be n ede n
. described i n the adu lt bat
the egg cells as formed from the germi n al epithelium L an e .

Cl ayp o n i n the ovary of the rabbit co n cl u ded that the fol ,

l i cl e cells an d t he i n terstitial cells are formed from the germi n al

epitheliu m These i n terstitial cells from their origin are p o
.
, ,

t en t i al egg cells an d u n der the proper stim ul u s are capable of

, ,

developi n g i n to ova V on Wi n iwarter an d Sai nm on t

. stated
that i n the cat at abo u t the age of three an d on e half to fo u r
,
-

mo n ths a re n ewal of the activity ( the third proliferatio n ) of the

,

germi n al epithelium prod u ces a n ew s u pply of germ cells which

develop i n to the defin i t i v e ova whe n all the egg cells of the first
,

( embryo n ic ) a n d seco n d ( shortly after birth ) proliferatio n s have

degen erated an d he stated fu rther that these defin i t i ve ova
come either e n tirely from the third proliferatio n or partly from it
an d partly from the u n di ffere n tiated cells left over from the sec

o n d proliferatio n .
43 8 HAYATO ARAI

Ki n gsb ury i n cli n es to the opi n io n that there is n o evi

de n ce of a n ew formatio n of ova by the third proliferatio n of
Wi n iwarter an d S ai n m on t Van der St ri ch t ( 1 1 ) do es n ot dis
.
’

c u ss at all the n ew formatio n of ova b ut simply states that i n

,

the ad u lt ovary i n cats the d efin i t i v e ova are derived from the
seco n d proliferatio n Felix
. i n describi n g the developme n t
of the ovary i n m an states that after the t un ica alb ugi n ea is
,

formed i n embryos of 1 8 0 mm i n len gth n o egg cells c an b e

, .
,

added to the i nterior of the ovary from the epithelial layer an d ,

thu s accordi n g to this au thor there is n o possibility of a n ew

, ,

f o rm ati é n of o v a from the germi n al epitheli u m .

Ki n gery i n his st u dies o n the white mo u se states that ,

there is a n ew formatio n of germ cells after birth an d that the ,

d efi n i t i v e ova come from the primary follicles Br i efly p u t .

,

Kin gery believes that at birth all the cells of the germin al epi
theliu m seem equ ally capable of developi n g i n to oocytes follicle ,

cells or epithelial cells thou gh it is n ot evide n t j u st what factors

,

are respo n sible for their even tu al fate A s the ovary becomes .

more matu re an d the cells better differen tiated this poten tiality ,

of the cells of the germi n al epitheliu m is lost an d after sex u al ,

matu rity n o more egg cells or follicle cells are derived from the
epitheliu m .

H e states also that all the egg c ells of the fi r st or embryo n ic

proliferatio n i n the mou se u n dergo dege n eratio n an d disappear .

In ovaries of mice from abou t seve n teen days after birth u p to

,

sexu al matu rity an d at the adult stage egg cells i n their folli
, ,

cles may be seen i n vario u s stages of dege n eratio n an d atresia .

This is the eviden ce of cou rse that a large nu mber of these

, ,

germ cells of embryo n ic origi n degen erate an d are resorbed .

The dege n eratio n of the d efini t i v e ova which i n all likelihood ,

sets i n before sexu al matu rity is co n ti nu ed thro ugh the whole

,

sex u al life of the i n dividu al as is well kn ow n Si n ce the n a large

,
.
, ,

n u mber of d efi n i t i v e ova degen erate an d si n ce these are sit u

,

ated more sup erfi ci ally than the primitive germ cells which are ,

mostly dege n erated it seems reaso n able to co n clu de that all the
,

primitive germ cells dege n erate an d are resorbed an d that all ,

the defini t i v e ova arise after birth from the germi n al epitheliu m .
440 HAYATO ARAI

fall . Followi n g this rapid fall the n ext period is represe nted by
,

o n e of co n sta n cy from abo u t twe n ty three u p to abo u t sixty -

three days This period of co n stan cy may be du e to a balan ce

.

between the dege n erati n g cells o n o n e han d an d the n ewly formed

germ cells o n the other A mo n g the dege n erati n g germ cells
.

m ay be some of the germ cells of the seco n d proliferatio n besides

the primitive germ cells an d tho u gh lacki n g evide n ce I am
, , ,

i n cli n ed to believe that at this time the maj ority of the primitive
germ cells have already disappeared .

The stateme n ts made by Wi n iwarter an d Sai n m o n t an d by

Ki n gery that all primitive germ cells are dege n erated an d the ,

View of the two former i n vestigators that all of the seco n d pro
literated germ cells have disappeared may very likely apply to ,

this prep u bertal period i n the rat as me n tio n ed above .

Followi n g this period of co n stan cy the c u rve agai n shows a ,

rapid decrease up to seve n ty days Si n ce duri n g this period the .

corpora l u tea begi n to appear we may assu me that at this period

, ,

also the cells are degen erati n g rapidly A ltho u gh eve n after .

p u berty there is some n ew formatio n of oocytes the s u dde n de ,

cre ase i n the nu mber of ova m u st be du e to a great excess of the

dege n erati n g cells .

Followi n g this seco n d rapid fall the curve n ow shows a gradu al

,

decrease up to 947 days thou gh there occ ur some slight fl u ct u a

,

tio n s probably merely i n dividu al especially at the earlier period

, ,
.

We may safely state that tho u gh some few ova might be n ewly
,

formed the decrease i n the total n u mber of ova is du e to the

,

dege n eratio n of the germ cells which represe n t those of the

seco n d proliferatio n .

I wish to emphasize the fact that the dege n eratio n of the prim
i t i v e germ cells begi n s from o n e day after birth a n d co n ti n u es u p
to abou t six ty three days at which time all of the primitive ova
-
,

may disappear c om p l et ely a res ult which i n ge n eral agrees with

w

the observatio n of Ki n gery .

Chart 2 also shows that the larger oocytes are relatively more
n u mero u s at a n earlier age tha n at p uberty .

This relatio n m ay i n dicate that the primitive ova before p u

berty especially at the earlier period e nlarge rapidly an d before
, , ,
 NUM BE R OF O VA : AL BIN O R AT 44 1

ripe n i n g may u n dergo dege n eratio n an d be resorbed From .

thirty days o n the proliferated defin i t i v e ova are added to the

primitive germ cells which are already prese n t a n d th u s the ,

d efi n i t i v e ova are very few at fi r st b u t i n crease rapidly c o n c om ,

i t a n t l y with the dege n eratio n of the primitive germ cells This -

.

process mai n tai n s an approximately co n sta n t nu mber of the

larger oocytes an d this co n ti nu es with more or less fl u c t u at i o n
,

u p to 9 4 7 days .

The e ffect of preg n an cy o n the total n u mber of ova is give n i n

table 4 b u t the n u mber of i n sta n ces is too small to f ur n ish a
,

b e sis for an y ge n eral s tateme n t .

Stratz co n siders that d uri n g preg n an cy there are small

follicles which become atretic before they can develop an d o n ly ,

toward the en d of preg n an cy the follicles begi n to grow to a

co n siderable S l z e an d come to mat urity O n the other ha n d .
,

L oeb mai n tai n s that while some of the large follicles dege n
er at e ,
the small medi u m an d some of the large follicles also
, ,

remai n witho u t dege n eratio n i n the last days of preg n an cy .

A ccordi n g to my ow n observatio n s the co n ditio n of the folli ,

cles as a whole agrees with part of the stateme n ts give n by the

two au thors me n tio n ed above tho u gh differi n g i n detail D ur ,
.

i n g preg n a n cy i n yo u n g rats the small follicles are f ew c o m p ared

with the mediu m an d large follicles In the later stages of preg .

n a n cy i n older rats the relative n u mber of small follicles i n cre a ses

co n siderably ( table while the nu mbers of the mediu m an d

large sized follicles are co n siderably redu ced whe n compared
-

with those fo u n d i n n o n preg n a n t rats of abo u t the same age

—
.

We i n fer from these facts that i n preg n an cy i n older rats the

maj ority of the mediu m an d large sized follicles have dege n er —

ated However this stateme n t is based o n a limited n u mber of

.
,

cases an d i n additio n cytological stu dies o n the cells were n o t

, , ,

made so that n o great emphasis c an be p u t o n the co n clu sio n

,
.

It is my hope to co n ti n u e the st u dy o n these problems i n the

fut ure .

TH E A M E R I C AN J O U RNA L or ANA O M Y
T , VO L 27 . NO 4
442 H AYA ro r
ARAI

O n the wei ght of the o vari es a n d the p resen ce f

o c or p ora l utea a nd

o f degen era te folli cles

The graph for the nu mber of ova which ill u strates the gro u p
i n which the ovaries possess the corpora l u tea grad u ally falls , ,

with some fl uctu at i on s from to 7 8 mgm i n the weight of

,
.

the ovaries ( chart

The two graphs which illu strate the n u mbers of corpora l u tea
show rather co n siderable fl u ct u atio n s b u t i n dicate that the ,

n u mber of small i n creases more rapidly tha n the nu mber of

,

large corpora l u tea a resu lt which is i n accord with expecta t io n

—
.

In this ( b ) gro u p the n the relatio n betwee n the weight of

, ,

ovaries an d the ova nu mber is also i n verse thou gh it is n o t so ,

striki n g as i n the ( a) gro u p an d the i n verse relatio n is du e i n

,

some measu re to the appearan ce of the corpora l u tea .

It is desirable n o w to disc u ss the i n flu en c e which the nu mber

of degen erate follicles may have o n the weight of the ovaries .

Previou sly it was me n tio n ed that an e ffort had been mad e to

divide the degen erate follicles i n to fo u r gro ups (p .

However on acco u n t of the great di ffic u lty i n maki n g eve n an

,

approximate estimate of the n u mber of the small dege n erated

follicles I have n o t attempted to record these i n the table For
,
.

co n ve n ie n ce the n u mber of the C an d D types of my cl assi fi ca

,

tio n or those which correspo n d to large dege n erated follicles an d

, ,

the B type or the middle sized dege n erated follicles are gro u ped
,
-
,

together an d the res u lts give n i n table 1 3 .

Table 1 3 s u ggests that i n the rat before p u berty the weight of

ovaries i n which the large dege n erated follicles are the more
,

nu mero u s te n d to be greater than that of the o v arl es i n which

,

these are less n umero u s A fter p u berty thou gh the similar re

.
,

latio n seems to hold n evertheless the prese n ce of the corpora

,

l utea obsc u res the relatio n G en erally speaki ng i n the ovaries

.
,

of yo u n g rats ( twe n ty six days old ) before the corpora l utea are
-

formed we fi n d a co n siderable nu mber of large de g e n erated

,

follicles .

From table 1 3 it is clear that before the formatio n of the cor

pora l u tea the nu mber of these follicles decreases slowly with t h e
444 HAYAT O ARAI

The nu mber of these small dege n erated follicles i n the ovaries

withou t corpora l u tea lies betwee n 3 00 an d 9 00 while i n the o v a ,

ries with corpora lu tea their n u mber varies from 600 to 1 40 0 .

It is a remarkable fact that the n u mber of these small follicles

is n ot partic u larly di ffere n t i n the ovaries of preg n an t as com
pared with the n o n preg n an t rats O fte n i n the ovaries i n which
-
.

corpora l utea are n o t prese n t the small dege n erated follicles are
ab u n dan t as for i n sta n ce i n the rat at 1 1 0 days i n which the
, , , ,

n u mber was 1 50 0 . A lso i n the ovaries with corpora l u tea the

n u mber of small dege n erated follicles may be very small as i n ,

the rat at seve n ty days i n which it was 300 The average n u m

,
.

T ABLE 14

Ta bu l a ti on of deg en er a ted f ol l i c les

M I DD L E I E D L A RG E O AL NUM E R
I S MA LL -
S Z T T B

F o u r t een c a s es i
w t ho u t cor p o ra l u t ea

Sv e en t een c as e s w thi cor p o ra l u t ea

bers of the differe n t sized dege n erated follicles in all cases are
show n i n table 14 .

O vu lati on

In regard t o the ovu latio n of mammals as well as the mat u ra

tio n of the ova there are n u mero u s observatio n s It is ge n erally
,
.

held that the mat u ratio n process takes place o n ly as the res u lt
of a sp ec i fic stim u l us which may follow cop ulatio n or the e n try
of the spermatozoo n i n to the ovid u cts Y et there are also n u .

m erou s observatio n s which show that the mat u ratio n process may
take place i n depe n de n tly of any s u ch stimu lu s For i n stan ce .
,

Weil i n the rabbit Sobotta ,

i n the mo u se Tafa n i ,
 NUM BER OF O VA : AL BIN O RAT 44 5

in the rat an d R u b aschki n

,
i n the g u i n ea pig stated that -
,

ovu latio n occ u rs spo n ta n eo u sly du ri n g heat an d is i n depe n de n t

of coitu s Iwan o ff
. showed that i n the rabbit preg n an cy is
also possible by the art ifi ci al i n semi n atio n an d Heape tried ,

a rt i fi c i al i n semi n atio n o n mares do n keys an d cows an d su c

, , ,

c ee d ed i n prod u ci n g preg n a n cy i n al l .

L oeb also believes that i n the g u in ea pig ov ulatio n occ urs —

i n the large maj ority of cases i n depe n de n tly of cop ulatio n .

Marshall an d J olly i n the dog an d i n the ferret described ,

the spo n tan eo u s ovu latio n at each of the earlier heat periods
du ri n g the breedi n g seaso n .

To test this matter i n the albi n o rat I u sed the co n trol females,

employed i n a stu dy of the su rvivi n g ovary after sem i sp ayi n g .

These co n trol female rats were separated from the males at

twe n ty days of age an d were kept together with the sem i sp ayed
,

rats of the same litter N evertheless i n the ovaries of these

.
,

two co n trol rats were fo u n d corpora lu tea at sixty two an d six ty -

n i n e days respectively ; that i s ov ulatio n occ u rred spo n ta n eo u sly

, ,

witho u t any associatio n with the male From all these res ults .

there is little do u bt that i n the maj ority of mammals ovu latio n

occ u rs reg u larly du ri n g the oestru s period an d is i n depe n de n t of
cop u latio n .

A ltho u gh ma n y i n vestigators have stu died the relatio n b e

tween me n stru atio n an d ovu latio n i n the higher mammals su ch ,

as the mo n key an d m an yet whether ovu latio n occ u rs before

, ,

du ri n g or after men stru atio n has n ot bee n determin ed It fol

, .

lows therefore that at least i n mo n keys as well as i n m an the

, , , ,

process of ovu latio n does n o t seem to be n ecessarily associated

with me n stru atio n tho u gh su ch a stateme n t is n o t d efi n i t ely
,

made by most of the observers .

A ccordi n g to Heape ovulatio n an d me n stru atio n are n ot

associated si n ce i n mo n keys me n stru atio n may occ u r periodi
,

cally all the year ro u n d b ut the seaso n for ovulatio n an d c on cep

,

tio n is limited V an H erw erden

. has also give n fu rther evi
de n ce that there is n o close co n n ectio n betwee n ovu latio n an d
men stru atio n either i n mo n keys or i n the aberran t lemur Tar
, ,

si u s spectr u m .
44 6 HAYAT O ARAI

In the case of the h u man female there are several opi n io n s as

to the u su al ti m e for the dischargi n g of the o vum Some au thors
.

report that ovulatio n occ u rs before me n stru atio n ; others d u ri n g ,

that process an d still others that it follows me n stru atio n

, , .

H erg esell holds that ovu latio n precedes me n stru atio n for
the reaso n that the most u su al period for ovu latio n i n the hu ma n
female as i n man y of the lower mammals w as du ri n g a defin i t i ve
, ,

oestru s followi n g the p reo est ru s ; for the period of most active
sexu al feeli n g is ge n erally j u st after the close of the men stru al
period whil e accordi n g to R ac i b orsky ( Traité de M en stru atio n
,
“
,

cited from Physiology of reprodu ctio n Marshall ,

” this ,

is also the commo n est seaso n for fertile coitio n Co n trary to .

this Bryce an d Teacher

,
hold that the ovu m is discharged
shortly after the cessatio n of the last men stru atio n .

To u chi n g the qu estio n whether a special stimu l u s i n du ces

ovu latio n i n woman O liver
,
regards the View that coitu s
accelerates ovu latio n as the more probable sin ce at this time ,

there is an i n creased blood su pply to the whole ge n ital tract .

Heape however main tai n s that the cau se which deter

,

min es the ru ptu re of the Graafian follicle is i n the rabbit the

, ,

stim u latio n of erectile tissu e du e to a n ervou s refl ex an d n ot

, ,

simply the result of i nter n al pressu re arisi n g from an i n creased

blood su pply or a greater qu an tity of liqu or follic uli .

Harper i n pigeo n s co n cl u des that ovu latio n requ ires

,

o n ly a slight stimu l u s me n tal an d that the prese n ce of sperm

,
‘
,
’

i n the ovid u ct cann ot be regarded as importan t K6lli k er

.

co n siders the cau se of r u ptu re as a simple mecha n ical process

becau se the irritatio n of vasomotor n erves i n creases the pressu re
of the liqu or follic u li th u s i n du ci n g the r uptu re .

From the foregoi n g it is evide n t that the real cau se of the rup
t u re of the Graafian follicle is n ot clearly established an d there ,

fore that the factors which may i n du ce ovu latio n requ ire fu rther
stu dy .

Whatever may be the fi n al co n clu sio n as to the cau se it is ,

clear that ovu latio n is closely related to the mat u ri n g of the ovary ,

as my prese n t stu dy i n dicates .

44 8 HAYATO A R AI

the n i nth week which probably correspo n ds to the age of p u berty

, .

M yers did n o t disc u ss the relatio n of this rapid growth to p u

berty The s u ggestio n is that these vario u s organ s might b e some
.
,

or all of them related to the process of ov u latio n i n the se n se of

,

formi n g stimu lati n g s u bsta n ces cau si n g ovu latio n a n d it is my ,

hope to make further stu dies alo n g this li n e i n the f u tu re .

Corp ora lu tea

I n ow wish to co n sider the relatio n betwee n the first appear

a n ce of corpora l u tea a n d p u berty .

For this p u rpose I have assembled the data from seve n ty n i n e —

albi n o rats ran gi n g i n age from thirty days up to abou t thirty

,

two mo n ths For some of them the weights of t he ovaries were

.

n o t determi n ed .

A s is show n i n table 1 5 the corpora l u tea fi rst appear i n a rat

,

at sixty on e days b u t after sixty on e days tho u gh corpora l utea

-
,
-
,

are prese n t i n most cases they are n o t prese n t i n all The ab

, .

sen ce of the corpora lu tea i n the rats after six ty on e days is always -

associated with a bo dy weight too small for the age showi n g poor ,

n u tritio n The weights of the ovaries are also small

. .

I have n ext rearra n ged some of the data give n i n table 1 5 , ,

accordi n g to the body weights of rats omitti n g those with body ,

weights less tha n gra m s an d greater th an grams ( table

We n otice the fi rst appeara n ce of the corpora l u tea i n the rats

w ith a body weight of grams b ut they are n o t always prese n t
,

u n ti l the rats reach 1 00 grams i n body weight .

Beyo n d 1 0 0 grams w e always fi n d corpora l u tea i n ovaries .

S o far the n as the body weight is co n cer n ed the fi rst appear

, , ,

a n ce of the corpora l u tea occ urs at from u p to 1 00 grams .

These body weights of to 1 00 grams correspo n d to sixty o n e -

an d seve n ty o n e days respectively as give n i n D o n aldso n s table

’
—
, , .

Thu s i n my series the appeara n ce of p u berty approximately c o

i n cides with the observatio n of D o n aldso n From these
data it appears that while p u berty is attai n ed as a ru le betwee n
sixty on e to seve n ty on e days i n rats that have grow n app ro x i
- -
 N U MBE R OF O VA : AL BIN O R AT 44 9

L
TA B E 15

The r el a ti on between the co r p o ra l u tea an d the a g e, bod y wei g ht an d bod y l en g th

B ODY B O DY W E I G HT
O O
CO R
A GE A GE
W EI G H T -
1
T
HTP R
L EN G H O VA R I E L U E A
OF B K
S T

mm mm
4 50 HAYATO A R AI

mately n ormally yet mal nu tritio n as represe n ted by a d efi ci en t

,

body w eight may delay its appearan ce or eve n e n t i rely preve n t i t .

This is i n accord with laboratory observatio n s o n breedi n g .

L
TA B E 16

The re l a ti on between the cor p o r a l u tea an d the bod y wei g ht . Body wei g ht, 7 5
, to
1 1 5 g r a ms

B ODY EI GH B O D Y L ENG H W E I GH T O F B OH C O RP O R A L U E A
A GE \V T T
O VA R I E S
T
T

g r a ms mm

Fi n ally I have arran ged the data give n i n table 1 5 accordi n g

,

to the body le n gth of the rats R ats whose body le n gths are less
'

than 145 mm an d greater tha n 1 55 mm have bee n elimi n ated

. .
,

an d the res u lts are show n i n table 1 7 .

4 52 HAY ATO ARA I

compared the res u lts obtai n ed by the three di ffere n t methods i n

table 1 8 .

It is at o n ce clear from table 1 8 that whe n body le n gth is take n

as a criterio n the ra n ge at which period the corpora l utea fi rst
,

appear is least n amely the age of sixty two to sixty fi v e days

, ,
- -
.

Corp or a lu tea i n p regn an t a n d n on - p regn a n t ra ts

In regard to the corpora lu tea there are nu mero u s observa

tio n s especially o n their origi n V o n Baer
,
co n siders that .

the corpora l u tea co n sist e n tirely of co n n ective tiss u e an d i n ,

their formatio n the follic u lar epitheliu m has n o share O h the .

T ABLE 18

Con d i ti on s deter m i n i n g the a p p ea r a n ce o f the fi rs t c or p ora l u tea

IME IR A G E I N D A Y R O M
M E HOD C O MP A R I ON A PP E A R ANC E C O R P O R A D O NA L D O N A BL E
TH E T O F F ST
TH E S F

L U EA
TH E T OF S OF ’
S S T
T

By ag e 62 t o 1 10 d y
a s

By b dy
o we i ght to g ra ms
1 48 t o l 50 mm .

other han d Bischo ff ,

co n cl u ded that the lu teal cells were
formed by the hypertrophy of the epithelial cells of the u n dis
charged Graafi an follicles E ver si n ce these stateme n ts were .

made by Vo n Baer an d by Bischo ff they have bee n t h e s u bj ect

of disc u ssio n .

Marshall stated that if the discharged ov u m fails to b e

come fertilized the corp u s lu te u m goes o n growi n g for a short
,

time an d the n dege n erates In the smaller a n imals it disappears

.

after a comparatively short time If o n the other ha n d c o n cep .

,

tio n follows ovu latio n the corp u s l u te u m co n ti n u es to i n crease

,

i n size u n til almost the middle of preg n a n cy .

L oeb 1 1 ) also fo u n d i n g u i n ea pigs that the corpora l utea -

which are formed withou t preg n an cy are mu ch smaller an d ,

shran k more rapidly than those the formatio n of which was fol
lowed by preg n an cy .
 NUM BE R OF O VA : AL BIN O RAT 453

On the oth er han d Sobotta from his st u dy o n the mo u se

, ,

expressed the View that the persiste n ce of the corpora lu tea an d

their u ltimate size are n ot alt ered by co n ceptio n .

In the prese n t st u dy we have exami n ed fo u r preg n a n t rats ,

an d fo u n d that as L oeb described the corpora l u tea i n the preg

, ,

n a n t rats were larger tha n those i n rats n o t preg n a n t How .

ever the di ffere n ce is slight The completely formed corpora

,
.

lu tea at abo u t t en days after co n ceptio n j u dgi n g from the size of ,

the fetu s are abo u t

,
mm i n diameter This size has dimi n . .

i sh ed to abo u t mm i n diameter abo u t seve n tee n days after

.

preg n an cy that is i n the later phase

, ,
.

S e lo n g as I did n o t collect the material with a View to stu dyi n g

the fate of corpora l u tea I am u n able to disc u ss their age Ii
,
.
,

however w e divide the e n tire pop u latio n of corpora l u tea i n to

,

two groups the larger an d t h e smaller o n es it is possible to

, ,

st u dy the i n flu en ce of these o n the weight of ovaries .

A s is show n i n tables 3 6 8 an d 1 0 both the total n u mber of

, , , ,

the corpora lu tea as well as the nu mber of small o n es show an

, ,

u n mistakable te n de n cy to i n crease despite great i n divid u al v ari ,

atio n accordin g to the age or other measu reme nts of the rats
,
.

O n the other han d the n u mber of corpora l u tea of large size

,

remai n s approximately co n stan t .

From these resu lts we co n clu de that after pu berty the acc u mu
latio n of the smaller corpora lu tea is mai nly respo n sible for the
i n crease i n the weight of the ovaries with i n creasi n g age .

The degen era ti on o f the folli cles

So far I have simply stated the relatio n which exists betwee n

the total nu mber of ova an d of the dege n erated follicles ; b u t I
n o w wish to co n sider these dege n erated fo l licles i n their relatio n

to the corpora lu tea A s has bee n me n tio n ed already ( table

.

the dege n era t ed follicles are n ot fo u n d i n the rat u n til abo u t

twe n ty six days The failu re to fi n d an y degen erated follicles
—
.

u p to this time may be du e partly to the extreme d iffi c u l t y i n

recogn izi n g them an d partly to a possible resorptio n immediately

after they are formed However this may b e after twe n ty six
.
,
-
4 54 HAYATO ARAI

days the dege n erated follicles were first fo u n d an d these probably

origi n ated from e n larged primitive follicles .

5
15; L oeb while stu dyi n g the cyclic chan ges i n the ovaries of
g u i n ea pigs fo u n d that at eightee n days of age the dege n erative
—
,

process i n some follicles had bee n e n tirely completed an d c on ,

h ective tissu e had beg u n to grow i n to the cavity Thu s the de .

ge n erative processes occ u r at abo u t the same phase both i n the

rat an d i n the more precocio u s gu i n ea pi g -
.

Moreover L oeb states that associated with ovulatio n all the

, , ,

follicles with the exceptio n of very small o n es degen erate The

, , .

ge n eral degen eratio n of the follic u lar granu losa cann ot be see n b e
fore ovu latio n This su dde n dege n erative process as well as
.
,

typical follic ular dege n eratio n is qu ite i n depe n de n t of coit u s

, .

H e fu rther f ou n d that i n n ewly r u pt u red follicles the dege n eratio n

of the granu losa is show n except i n the very small follicles while
, ,

i n a large maj ority of the follicles almost the whol e gra nu losa is
fo un d i n the process of dege n eratio n an d this degen eratio n is es
,

sen t i all y i n depe n de n t of cop u latio n an d of preg n a n cy b ut di ,

re c t ly co nn ected with o v ulatio n Fu rthermore from the age of

.
,

the corpora lu tea the ultimate fate of the follicles i e whether

, ,
. .
,

these will disappear or n o t may be i nferred A t a given time .

,

approx imately t en days after ovu latio n a certai n equ ilibriu m is ,

reached between the follicles which u n dergo dege n eratio n an d

those which grow further A mo n g those which were growi n g
.
,

the degen erative process may also take place .

Tho ugh i n my st u dy the exact age of the corpora l utea cann ot

be determi n ed w e c an ass u me for the two preg n an t rats whose
, ,

ages are eighty an d n i n ety fiv e days respectively that the cor

-
, ,

pora l u tea ( from the estimated age of the fet us ) were formed
withi n the t en precedin g days In these cases the n umber of
.

the dege n erated follicles of middle an d large sizes is far greater

than i n the other rats at abo u t the same age A ltho ugh i n these .

cases the corpora l u tea may have bee n prese nt withi n t en days
after ov ulatio n as w as i nferred yet it is premat ure to co n cl ude
, ,

from my st u dy that all of the follicles of large a nd medi um size

u n dergo dege n eratio n after the appeara n ce of relatively n ew

corpora l utea as L oeb stated The n umber of middle sized

,
.
-
4 56 HAYAT O A R AI

li c l esis co n siderably i n creased as compared with their n u mber

before p u berty This fi n di n g is di ffere n t from that of L oeb b u t
.
,

the di ffere n ce may be du e partly to the differe n ce i n the a n imals

u sed However it seems to me more probable that the ap
.
,

p eara n ce of the corpora l u tea gives an imp u lse to the dege n era
tio n of the small follicles .

A t a ny rate the p erce n tage of dege n erated follicles after

,

p uberty is higher than before p u berty as is show n i n the above ,

calc ulatio n where the nu mber of dege n erated follicles is as high

,

as abo u t 1 6 per ce n t of the total nu m ber of ova Therefore the .

,

decrease i n the n u mber of ova accordi n g to the age seems to be

cau sed pri n cipally by the dege n erated follicles We c an n eglect .

the ripe n ed ova eight to t en of which are discharged i n to the ovi

,

d u cts at every ovulatio n period as this nu mber is i n si g n i fi can t

,

i n relatio n to that of the dege n erate follicles .

Sobotta an d Burc k ar d stated that i n white rats the maxi

m u m n u mber of ripe ova which e n ter the ovid u cts is altogether
thirtee n from both ovaries a n d the mi nimu m is fo u r altho ugh ,

eight to t en ova are more u s u al The maximu m n u mber of ripe

.

ova from o n e ovary is said to be eight .

A ccordi n g to D o n aldso n the largest litter n oted i n the com

,

m o n albi n o is seve n tee n while Ko l a z y

,
also reported a litter
co n sisti n g of seve n tee n y o ung .

A c cordi n g to these observatio n s the highest n u mber of ova

,

discharged i n to the fallopia n t ubes is seve n tee n a n d the lowest

is fo ur U s u ally it is eight to t en The average nu mber of the
. .

mat ure follicles i n the ovaries at several ages however is twe n ty , ,

o n e an d therefore abo u t half of these mat u re follicles m u st r u p

,

t ure an d discharge their ova i n to the ovid ucts and the remai n i n g ,

half m ust un dergo atresia .

 NU M BE R OF O VA : AL BIN O R AT 457

Comp ari son between man an d the ra t in r egard to the p ostn a ta l

c han ges i n the ovar i es

It may be worth while to co mpare the res ults obtained from

th ese st u dies o n the developme n t of the ovary i n the rat with
those i n m an especially d uri n g the period i n which the dege n er
,

atio n of the primitive germ cells an d the new formatio n of deh

n i t i v e ova is most active .

If we take six ty fi v e days as the mean ag e for the first ovula

—

tio n we fi n d that the rat is disti n ctly precocio u s for sixty fiv e

,
-

days correspo n ds to abo u t sixty hy e mo n ths of h u ma n life or

,
—
,

ro ug hly hy e an d a half y ears A ccordi n g to Vi er o rdt s table

,
.
’
,

the first me n str u atio n i n m an may occ ur i n tropical co untries

at eight years b ut more commo n ly a y ear or two la ter
,
.

0 n the other ha n d the cessatio n of the breedi n g period i n

,

the female rat at eightee n to twe n ty mo n ths 4 5 to 50 years )

agree very well with the appearan ce of the me n opau se which
occ u rs i n m an betwee n forty hy e an d fift y years -
.

A s is to be expected i n divid ual rats may breed for a lo n ger

,

period an d Ki n g
,
has reported a female beari n g a litter of
o n e at twe n ty two mo n ths an d as table 1 5 shows the rat at 9 4 7
—
, ,

days thirty o n e mo n ths) had n ewly formed corpora l utea i n

-

its ovaries .

SUMMAR Y
1 The total n umber of ova i n both ovaries w as co un ted i n
.

thirty n i n e albi n o rats ran gi n g i n age from birth to 947 days

-
.

2 In relatio n to the body weight the size of the ovaries i n

.

creases to a maximu m at 3 3 grams of the body weight the n de ,

creases up to p uberty after which it i n creases rapidly an d reaches

,

the seco n d maximum .

The ovary weight accordi n g to age shows co n ti nu o u s i n crease

up to thirty o n e mo n ths ( table
-

3 The weight of the right ovary is less than that of the left
.

abo ut 90 per ce n t while the total n umber of ova i n the right

—

ovary is slightly more than i n the left tho ugh the di ffere n ce is ,

sm all ( table
TH E A M E R I C AN J O U RNA L or ANA O M Y V OL
T , . 27 , NO . 4
4 58 HAYAT O A R AI

4 . In ge n eral the n u mber of ova decreases with age The .

t otal n u mber of ova i n both ovaries decreases rapidly from

at birth to abo u t at twe n ty three days From twe n ty
-
.

three to sixty three days the n u mber is n early co n sta n t

-

to It the n decreases agai n rapidly ( to abo u t 6600 ) at

seve n ty days D u ri n g this last period ovulatio n u s u ally occ urs
. .

From seve n ty days up to the thirty o n e mo n ths there is a slow —

decrease to abo u t 2 00 0 ova In ge n eral this decrease res ults

.
,

mai nly from the dege n eratio n of the primitive ova b u t i n part ,

from that of the defi n i t i v e ova ( tables 2 an d 3 charts 1 an d ,

5 In n o n preg n a n t rats the perce n tage of larger sized ova to

.
— —

”
the total n u mber remai n s n early co n sta n t In yo u n g preg n a n t .

rats the perce n tage of ova more than 2 0 M i n diameter is greater

than i n the n o n preg n an t rat b u t i n older preg n an t rats this per
—
,

ce n tage decre as es These co n clu sio n s are however based o n

.
, ,

fo u r i n stan ces o n ly .

6 The graphs ill u strati n g the cha n ge i n the total n u m ber of

.

ova are similar i n form whether they are based on the body
weight or the body le n gth ( table 6 chart ,

7 With the i n creasi n g weight of the ovary the total n u mber of

.

ova decreases The i n crease i n weight is associated before p u

.

berty pr i n c i pally with the formatio n of lar g e dege n erate follicles

together with mat u re follicles an d growth of co nn ective tiss u e ,

b u t after p u berty the i n crease depe n ds mai n ly o n the ac cu mu

l atio n of small corpora l u tea i n additio n to the mat u re an d dege n
era t e f o l li c l es an d co n n ective tiss u e A fter p u berty the n u mber of
.

large corpora l u tea is abou t the same i n all ovaries an d these , ,

therefore are n o t respo n sible for the reg u lar age cha n ges i n the
,

w eight of ovaries ( table 1 0 chart ,

8 In albi n o rats the n ew formatio n of the egg cells takes place

.

after birth from the germi n al epitheliu m These ova grow i n .

situ an d as developme n t proceeds are covered by the adj ace n t

, ,

epithelial cells an d exte n d i n to the stroma passi n g thro u gh the ,

t un ica alb ugi n ea From these n ewly formed germ cells the defi n i
.

tive ova appear to develop begi nn i n g from abo u t the seco n d

,

week after birth an d the formatio n of them is most active i n the

,

period betwee n the third a n d n i nth weeks D uri n g p u berty .

4 60 HAYATO ARAI
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d es rat s . A rch . It al . de Bi l o .
, vol . 11 .

V I E R O R DT , H . 1 906 D a t en un d T b a el l e n fu r M edi z i n e r . 3 A n ti . G u s t av
Fi sc h e r . J en a .

WA L D E Y R W D ie G esc h l ec h t sz el l en In H db u ch v gl i
”b y
E , . 1 906 . an der er e ch

en d xp i m t ll E t w i k l
en u n d e er en e en n c e u n g s l eh r e d er Wi b r el t er e , i
O k sH tw i g t B d t T i l
ar er ,
e rs e r an , ers er e ,
ers t e H a li te, S 351 .

W IL E ,
C . 1 8 7 3 B i t ag K t i d B f e r e zur en n n s er e ru c ht u n g und E n tw c ik el u n g d es
Kan n i n c h en e i es . Wi en M e d J a h rb u c h . .

W I E DE R S H E I M R ,
. 1 8 97 El em e n t s o f t he c om p ar at iv e an at o m y fv o e rt e b r a t es .

2 md ed . T r an s . by W N . . P k ar er . Mac m i ll an , L d n p
on o ,
. 3 68 .

VON WI NI W A R R TE ,
H . ET S A IN MO N T, G . 1 908 N v ou e ll es r ec h er c h e s su r l
’
o v o

g en ese et l o r g an o g en ese de l
’
ov a i re d es m am m i f er es ( ch at ) . A rc h .

Bi ol .
,
T 24 . .

1 908 Ub e er d ie au ss c h l e s sl c h i i p o s t f et a l e B i l d g d d fi i t i v
un er e n en Ei er

b e i d er K tz a e . A n at . Anz .
,
Bd 3 2 . .
R es ume n por a utor L eo n Au g u st u s Hau sman
cl , .

U n iversidad Cor n ell Ithaca , .

In v est i g a c i é n microl ogica de la estr u ct u ra del pelo eu los

M onot rem as .

L o mismo O r n ithorhyn ch u s Tachygloss u s posee n varios

q ue
tipos difere ntes de pelo en su c uerpo El tipo de pelo q ue parece .

ser c ara ct eri st i c o de los M o n o t rem as es el d e tipo muy aplanado ,

represen tado por los pelos de n omin ados pelos esc ut eliform es en
O rn ithorhyn ch u s y por los pelos llamados pelos o n d ulosos apla
n ad o s y pelos espi n osos eh Tachygloss u s O rn ithorhyn ch u s posee .

un re c u b r1 m 1 en t o de pelos semej a n tes a los q u e forma n la piel

de otros mami fero s los c u ales faltan en Tachygloss u s L a es

,
.

t ru ct ura de dichos pelos es semej an te a la del tipo ge n eral de

pelos c ili ndri c os qu e se e n c u e n tran en la mayor parte de los
mam i fero s .

T ra n s l a t i by J é F N m d
on os o ez

C d i l C ll g N w Y
.

o r n ell Me ca o e e, e o rk
A U HOR A
T
’
S BSTRAC HI P AP E R
T OF T S i s s u nn

BY TH E B I BLI O GR A P H I C E RV IC E J LY
S , U 12

A M IC R O L O G IC A L I N VE STI G A TI O N O F TH E H A I R
ST R U CT U R E O F TH E M O N O T R E M A T A
LE O N AU G USTUS HAUS MAN
Z oo l og i ca l L a bor a tor y, Co rn ell U n i vers i ty

F O U R P L A T E S ( N I N E T Y E I G H T F I G UR E S )-
AN D T H REE TE X F I G UR
T ES

C O N TENTS
I n t ro du t i c on

C o l l ec t i o n of ha i r sam pl es .

P pre i
a r at o n o f ha i rs for m i c ros c o pi c al e x a m in t i
a on

T he h a i r of Oi ni th m hyn ch us a n a ti n u s . 0 0 0 0 0

S u mm a r yf or O r n i tho r hyn chi t s u

T he ha i r an d s pi n es o f Ta c hyg l oss us hyst i i x

S mm y f
u ar or Ta c hyg l oss u s
B ib l i g p h y
o ra

I NTR O DU C TI O N
The observatio n s an d co n cl u sio n s embodied i n the prese n t
co n tribu tio n are the o utgrowth of a comparative stu dy of the
microscopic stru ctur e of the hairs of the mo n otremes O rni tho ,

r hyn chu s a n a ti n u s a n d Ta chyglossu s hys tri x a n d i n cide n tally ,

a n d for compariso n of members of all of the existi n g families of

mammals ex cept the Cetacea ,

.

Col lecti on of hai r sa mp les

The maj ority of the hair samples were collected by the writer
from ski n s an d mou n ted specime n s i n the collectio n s at Corn ell
Un iversity a n d from those i n the A merica n M u se u m of N at ur al
History i n N ew York City an d i n the Un ited States N at i o n al
Mu seu m i n Washi ngto n O ther i n stitu tio n s ki n dly fur n ished .

man y samples .

Withi n each family samples were exami n ed from species rep

resen tin g variou s types of e n viro n me nts an d from each family ,

463
4 64 LE O N AUGUSTUS HAUS MAN

three species were selected w here possible to ill u strate the t ypi , ,

cal trichologie str u ct u res i n their vario u s m o di fi c at i o n s .

It was fou n d that after collectio n the hair samples were most
, ,

satisfactorily disposed of by placi n g them together with their ,

data slips i n gelati n veteri n ary caps ules 0 n e o u n ce caps u les

, .
-

were u sed for the lo n ger hairs a n d half an d qu arter o u n ce for ,

-

the shorter o n es These co n tai n ers were m u ch more satisfactory

.

than glass Vials for they co uld be tra n sported loosely with little
, ,

care witho u t da n ger of breakage ; were less expe n sive an d mu ch

, ,

lighter i n weight .

P r ep ar a ti on of ha i rs f or mi cr oscop i ca l ex a mi n a ti on

1 . P r ep ar a ti on f or First
ex a mi n a ti on o f cu ti cu lar sca les .

method dry mou n tin g Some au thors recomme n d placi n g the

—
.

hair directly be n eath the microscope o n a slide an d maki n g the

exami n atio n dry withou t previo u s preparatio n This method
,
.

was tried an d while it revealed i n ge n eral the arra n geme n t of the

,

scales i n those hairs which are coarse an d i n which the scales are
un u su ally promi n e n t it failed to yield acc u rate res u lts It was
,
.

fo u n d to be qu ite esse n tial to have the hair shaft perfectly clea n ,

otherwise du st fib ers which adhered to it were qu ite likely to b e

mistaken for the fi n e tran sverse marki n gs which i n dicate the
o u tli n es of the scale edges The simplest method of preparatio n .

employed w a s that of washi n g the hair carefu lly i n a solu tio n

composed of equ al parts of 95 per ce n t alcohol a n d ether This .

removed all the oleagi n o u s matter from the su rface of the hair
shaft an d made it diffi cul t for du st fib er s to hn d lodgme n t u po n
it
. It w a s the n tran sferred to a clea n slide covered with a ,

cover glass an d allowed to sta n d o n a tripod over a low alcohol

-
,

flame u n til the whole had become perfectly dry Throu ghou t .

the e ntire man ipulatio n of hairs it was fo un d to be absol u tely

imperative to keep all u te n sils an d i n stru me n ts an d especially ,

all glassware scr up u lo u sly clea n A dry exami n atio n of the hair
,
.

was the n made This sort of treatme n t was fou n d to be eft ect i v e
.

i n the exami n atio n of hair whose scales were large or promi n e n t ,

s u ch as the hair of some of the Cervidae or of the Camelidae In ,

.

the great m aj o rl ty of cases however it became n ecessary to have , ,

4 66 L E O N AUGUSTUS HAUS MAN

face of the hair shaft prese nts an alter n ati n g series of tra n sverse
ridges an d tra n sverse depressio n s du e to the overlappi n g i mbri
, ,

cate c utic u lar scales The method devised for maki n g clear the
.

o u tli n e of the scales is i n pri n ciple to lod g e fi nely divided color

, ,

i ng matter i n all of these tra n sverse depressio n s leavi n g the ele ,

v a t i on s u n colored
. This was accomplished i n the followi n g m an
n er : The hair shaft was fi r st washed i n a sol u tio n composed of

equ al parts of 9 5 per cen t alcohol an d ether to free its s urface ,

from oily matter It w a s then heated very slightly or fa nn ed

.

ge n tly to in su re complete dryi n g an d the n immersed i n a 9 5

, ,

per cen t alcoholic sol u tio n of ge n tian Violet or safra n i n After .

remai n in g i n this solu tio n for a mi nu te or more it was removed ,

with forceps an d held i n a ge n tle draft of air or i n the c urre n t of

.

warm air risi n g from a b un sen fl am e un til the alcohol had com
p l et el y evaporated . The ge n tia n Violet or safra n i n it was fo un d , ,

had been deposited from the sol utio n an d had gathered i n all the
tran sverse depressio n s o n the s urface of the hair thu s marki n g ,

o u t clearly the o u tli n e i n sharp color of every c u tic u lar scale

, , .

The most delicate scale sc u lptu ri n gs which are capable of deter

mi n atio n u po n the fi nest of the hairs with the immersio n ob j ec
t i v es were by this method of treatme n t re n dered plai n tho u gh ,

for the very finest of the hairs this method was combin ed with
exami n atio n u n der obliqu e illu mi n atio n hereafter described It
, .

was foun d however that while this method gave almost ideal
, ,

results with some hairs at the very fi rst trial it was n ecessary ,

with other hairs to subj ect them to the processes again and agai n
before the evaporatio n of the alcohol deposited the pigmen t
u n iformly i n the c u tic u lar grooves over an y co n siderable portio n

of the hair s urface .

O ther stai n s which g o eas i ly i n to sol utio n i n the 9 5 per cen t

alcohol an d are readily deposited u po n its evaporatio n s u ch as ,

iodin e (with potassiu m iodide) methyl gree n methyl and methyl

, ,

en e bl u e an d tol u idi n bl u e were also u sed b u t for some u n k n ow n

, , ,

reaso n the best su ccess was obtai n ed with the ge ntian Violet and
safran i n.

In the case of those hairs i n which the scales are so promi n e n t

as to proj ect n otably ou t from the shaft a very striki n g pro file
,
 HAI R STR U C TU R E OF THE MO N O TR E MATA 467

was obtai n ed by soaki n g the whole hair ( after washi n g i n the

ether alcohol sol utio n ) for several mi nu tes i n a 1 0 per ce n t aqu e
-

o u s sol u tio n of ca u stic soda yet n o t lo n g e n o u gh to re n der the

,

ha i r slimy or to distort the scales an d the n dippi n g it i n to a

,

pin k solu tio n of safra n i n an d 8 2 per ce n t alcohol The e n tire .

su rface of the hair assu med a pi nk color an d while t he scale ,

sc ulptu r i n gs over the surface of the hair were obliterated by the

u n iformity of the coloratio n yet the p r o fil e of the hair shaft stood
,

o u t with great clarity agai n st the white light of the microsc o pic
h eld or eve n more striki n gly agai n st the black backgro u n d whe n
,

very obliqu e ill umi n atio n an d a low power obj ective were u til -

i z ed
. This man ip u latio n w as ofte n resorted to to determi n e the
relatio n of the tran sverse scale sc u lptu ri n gs o n the su rface of the
hair to the profil e of the serrate edge .

Fig ur e A
For immediate exami n atio n the hairs stai n ed to show the scale
sc u lptu ri n gs o n the surface were pu t i nto temporary dry moun ts
by fasten i n g over them a cover glass to u ched abou t its edges with
-

balsam Perman en t mo un ts were obtai n ed by ri n gi n g the cover

.

g lasses with cemen t u po n a turn table They were often for i m

-
.
,

mediate u se also mou nted i n very Visco u s gelati n g um dammar

, ,

or balsam It was fo u n d that if very flui d balsam dammar or

.
, ,

gelati n were u sed the colori n g material was qu ickly dissolved

,

from o ut the scale depressio n s an d distrib u ted throu ghou t the

whole of the mou n ti n g medi u m For the coarser hairs dry .

mo u n ts were the most su ccessfu l ; for the fi n er the glyceri n bal , ,

sam an d dammar mo u n ts
,
.

It was frequ e n tly fo u n d desirable to trace completely arou n d

the hair the co urse of the marki n gs i n dicati n g the edges of the cu t i c
u lar scales partic u larly i n the st u dy of the form of the coro n al
,

type of scales To re n der the rotatio n of the hair u n der the high
.

est powers of the microscope practicable the apparat u s show n i n ,

fi g ur e A was devised a n d termed hair rotator It was c on

‘ ’
.
,
468 L E O N AUGUSTUS HAUS MAN

stru cted as follows : Upo n a glass slide were faste n ed with Can ada
balsam two small corks of firm text u re tra n spierced by fi n e cop
, ,

per wires with their opposite e n ds be n t i n to loops Betwee n the

,
.

i n n er loops of these wires a n d betwee n the corks the ha i r u n der ,

st u dy was stretched an d faste n ed at each en d with droplets of

balsam or Visco u s mu cilage The copper wires were n ow draw n
.

o u t carefu lly away from o n e a n other u n til the hair betwee n them

w a s stretched tau t a n d the whole device placed upo n the stage

,

of the microscope By ge n tly tappi n g the o u ter loops of the

.

copper wires with a dissecti n g n eedle the hair co u ld be with the ,

greatest delicacy t ur n ed i n either directio n while u n der exami n a

tio n It was discovered that a small drop of a 2 5 per ce n t aqu e
.

o u s sol u tio n of cau stic soda or potash placed n ear o n e extremity

of the stretched hair softe n ed this portio n to s u ch an ex te n t that

the hair co uld be more easily rotated by t u r n i n g b u t o n e of the
copper wire loops N ot o n ly co u ld the hair be rotated b ut also
.
,

stretched slightly an d this was a n advan tage ofte n i n asmu ch as

, ,

the le n gthe n i n g of the cortex slightly separated the c u tic ular

scales an d re n dered the depressio n s betwee n them a t ri fle deeper .

This device proved equ ally u seful also i n the exami n atio n of
the medu lla The hair was fir st washed i n the ordin ary w ay as
.
,

has bee n described an d then cleared i n clove or cedar oil after

, ,

which it was dried betwee n two pieces of le n s paper an d stretched

i n the hair rotator as previo u sly described The c on fig u rat i on .

of the medullary cells an d the relatio n of i n dividu al an d gro u ps

of cells to each other was by this mea n s bro u ght ou t i n the most
satisfactory ma nn er Several of the hairs were after havi n g
.
,

been cleared i n xyle n e allowed to remai n i n a bath of balsam for

,

several hou rs an d the n take n out hu n g up like ca n dles an d

, , ,

allowed to dry covered with a thi n hlm of harde n i n g balsam .

These were the n mo u n ted i n the hair rotator an d th u s a com ,

p l et ely balsam mo u n ted hair sec u red for exami n atio n It was
-
.

fo u n d however that i n the mai n this precedure gave n o more

, ,

satisfactory resu lts tha n simply cleari n g i n clove or cedar 0 11

before mo un ti n g i n the rotator tho u gh there were i n sta n ces i n
,

the cases of some of the larger hairs i n which it was thou ght that
the cleari n g i n xyle n e i n su red a slightly greater cl ari fic at i o n of
47 0 L E O N AUGUSTUS HAUS MAN

eati n g the ex te n t of so me of the c u tic ular scales The disposi .

tio n of the colors show n i n dicated ofte n the locatio n an d relative

thick n ess of the vario u s hair eleme n ts an d n ot i nfrequ e n tly ,

cau sed the tra n sverse edges of the c u tic ular scales to sta n d ou t
promi n e n tly i n black tracery agai n st the b ackgro u n d of some
vividly co n trasti n g color The state of fu sio n of the cortical
.

cells w a s ofte n also i n dicated by the appeara n ce of certai n char

a c t er i st i c color associatio n s By arra n gi n g the hairs u n der
.

exami n atio n at di ffere n t a n gles o n the slide an d by i n cl u di n g i n

o n e mo u n t hairs of ma n y differe n t species vario u s beau tifully ,

colored demo n stratio n s of differe n ces an d similarities were ob

t ai n ed . It was fou n d most satisfactory to mo un t hairs desig n ed
for polariscopic exami n atio n i n some heavy oil castor oil )
or balsam This treatme n t was i n fact imperative for those
.
, ,

hairs whose medullas were the obj ects of i n vestigatio n .

2 P r ep a r a ti on for the ex ami n a ti on of the medu l la

. The meth .

ods u sed to re n der the scales of hair promi n e n t obsc ure the med '

u lla . Co n sequ e n tly it was fo u n d n ecessary to devise some mea n s

of re n deri n g the hair shaft tran spare n t i n order to bri n g the med
ul l ary cells or chambers i n to Visibility This was ac complished .

i n the followi n g ways :

First method cleari n g i n water This is the m et ho d of clari
—
.

fi c at i o n commo n ly u sed for hairs of a n o t too great diameter i n

ge n eral It works fairly well with the fin er hairs which lack pig
.

m e n t i n the cortex a n d whose c u tic ular scales are n o t close ly set

together which is ta n tamo un t to sayi n g that it works well with
,

b u t very few .

Seco n d m et h o d cl eari n g i n oils It was fo u n d that cleari n g

~
.

the hair i n oils of vario u s sorts s u ch as oil of bergamot oil of cloves

, , ,

oil of cedar oil of origanu m an d cast or oil te n ded to obsc u re

, , ,

almost e n tirely the marki n gs of the scales an d to make the hair

shaft i n e ffect a glassy cyli n der thro u gh which the med ullary
, , ,

cells co u ld be see n with great disti n ct n ess The hair was fi rst .

washed i n the ether alcohol sol u tio n as before a n d the n tra n s

-
,

ferred to a bath of oil where it was allowed to remai n for several

,

mi nu tes It was the n mo u n ted i n the same oil for microscopical

.

examin atio n In the case of the larger hairs it was ofte n n ec es

.
,
 HAI R STR U C TU R E OF THE MO N O T R E MATA 47 1

sary to tra n sfer the hair from the ether alcohol sol utio n to 9 5
“
—

per ce n t al c oh ol th en to a sol utio n composed of equ al parts of

,

9 5 per ce n t alcohol a n d the mo u n ti n g oil a n d the n to the mo u n t ,

i ng oil itself In each of these sol u tio n s the hair was allowed to
.

remai n for several mi nu tes before tran sferri n g it to the mo u nti n g

oil The latter was ofte n heated to 1 00 C or thereabo u ts to
.
°
.
,

i n sure its pe n etratio n i n to all of the tra n sverse ridges of the c u tie
u lar scales .

Third method cleari n g i n balsam O fte n the fin er hairs were

—
.

cleared an d mo u n ted i n balsam After havi n g bee n washed i n the .

ether alcohol solutio n they were dried an d immersed i n a bath

—
,

of xyle n e a n d the n tra n sferred directly to a mo un t of very thi n

balsam With hairs as those of the bats shrews an d man y of
.
, ,

the rode n ts this treatme n t proved to be the best

,
.

3 S ecti oni ng a n d mou n ti ng

. Tra n sverse an d lo n gitu di n al sec
.

tio n s of the larger spi n es s u ch as those of the spi n y an t eater ,

-

( Ta chyglossu s hystri x ) (h g s 52 to were sec ured by faste n i n g

.

the spi n e betwee n two pieces of very hard fi rm grai n ed cork i n ,

—

a n immovable fashio n a n d the n h li n g the spi n e to the desired

,

thi n n ess with a smooth parallel grooved fil e The sectio n was ,

—
.

the n removed from the cork an d gro u n d ge n tly u po n a ho n e

moisten ed with water to re n der the su rface perfectly smooth ,

a fter which it was dehydrated i n several alcohols impreg n ated ,

with xyle n e an d xyle n e balsam an d mo u n ted i n balsam i n the

-
,

u s u al way Those possessi n g the pithy type of medu lla were

.

differe n tially stain ed with eosi n the medu lla taki n g the color
'

, ,

the firm er cortex an d c u ticle n o t .

For hairs i n ge n eral the u su al processes of dehydrati n g i m ,

p reg n at i n g with x yle n e p ar affin xyle n e i n fi l t rat i o n with paraf

—
, ,

fi n mo u n ti n g o n blocks sectio n i n g stai n i n g a n d mo u n ti n g were

, , , ,

employed It was fou n d advisable to u se xyle n e an d p araffi n

.

x ylen e baths hot an d to keep them so i n the ove n allowi n g the

, ,

hairs to remai n therei n for several days each In stai ni n g with .

eosi n safran i n ge n tian violet or methyl gree n the medu lla always
, , , ,

took the stai n the cortex an d c u ticle remai n i n g u n colored

,
.

With most hairs an d more partic u larly with the fi n er o n es a

, ,

very hard p ar affi n gave the best resu lts Whe n it occ u rred t h at .

TH E AM E R I C AN J O U RNA L OF ANA O M Y V O L
T . 27 , NO . 4
47 2 L E O N AUGUSTUS HAUS MAN

the hairs p ulled from ou t the block d uri n g the sectio n i n g a lo n ger ,

period i n the hot xyle n e an d z yl en e p araffin baths was fo u n d to

-

be the re m edy .

The au thor here wishes gratefully to ack n owledge his i ndebted

n ess to the M u se u m of V ertebrate Zoology of the Un iversity of

Californ ia a n d to the Peabody M u seu m of Yale Un iversity for

ha i r samples se n t him an d to D r G S M iller of the Un ited
,
. . .
,

States N atio n al Mu seu m at Washi n gto n an d D r J A A lle n ,

. . .
,

of the A merican M u seu m of N atural History i n N ew York City ,

both for se n di n g ha i r samples an d aids of o n e ki n d an d an other

a t vario u s times a n d for allowi n g the au thor the free u se of their
,

ex te n sive collectio n s of ski n s D r A H Wright of the D epart

. . . .
,

men t of Zoology at Corn ell Un iversity le n t his ki n dly aid i n the ,

determi n atio n of some of the ver n ac ular n ames of species m en

t i on ed i n this paper .

E special tha n ks are du e to Professor H D R eed an d to P r o f es . .

sor Simo n He n ry G age for g uidan ce an d co u n sel d uri n g the

cou rse of the i nvestigatio n .

TH E HAIR OF O RNI T H O RH Y N C HUS A N AT I N U S

The hai r types f O rni thorhyn chu s ( p la tes

o 1 a nd 2)

The fact that O rni thorhyn chu s possesses two disti n ct types of
hair was made k n ow n i n 1 8 02 whe n Bl ume nbach ( 1 80 2 ) described
,

its gross appearan ce L ater Home ( 1 8 0 2 ) recorded the same

.

facts A ppare ntly i n depe n de n tly G lock n er

.
,
v a n de Hoe

V en a n d Pero n a n d L esseu r made a more careful st u dy

of some of the stru ct ural featu res i n a ge n eral way A s ummary
,
.

of the observatio n s up to the year 1 8 2 5 was made by M eckel

The relatio n s of the hairs to each other withi n the follicle
w as the s u bj ect of the i n vestigatio n s of L eydig while the
s ize an d ge n eral gross appeara n ce of the hairs occ u pied the r e

searches of Welcker
The fir st systematic attempt to review the histology of the hair
is that of Waldeyer w h o w as followed i n this same e n deavor
b y Po u lto n The latter first clearly i n dicated the relatio n
47 4 LE O N AUGUSTUS HAUS MAN

Coveri n g t h e ve n ter is a type of the shield hair somewhat si m i

lar to that which clothes the head b ut beari n g a m u ch lo n ger ,

shield an d as might be ex pected co n tai n i n g less pi g m ent ( fi g

, , , .

1 D)
,
.U po n the sides of the ve n ter ( the regio n of the tra n sitio n
i n color from the dark brow n of the back to the creamy white of
t h e abdome n ) the pigme n t is c o n fi n ed to the proximal fo u rth of
the shield In all cases the shaft of this ve nter hair co n tai n s so
.

few of the brow n pigme n t gra nu les as to show n o color except

u n der the microscope .

From the middle of the back to the base of the tail the shield
u n dergoes a progressive i n crease i n le n gth a n d rigidity a n d the ,

shaft becomes correspo n di n gly shorter (fi g Wh e n the tail .

is reached each hair co n sists almost wholl y of the flatte n ed sti t ,

te n ed shield eleme n t a co n str u ctio n which gives it the character

,

of very fi n e bristles N ear the base of the tail the top of the
.

shield is weakly pigme n ted while n ear the en d of the tail the ,

shields of the hairs exhibit an i n creased nu mber of pigme n t

granules .

The tail hairs are similar to those of the feet an d are su bj ect ,

to mu ch attritio n They are co n sequ e n tly co n siderably wor n

.

a n d frayed an d are held closely agai n st the ski n

,
The fore .

feet are almost e n tirely shor n of their hair as far as the wrist ,

si n ce it is upo n these feet that the bru n t of the labor of swimmi n g

a n d of b u rrowi n g falls U po n the hi n d feet more hair is prese n t
. .

O n both feet the hair is of a light brow n color Un like the shields .

of the hair upo n the ve n ter the shields of the hair of the feet are ,

pigmen ted with their characteristic color thro u gho u t their

le n gths This su ggests that the lighter color of the hair upo n
.

the fore feet is n o t du e to the fact that the tips of the shields have
bee n wor n away .

In all these varieties of the shield hair the three eleme n ts ,

shield isthm u s an d shaft are i n variably prese n t ( fig 1 4 B C

, , ,
.
, , ,

D ) tho u gh differe n t eleme n ts rise i n to promi n e n ce a n d determi n e

,

t h e gross appeara n ce of the hair u po n di ffere n t parts of the body

( figs 3 to
. In all of these varie ties likewise the relatio n ships , ,

betwee n the cortex med ulla an d c uticle are f un dam en t al l the

, ,

same b ut i n certai n of the hairs as will be show n the med u lla

, , ,

i s lacki n g altogether .
 HAIR STR U C TU R E OF THE MO N O T R E M ATA 47 5

There exists a variety of the shield hair w hich di ffers so

markedly i n its appeara n ce from all the other types as to su ggest
at fi r st the n ecessity of accordi n g it a separate c la ssi fi c at i o n This .

aberra n t type is restricted to a meager tract abo u t the base of the

s n o u t m ore partic u larly to a n area j u st ve n trad of the base a
, ,

regio n w hich may be termed the chi n ( fi g 1 A ) Fig u re 8 shows .

,
.

the gross appearan ce of this hair an d fi g ur es 3 2 to 3 5 its stru ct u re

,
.

A ll disti n ctio n s bet w ee n sh ield isthmu s a n d shaft have di sap

, ,

p ear ed b u t the c o n fig u rat i o n of the c u tic u lar scales their mode

, ,

of imbricatio n a n d the character of those dissipated medullary

,

cells which still remai n are the same as those of all the other
types of shield hairs The fact that these u ltimate stru ct u ral ele
.

me n ts of the hair preserve their i ntegrity of appearan ce eve n

tho u gh the ge n eral aspect of the hair as a whole may be radically
m o difi ed wo u ld seem to s u bsta n tiate the ass u mptio n that i n
, ,

O r ni thor hyn c hu s at least the c u tic u lar scales an d the med u llary
,

cells o ffer the readiest mea n s of i den t i fi c ati on an d the most

tr u stworthy criterio n for the cl assi fic at i o n of the variou s types of
hair .

The fin er hair fo u n d n ex t the ski n an d u n der n eath the shield

,

hair an d which will be termed the fur hair is of a grayish

,
‘
,
’

brow n color a t ri fl e darker o n the dorsu m tha n o n the ve nter

,
.

It covers the e n tire body with the exceptio n of the tail an d feet
, ,

a n d is everywhere approximately from o n e third to o n e half the — -

le n gth of the shield hair (fig V estiges of this fu r hair may

.

sometimes be fo un d upo n the tail i n adu lts an d a co n siderable

amo u n t is presen t o n the tail of yo u n g i n divid u als This fur .

h air soo n disappears as a rule early i n adu lt life over su ch por

, ,

tio n s of the body as the tail an d more distal portio n s of the

limbs where its prese n ce is u n n ecessary for warmth an d p ro t ec
,

tio n from water L i ke the sh i eld hair the fur hair occ urs upo n
.
,

o n e restricted portio n of the body i n a m u ch m o difi ed form ,

n amely u po n the area immediately s u rro u n di n g the pi n n ae of

,

the ears Here it di ffers from the typical form i n both le n gth
.

a n d diameter tho u gh its characteristic histological str u ct u res are

,

u n cha n ged It is as has bee n said the softest an d fi n est hair

.
, ,

which the creatu re possesses .

47 6 L E O N AUGUSTUS HAUS MAN

The med ulla of the fu r hair is somewhat similar to that of the

shield hair ( fi g its pri n cipal di ffere n ces lyi n g i n the co nfor
.

matio n of the i n divid u al cells a n d i n their closer compactio n

i n the shaft The shield hair h owever possesses a stru ct urally
.
, ,

disti n ctly differe n t med ulla i n the regio n of the shield ( fig s 2 3 .

,

24, an d

The hairless areas of the body are the ve n tral s u rface of the
tail an d the distal po rtio n s of the fe et The denu ded state of .

t h e former is acqu ired by the weari n g away of the few hairs

w hich make their appeara n ce i n the yo u n g Stu mps of these .

early hairs remai n imbedded i n the follicles thro u ghou t the life
of the a n imal an d appare n tly co n ti nu e to grow bu t are c o n ,

st an t l y wor n away A similar co n ditio n obtai n s o n the distal

.

po r tio n s of the feet .

The cu ti cu lar sca les of the s hi eld hai r A t the base of the shield
.

hair shaft the exposed portio n of the c u tic u lar scales is tria n gu lar
i n form with the ac u te apex directed distad ( fi g
,
A s the .

middle of the shaft is approached the scales become more obtu se

, ,

u n til their edges form a series of fi n e tra n sverse ro u ghly parallel

, ,

ridges exte n di n g obliqu ely across the shaft (fig N o differ .

e n ce i n the thick n ess of the c u ticle of the ex t al an d e n tal s u rfaces

of the cyli n drical shaft are appare n t ( fi g .

The scales of the isthmu s ( fig 1 4 C) are like those of the shaft
.
, ,

i n o u tli n e b u t are smaller a n d more compactly gro u ped

,
The .

tra n sverse marki n gs i ndicati n g their edges are i n co n sequ e n ce , ,

more crowded Du e to the a n gle at which the isthm u s is be n t

.
,

the scales i n the co n vex regio n of the be n d are m u ch more closely

imp acted tha n those above an d ofte n prese nt the appeara n ce o f '

havi n g bee n fu sed i nto a solid plate This is du e to the fact that
.

the edges are so closely cro w ded together that they are n o t dis
t i n g u i sh ab l e except u n der very carefu l exami n atio n Upo n the .

o u ter or co n vex side of the be n d where the wear occ u rs the

, , ,

scale edges are m u ch m o d i fi ed i n o u tli n e a n d prese n t a m u ch

more eve n ly parallel appeara n ce tha n a ny of the others From .

t hi s a n d from similar observatio n s o n other hairs i n vario u s sit u

a t i o n s u po n the bodies both of O r n i thor hyn c hu s a n d of other

species of ma m mals it is co n cl u ded that the cha n ge i n form b e

,
47 8 L E O N AUGUSTUS HAUS MAN

shaft of the typical shield hair rather tha n the shield It may .

be that these hairs represe n t a n atomically a n expa n ded a n d flat

, ,

te n ed portio n of t h e sh aft eleme n t rather tha n a n added shield

eleme n t The n ame shield hairs however i n this case is n o t a
.
, ,

m is n omer whe n take n to refer to the form of the hair a n d n o t to

its derivatio n .

The cor tex of the s hi eld hai r The cortical eleme n t is that to
.

which is primarily du e the variatio n s i n form of the shield hair .

In its si m plest co n ditio n i n the shaft of the hair it co n sists of

.

elo n gated distorted fu sed nu cleated cells which are compacted

, , , ,

lo n gitu di n ally as shown i n fig u re 22 In the shield the cortex.

expa n ds an d c omprises the maj or portio n of that str u ct u re ( fig s .

1 7 an d It is this eleme n t of the hair str u ctu re also which

bears the pigme n t to which the color of the hair is du e In all .

bu t the cortex of the shield this pigme n t is distrib u ted u n iformly ,

b u t here it occ u rs most ab u n da n tly i n the form of masses of

gra nu les i n the ect al half of the shield ( fig .

'

The relatio n of the co r tex to the med ulla an d c u ticle i n the

shield hairs of the feet an d tail is show n i n fig ures 2 5 to 2 9 .

The cu ti cu lar sc a les of the f ur ha i r The c utic ular scales of

.

the fur hair also differ i n form from the base to the tip of the
hair N ear the b a se the sc ales are very elo n gate ( hg s 38 an d
. .

3 9 ) a n d their free distal edges proj ect away from the shaft giv ,

i ng to it a disti n ctly serrate p r o fil e ( fig .

N ear the middle of the hair the wor n edges of the scales give
the hair the appeara n ce show n i n fig ure 4 2 a n d n ear the tip that ,

ill u strated i n fig ure 3 7 .

The medu l la of the f ur ha i r N o marked di ffere n ce betwee n

.

the medu lla of this type of hair a n d that of the shaft of the shield
hair exists T hro ugho u t fo u r fift hs of the le n gth of the shaft the
.
-

med u lla persists ( fi g . an d n ear the tip where it does grad u ,

ally thi n o u t a n d disappear there is n o place w here it show s a ny

,

m o di fi c at i o n i n form s u ch as that w hich is n oted i n the c a se of

the shield of the shield hair of the dors um .

The cor tex of the f ur ha i r This eleme n t prese n ts n o di ffere n ces

.

fr om its homologo u s st ru ct u re i n t he shaft of the shield h air .

 H AI R STR U C TU R E OF TH E MO N O T R E M ATA 47 9

S u mmar y f or O r n i thor hyn c hu s

O rn i thor hyn c hu s possesses six well d efi n ed areas u po n
an a ti n u s -

th e body each of which is characterized by the growth of a dif

,

f er en t ki n d of hair These differe n t ki n ds or varieties have

.
, , ,

appare n tly j u dgi n g from the form of the c u tic ular scales an d
,

medu lla bee n derived from two str u ct u rally disti n ct types of
,

hair the fur hair an d the shield hair

,
.

The shield hair becau se of its str u ct u re acts as a protectio n

, ,

agai n st physical i n j u ry to the ski n a n d precl u des the e n tra n ce of

water while the fu r hair serves as an i n s u lati n g mediu m g u ard
, ,

i n g the body agai n st cha n ges i n temperat u re .

THE HAIR AND SPI NES OF TA C H Y G L O SSUS ( E C H IDNA ) H Y S TRI X

The ha i r a n d sp i n e typ es of Ta chyg loss us ( p la tes 3 a nd 4)
The systematic acco u n t of the body coveri ng of Tachy
fi r st
g loss a s w as give n by M ei j er e i n 1 8 94 a n d i n 1 8 9 8 a n acco u n t ,

virt u ally similar b u t with especial co n sideratio n to the disposal

,

of the gro ups of hairs an d spi n es was p u blished by R fi m er D ur , .

i n g 1 8 9 8 also Spe n cer a n d Sweet had iss u ed their work o n the

,

develo pme n t of the hairs a n d spi n es i n the mo n otremes a n d mar

su p i al s i n which a carefu l st u dy is made partic u larly of the de
,

v el o p m en t of the n ew hairs a n d spi n es an d of the vario u s f o lli cu

lar layers taki n g part i n their growth .

Toldt i n a disc u ssio n of the ge n era w hich he mai n tai n ed

as P roechi dna an d Tachyglossu s w as the first to s u ggest that ,

the criterio n of hair str u ct u re an d form is at least a s far as these ,

ge n era are co n cer n ed si g n i fic an t from the stan dpoi n t of their

,

phyloge n etic stu dy a n d may be u sed as aids i n determi n i n g the

,

positio n of the species In 1 9 0 6 he described the hair an d spi n e

.

coveri n g of a species allied closely to the o n e n o w u n der co n sidera

tio n Z ag lossus br uij j ni i bruij j n i i
,
.

A fu rther co n trib u tio n to the k n owledge of the growth a n d

developme n t of the hairs an d spi n es with especial emphasis u po n ,

their gro upi n g i n the ski n was made by Pi n ku s i nzl 906 ,

.

The body coveri n g of Ta c hyglossu s has bee n commo n ly de

scribed as of two types spi n es a n d hairs This is however a,
.
, ,
48 0 L E O N AUGUSTUS H AU S MAN

very ge n eral classi fi catio n a n d m ay be u sed o nly i n a ro u gh way

to i n dicate the ge n eral n at u re of the coveri ng; of the dors u m
a n d ve n ter respectively
,
The coveri n g of the latter co n sists of
.

lo n g sti ff hairs while the former is beset with lo n g sharp rigid

, , , ,

S pi n es so thickly placed as to cover e ffectively both the ski n

a n d their o w n bases .

A closer exami n atio n of these spi n es a n d hairs reveals the fact

that n o defin i t e li n e of demarcatio n c a n be draw n be t wee n them .

From the lo n gest an d most rob u st of the spi n es of the back to

the fi n est of the hairs which c a n be fo u n d o n the ve n ter there ,

exist all gradatio n al estates M ore will be said later regardi n g .

this tra n sitio n i n form from the hairs to the spi n es It w ill .

fir st be n ecessary however to pass i n review the vario u s parts

, ,

of the body with refere n ce to the type of hair a n d spi n e coveri n g

which each possesses .

The dors u m (fig 43 G) is as has bee n said clothed with a for

.
, , ,

m i d ab l e pa n oply of lo n g rigid ac u mi n ate spi n es closely massed

, , ,

together The largest of these are those which occ u r i n what we

.

shall term the hip t ufts ( fig 43 F ) These spi n es are app ro x i

‘ ’
.
,
.

mately 50 mm i n le n gth a n d abo u t 4 mm i n diameter i n t heir

. .

thickest portio n s They are of a light yellowish white hu e with

.
-

very dark brow n almost black tips the color exte n di n g dow n
, , ,

the spi n e for a dista n ce of abo u t 7 mm i n the case of the lo ngest .

spi n es ( fi g . The smaller dorsal spi n es are like the larger ,

with the exceptio n that their tips are pigme nted for a greater
dista n ce dow n the shaft ofte n exte n di n g to o n e fo u rth its e ntire
,
—

le n gth ( fig . Becau se of the close n ess with which the larger
spi n es are set together the lower three fo u rths of the shafts of
,
-

the smaller spi n es are hidde n Th u s it appears as tho u gh the .

dors u m possesses spi n es of two colors large yellowish spi n es a n d ,

smaller black o n es .

The hair of the dors um i s obsc u red by the spi n es a n d co n sis t s ,

of two ge n eral types : o n e fl at t en e d straig ht an d some w hat s t i ff , ,

a n d spi n o u s ( fi g a n d the other smaller

. also fl at t ened a n d ,

somewhat sti ff bu t i n additio n slightl y wav y ( fi g

, , ,
.

U po n the ve n ter occ u r three types of hair : a spi n y flatte n ed ,

type ( fig . a flatte n ed w avy type ( fi g a n d a still fi n er .

48 2 L E O N AUGUSTUS H AU S MAN

The c u ti cu l ar sc a les o The fi n est of the

f the ha i rs a n d sp i nes .

hairs the wavy hairs from the au ric ular depressio n (fi g 4 3 A
—
.
,

a n d E ) dors u m ( fi g 4 3 G) a n d ve n ter ( fi g 4 3 I bear the larg —

. .
, , , ,

est c u tic ular scales ( fig a n d as w e pass i n o u r exami n atio n

.

from the wavy hair to the thicker sti ffer straighter type ( the , ,

type here called the fl at t en ed spi n y hair ) w e fi n d that the scales ,

decrease i n size ; their edges become more closely set together

The greater the thickn ess of the spi n y flatten ed hair ( an d the
'

c o n se q u en t l y g r eat er spi n osity ) the smaller relatively an d the

, , ,

m ore irreg u lar i n ectal o u tli n e are the c u tic u lar scales .

This alteratio n i n the form of the scales is illu strated by the

three types of the spi n y flatte n ed hair each on e i n the order , ,

give n bei n g thicker an d co n sequ e n tly a trifle more sti ff a n d spi

,

n o u s tha n its predecessor These types are : the spi n y flatten ed

.

hair of the dors u m ( fig the short spi n y fl at t en ed hair of the

.

t o p of the head ( fi g a n d the lo n g spi n y flatte n ed hair of the

.

ve n ter (fi g . A compariso n of the fi g u res referred to will .

make clear the relatio n s of the scales It will be n oted that the .

larger an d sti ffer an d more spi n o u s the hair becomes the smaller , ,

relatively are the c u tic u lar scales an d the more closely are their
,

ectal edges crowded together .

A s the tra n sitio n from the spi n y hairs to the tru e spi n es pro
g r esses so also does the decrease i n the relative sizes a n d i rr eg u
,

l ar i t y of the scales u n til upo n the s u rface of the tr u e spi n es s u ch

, ,

as the largest of those fo u n d u po n the dorsu m the scales are so ,

closely m assed together a s to re n der the traci n g of their i n di

vid u al edges ext remely di ffi c ult h ese c a n be see n o nly n ear
.
r
I

the base of the spi n e ; higher up the attritio n to which so stiff an ,

appe n dage as a rigid immobile spi n e is s u bj ected leavi n g the

, ,

s u rface smoothly polished a n d obliterati n g all traces of the edges

of the scales .

The medu ll a of the hai r s an d s p i n es The med ulla likewise .

shows reg u lar tra n sitio n al forms betwee n the wavy hairs a n d
the spi n es In the fi n e wavy hairs it is to be fo u n d o n ly as
.

streaks of mi nu te isolated cell fragme n ts i n va rio u s portio n s of

,

the hair shaft which as has bee n i n dicated i n the disc ussio n of the
, ,

hairs of the mammals i n ge n eral may de n ote that this type of ,

 HAI R STR U C TU R E OF THE MO N O TR E MATA 48 3

hair has bee n derived from a still fi n er variety which possessed

a complete medu lla These isolated medu llary cell fragme n ts
.

i n the shaft are n ever i n a co n ti n u o u s ba n d Seldo m i n d eed .

, ,

are they discoverable O fte n a close search of the e n tire le n gth

.

of the hair shaft with the highest powers of the microscope is

n ecessa r y before the mi nu te vestiges of the med u lla c a n be de

scribed They are I thi n k prese n t i n some degree i n all w avy

.
, ,

hairs ( fig s 8 5 an d
.

In the spi n y fl at t en ed hairs more of the medu lla appears a l ,

tho u gh it still exists i n most i n sta n ces as streaks of isolated cell

, ,

fragme n ts (fi g . A t the base of the shaft i n this type of ,

hair an d j u st below the mo u th of the follicle there ofte n occ u rs

, ,

a large gro u p of medullary cells which resembles the fu lly formed

med ulla fo u n d thro u gho u t the shaft of the smaller spi n es This .

gro up of cells pi n ches o u t an d disappears j u st before the emer

ge n ce of the hair shaft from the mo u th of t h e follicle (fi g .

Waldeyer observes that i n the ce n ter of the large stro n g spi ny ,

hair there may occasio n ally occ u r l arge n odose pigme n t masses , ,

( as he calls them ) which perhaps may be regarded as the r u di

me n ts of a medu lla That these masses are medu llary cells I
.
,

thi n k there is n ow n o do u bt .

A t the ectal extremity of the spi n y fl att en ed hair of the dor

su m there commo n ly occ u rs for the first time i n the series a
, ,

complete medu lla (fi g s 7 1 a n d 7 2 ) which persists for a short

.

dista n ce dow n the shaft a n d the n fragme n ts i nto mi nu te particles

as the middle of the hair is approached ( fig These fragme n ts .

become progressively smaller an d fewer i n nu mber i n the lower

portio n of the hair (fi g b u t gather together agai n i n to a
.

co n ti nu o u s medulla at its base (fi g .

In the smaller spi n es the medu lla is complete a n d i n the larger ,

it reaches its maximu m expan sio n occ upyi n g an importan t posi

,

tio n i n the str u ct u re of the spi n e (hg s 9 4 95 9 6 a n d .

, , ,

The cor tex of the hai r s a n d sp i n es The cortex i n all the hairs
.

an d spi n es is a homoge n eo u s hyali n e hor n y mass i n which n o

, ,

disti n ct evide n ces of its fu sed co m po n e n t cells co u ld with cer

tai n ty be determi n ed Whe n macerated i n a 2 5 per ce n t aq u e
.

o u s sol u tio n of sodi u m hydroxide it splits u p i n to a fi b ro u s m a ss ,

48 4 L E O N AUGUSTUS H AU S MAN

which may be s u ggestive of the c o n fi g u rat i o n of the f used cells .

Similarly whe n a spi n e is broke n obliqu ely across the irreg u larity
, ,

of the fractu re of the broke n cortex eleme n t may also be i ndica

tive of the origi n al cell form (fi g 9 8 . tho ugh it may be
,

that these cortical cells have lost all origi n ality of co n to u r an d

have completely fu sed i n to a tru ly homoge n eo u s s u bsta n ce A t .

prese n t n o stateme n t c an be made regardi n g their form i n the

vario u s hairs an d spi n es .

While it is n ot withi n the provi n ce of this paper to disc u ss the

developmen t of the hairs an d sp m es from what has bee n observed
,

it may n ot be amiss to call atte n tio n to the s uppositio n that the

differe n t varieties of hairs an d spi n es of Tachyglossu s may have
bee n derived from a si n gle hair type possibly of the wavy v a
,

r i et y by an i n crease i n the thick n ess of the middle portio n of the

,

shaft Their developme n t c an be determi n ed with certai nty

.
,

however o n ly by a c areful stu dy of the stages of their growth

,
'

from the follicle i n the embryo an d yo u n g .

S ummary f or Tachyg lossus

In all esse n tial respects the hairs of O rni thorhynchus an d .

Tachyglossus are similar The c u riou s flatte n ed type of hairs is

.

the characteristic form for this order In both O r ni thor hyn chu s
.

a n d Ta chyglossu s this type comprises abo u t 50 per ce n t of the

body coveri n g In Ta chyglossu s the s u ggestio n is adva n ced

.

that the spi n es are developed from the wavy hairs N ot o nly i n .

str u ctu re bu t also i n developme n t are the hairs of O rni thor hyn
“
,

c hu s an d Ta chyg l ossu s alike Spe n cer a n d Sweet observe S o

.
,

far as esse n tial poi n ts are co n cer n ed the developme n t of the

,

large an d small hairs alike agrees i n both O rni thorhyn chu s an d

Ec hi dn a
The type of hair therefore characteristic of the Mo n otremata
, , ,

is the fl at t en ed type represe n ted by the shield hairs of O r ni tho

,

r hyn chu s a n d by the fl att en ed wavy an d spi n y hairs of Ta chyglos

si t s
. O n the other ha n d the fu r hairs fo un d i n O rn i thor hyn chus
, , ,

b u t n o t prese n t i n Ta chyg l ossus are like the ge n eral type of

,

cyli n drical fu r hairs possessed by the maj ority of the mam

48 6 L E O N AUGUSTUS H AU S MAN

FRI D N H A L
E E T . H . 191 1 Zur T ec hn ik der U n t e rs u c h u n g d es l l a a r kl e i d e s u nd

d er H a ar e d e r S au g e t i e re . Z e i t sc h r fur Mo r ph . u . A n t h ro n ,
Bd 1 4 .
,

S 44 1 . .

191 1 T i e rh a a ra t l as . J en a .

G A RCI A , S . A . 1 8 92 B e i t ra g K e zur enn t n s i des H a a rw ec h se l s b e i men sc h l i c hen

E mb y r o n en u n d N g b eu e o r e n en Mo r ph A rb Bd 1 . .

G L O CK N E R ,
E . F . 18 19 Ub d i H er e a ar e d e s O r n i th o r h yn c hu s . I i s s, S 651 . .

HAG E R ,
H .
,
A ND M EZ , C . 1 899 D as M i k r o sc o p u n d se n e i A n w en du g n . B li er n .

H ES S E UND D O F L EIN 1 914 Ti b er au u n d Ti erle b en . L ipe . u . B l in

er .

H M O E 1 802 A d es c r ip t i on of t he a n at o m y of O r n i th or h yn c h u s
Phil T . ran s p . 67 .

JAPHA ,
A . 19 1 1 D ie H a ar e d er W a l t i ere . Zool . J a h rb .
(A n at Bd 3 2 . .

19 1 1 Ub e er d ie H a a re d er W al e . V b er . G es . d e r N a t u rf o r ,
L ip e .
,

Bd 8 2 S 1 68 .
,
. .

KA Z Z A N DE R J 1 9 10 o c hm a l s z u r i o l o i e d e r Ta l p a e urop aea ( H a a r ap p a ra t
,
. N B g
am u ss ) n at A n z J e n a Bd 3 7 S 4 F . A . .
, ,
.
,
. .

KE I BE L , F 1 8 95 O n t o en i e u n d . h y l o en i e v o n aa r u n d ed er Erg b der g P g H F . .

n at , Bd 5 A
61 9 . . S , .

K I DD ,
W . i n h i t n i l l u t t d b y t h di t i
1 90 1 U se er a ce s ra e e r ec on o f t he h a i r on t he
b di f im l L d n
o es o an a s . on o .

1 90 2 H i l p i t i n w i t h h bi t P a r s o e n c on n e c o a s . ro c . Zool . Soc .
,
L d on .
,

vol . 2, p . 1 45 .

1 903 N o t es o n t he h a i r sl o p e of f ou r t yp i c al m amm a l s ( o t t e r , d o g , ox ,

an d h o rse ) . P ro c . Z ool . Soc .

,
L d on .
,
vol . 1, p . 79 .

1914 T he di r ec t o n o f i ha i r in an im l a s an d m an . N ew Y o rk .

K U KEN TH A L ,
W . 1 90 9 U n t er su c h u n g en a n W al en . J en . Z e i t sc h r . fu r N at u r

w i ss .
,
Bd 45 S 545 .
,
. .

L AM PE RT M , . 1 9 10 C on t r ib u t i on s a l et u d e d es p il o s d e l h o mm e
’
et d es an i

m au x P i . ar s .

L A NG L EY J N . . 1 90 1 P r ac t c a l i h i st o l o gy .

L A T TE U X , P . 1 887 M anu el d e t ec h n i qu m i e c ro s c o p iq ue . P i ar s .

L YD I G
E Ub 1 8 59 e er die a u s se r en B d kun g
e ec en d e r S a u g e t h i e re . A rc h iv . fur
A t Phy na . s . u . Wi ss . M ed , S 67 7 . .

L O RE N L 1 909 B
Z, . e t r ac h t u n g ue b er d as H aar k l id d e er S a u g et i er e . V b er . d er
Zool -
Bo t . G esel l . zu W i e n .
,
Bd 59 S . . 27 1 .

M E CK E L 1 8 26 O rn i th o r hyn ehi p a r a d ox i esc r t o D ip i A n at o m c a i .

M E IJ E R E 1 8 94 e er d ie Ub
aa re d e r S a u g e t h i e re , H b e so n d e rs ue b e r i h re A n o rd
nun g . M o rp h . J ah rb .
,
Bd 2 1 S 3 1 2 ,
. .

M E R T S CH I N G ,
A . 1 888 B it g e ra e zur Hi st o lo gi e des H a ar es und Il a a rb a l g es .

A rc h iv f ur m ik A t r . S 32 na .
,
Bd 3 1 .
, .

MEYER -
L I RH IM E E ,
F . 1 9 1 0 D i D i h t ig k i t d B h g b im F t d M e c e er e a a run e e us es en

s c h en u n d der A ff Z i t h M p h S t t t g t Bd 1 3 S 1 3 1
en . e sc r or u ar ,
.
,
. .

P E RO N ET LES SU EU R V y g d D o t
a T A t l p d t l
e e ec o uver es au x o rr e u s ra e en an es

an n és 1 8 80 1 88 4 —
.

P I N KU S, F . Ub d H 1 90 6 e er er a a rs c h e i b en der M o n o t r em e n . Z oo l . F o rsc h u n g s
i i
r e s enA t li n us ra en und d e m m a l ay i sc h e n A rc h ip el, R . S e men , 3 .

B d 3 Li f ang (i . . e e ru n n D enk s c h r d e r . Med . N a t u rw i s s G e s e ll . .

, J en a) .
 HAI R STR U C TU R E or THE MO N O TR E MATA 48 7

P OU L T ON 1 8 94 St r u c t u re o f t he bi ll an d ha i rs of O r n i thor hyn c hu s p a r a do x us ,

et c .
Q u a rt J o u r M i c S c i . . . .
, p . 1 43 .

R AW ITZ , B 1 90 6 B i t ag e r e z ur m i k ro s 0 0 p i sc h e n A n at om e i d e r C e t a ce en ( Ub e er

der F i e n e r en Bau d e r H a ar e , u . s . I n t e rn . M o n a t sc h r , . A t na .
,

L ip e .
,
Bd 2 3 S 1 9 .
,
. .

R O ME R ,
F . 1 8 98 S t di u en ue b er d as I nt e gum en t d er S au g e t i er e . II . D as In
g
t e um en t d e r M o n o t r em en . Zool . F o r sc h u n g i s r e sen in A t li u s ra en u n d

d e m m a l a y i s c h en A rch ip el , R . S e mo n ,
3 . B d an ,
2 . Li f g e eru n ( in
D en k sc hr d e r . M ed . N a t u rw i ss . G esel l J en a ) .
,
.

S ARA IN S ,
P . 1912 Ub e S h tz g d
er d ie z oo lo g s i c he c a un e r so g en a n t en H a arm en
s c h en . u . s . w Z l J h b S pp l S 289
. oo . a r . u .
,
. .

S H W A LB
C E, G . 1 90 9 U b d i R i ht g d H
e er e c un er a are b e i S au g et i e r e , s p zie el l

b e im M e n sc h en M u n c h er m ed W o c h en sc h r Bd 56 S 3 1 5 . . .
,
.
,
. .

191 1 U eb er d i e R i c h t u n g d e r H a a re b e i d en A fi en em b ryo n en u s : w ,
. .

1 9 1 2 M i t t e i l u n g en u e b e r d i e H a a r e b e s o n d e r s u e b e r i h r e R i c h t u n g ,
.

Mit t . Phil o m at h . Ges . St ra s s ,

Bd 4 S 51 7 .
,
. .

S N
PE CE R A ND SW EET 1 898 9 9 —
T he s t ru c t u r e a n d d v e el o pm en t o f t he ha i rs o f

m o n o t rem es an d m a rsu pi al s . P ar t I . M o n o t r em es .
Q u art . J ou r .

M ic S c i . .
p 549
,
. .

T LDO T, K . J UN . 1 90 5 e Ub er d as G en u s P r o ec hi d n a . V er . der E . K . Z o ol .

Bo t . G esel l . zu . Wi en , Bd 55, S 5 . . .

1 90 6 Ub e er d as H a ar und St ac he l k l id e on Z ag l oss u s , G il l ( P r oechi dn a

G erv a i s ) . A n n d e s K K N a t u rh i s t H o f m u s , Bd 2 1 , S
. . . . . . 1 .

1 90 6 I n t e ress a n t e H a a r o rm e n f b ei e n em i k u r z sc h n ab e l i g en A m e i sen i
g el o o l A n z , Bd
. Z . . . 3 0 , S 30 5 . .

1 90 7 — 08 t u en u e S di b er d as H aar k l id e v on Vu l p es vu l p es , L . A nn d es .

K K N a t u rh i s t H o f m us
. . . Wi en , Bd . 22 , S . 1 97 .

1 909 B e t r ac h t u n g ub er d as H a ar k l id e d e r S au g et i e re . Wi en . V b er .

Z ool . Bo t . G esel l .
,
Bd 59 S .
, . 27 1 .

1910 Ub e er e n e i B ea c h t e n sw e r t e H aa r s o r t e u n d u e b er d as H a arf o r
m en sy s t em d e r S au g et i e re . Wi en An n . N a t u rh i st . H o f mu s .
,
Bd . 24 ,
S . 1 95 .

1912 B it g
e ra e zur K en n t n s i d e r Bah a a ru n g d e r S a u g et i e r e . Z ool .

J ah rb . J en a
A b t f S y s , Bd 33 , S 9, . . . . . .

1912 W i l t z e i c hn un g in d er En
Ep i d e rmu sst r e if en , H aa re i b en , un d

t w i c kl u n g d e r au s a t e Wi en V erh Z o o l Bo t G e sell , Bd 1 6 H k z . . . . . . . .

19 13 U
e b e r d i e au sse r e Ko rp e r g e s t a l t e i n es et u s v o n El ep has ma x i F
mu s Wi en , D en k sc h r d e r ad Wi ss Med N at K l ass e, Bd 90 , S
. . Ak . . . . . .

259 .

VAN DE H V NOE E 1 8 23 M em o i r e su r le G en r e O r n i thor hyn qu e, N v A o a ct a .

A d C ca . ae s . L e op .
,
vol 1 1 , p 3 51
. .
‘

W A L D EYE R 1 884 Atl as der m en sc h l c h en i und t i i er s c hen H a a r e , s ow e i d er a hn

l i c h en F ase rg eb i l d e . L ah r .

W E BE R M
, . 1 904 D i e S au g e t h i ere . J en a .

TH E AM E R C AN J O U RNA L
I OF ANA O M Y V O L T , . 27 , NO . 4
 PLAT E 1
E X LANA I N
P T O OF F I G UR ES

1 i n d w i g f O th hy h n t n t h w t h l t i
O utl e ra n o f t he rni or nc us a a i us , o s o e oca on o

v i u h i t t A m di fi d h i l d h i f t h h i n ; B f u h i f t h
ar o s a r r ac s , ; o e s e a r o e c ,
r a r o e ear

C ,
f p i gm
a re a o t d t un t d h i ld h i ; D
en e f un p i g m n t d t u n t d
,
r ca e s e a r ,
ar e a o e e ,
r ca e

s h i ld h i ; E t u
e t d hi ld h i
a r ,
f f t nd v nt
r nca e f t il ; F hi ld h i sf e a rs o ee a e er o a ,
s e a r o

d u m ; G l g h i l d h i f d um f t i l T h
o rs ,
ar e s e E nd G l k t h f u a r o o rs o a . e are a s a ac e r

h i a r N n f th
. o h p l y d fi n d (w i t h t h
e o x pt i f th
e a re as a r e s f th ar e e e e ce on o e a re a o e

e ar ) b ut m g g d u l ly i t
. ern th
e X ra a n o o e an o er

2 A g m n t f t h f l l i l f t h h i l d h i n d fu h i n t h d um
rr an e e o e o c es o e s e a r a r a r o e o rs

( f t M ij
a er ) 4 h i l d h i ; B b un d l
e er e . f fu h i r
1
,
X 200
s e a r ,
es o r a . .

3 t 1 1 i l lu t t t h v
o i u t y p f h i f u n d n O th hyn h ( i n t
s ra e e ar o s es o a r o o rn i or c us c r . a .

S iz ) e .

S h i l d h i f m t h d um
O
C e a r ro e o rs .

P S h i l d h i f m m idw y b tw n t h y
I - e a r ro a e ee e e es .

S h i l d h i f m t h p x im l nd f t h t il
H
Q e a r ro e ro a e o e a .

Q S h i l d h i f m t h di t l n d f t h t i l
e a r ro e s a e o e a .

N S h i l d h i f m t h v nt
e a r ro e e er .

Mo d i fi d h i l d h i f m t h h i n
O
O e s e a r ro e c .

9 Fu h i f th d r um nd v nt
a r o e o rs a e er .

1 0 Fu h i f m t h r f th
a r ro e a r ea o e ea r .

1 1 Shi ld h i f m th e f th a r ro e a r ea o e ea r .

1 2 G u id fi g u e re .

13 S hi ld h i t p inte fig u 12 h wi g th
a r a t i ul l
o M m d u ll ; 0 ,
re ,
s o n e cu c a r sc a es .
, e a

CO , c o rte x X 800
. .

14 Sh i ld h i t m fig 1 2
e a r a —
n, u re . B, sh a f t ; C ,
i st h m u s ; D , sh i ld ; M
e ,
me dul l a ;
CO , c or e t x X 800
. .

15 Sh ft f hi ld h i t p fig
a o s e a r a ,
u re 12 . M m e d u ll a ; CO c o r t e x
,
X 800 ,
. .

16 S h ft f h i ld h i b l w q
a o s e a r e o ,
fi g u re 12 B b u l b ; M m e d u l l a ; CO c o r t e x
.
, , ,
.

X 800 .

17 Shi ld
i d h i M m d ul l ; CO
e o f s h el t x ; CU uti l ; M v t ig a rf .
, e a ,
cor e ,
c c e
'
,
es es o

t h m d ul l w h i h f t n p
e e i t n th b
a f th
c h i ld
o X 400 e er s s ea r e a se o e s e . .

1 8 G u id fi g u e re .

19 T n v t i n t h u gh v
ra s i u p ti
e r se sec f th hi ld h w
o by ro ar o s or on s o e s e as s o n

t h g u id fi g u
e fig u 18e L t t in di t w h t h t i n w m d N t
re , re . e e rs ca e e re e sec o s ere a e . o e

th g e t t hi krea e r f th ut i l n th c t l u f
n ess o f th h i ld X 90 e c c e o e ec a s r ac e o e s e . .

20 T n v t i n t h u gh t h
ra s h ft f th
e r s e se c h i l d h i m id w y f m t h
o s ro e s a o e s e a r, a ro e

b f th
as e o h f t t t h i t hmu
e s M m d u l l C U u t i l ; CO
a o t x
e X 2 80 s s .
,
e a, ,
c c e . cor e . .

21 T nv t i n t h ugh t h
ra s h ft f th
e rse se c hi ld h i n th b o f ro e s a o e s e a r e ar e as e o

th h ft j u t b v t h m ut h f t h f l l i l
e s a s a M m d u l l ; C U u t i l ; CO
o e e o o e o c e .
,
e a ,
c c e ,

c o rte x X 280 . .

22 S h ft f t h h i l d h i di
a t d CU
o ti l i t t ; CO
e s e a r, ss ec e .
, cu c e, n ac ,
lon g di st o r t e d
ce ll f th
s o t x xp d ft th m v l f t h ti l
e co r e l
e o se , a er e re o a o e cu c u a r s c a es ; M m e d u ll a r
, y
ce ll xp d f t t h m v l f t h
s e o se t x l m
a t X 2400
er e re o a o e co r e e e en . .
 PLATE 2

E X LANA I N
P T O OF F I G UR S E

23 M e d u ll a of t he sh el i dh i a r i n t he sha ft b l w t h i e o e s t h mu s . M m ed u l l ary
,

c el l i ii
; I , n t e r s t t a l o r i n t e rm e d u l l a r y s a c e 4000 p . X .

d
24 M e u l l a o f t h e s h e l h a r m w a i d
e t w een t h e i id y b i s t h m u s an d t h e t ip of t he
i d
sh e l . M m e d u l l a ry
,
c el l . X 200 .

25 S h i ld h i
e a r of i
t h e t a l , n ea r t h e b a se . X 535 .

26 Shi ld h i
e a r of t he t a il
m wa , id y f rom b a se t o t i p M , me ul l a 45 . d . X .

27 Shi ld h i
e a r of t he f eet m w a ,
id y f r om b a s e t o t i p M , m e u ll a 45 . d . X .

28 S ti th
ec on ro u gh t he s hi ld h i
e a r o f t he t a l , m wa ro m b a s e t o t i pi id y f .

M med ul l a ; C U
, ,
c u t cle ; i CO , c o rt e x X . 73 .

29 S ec t o n i t h ro u h g sh e l i d ha i r of t he f ee t , m idw y f a ro m b as e t o t ip . M ,

me d u ll a; C U , c u t c l e ; CO , c o rt e i 73 x X . .

30 S i g
h o w n t h e re l a t o n s o f t h e s h e l i i d an d fu r ha i r on t he d o r su m o f O r n i tho
r hyn c hu s . A, sh e l i dh i a r; B fur hair ; C , ,
s kin . X
31 Sh w i g th
o n e r el at o n s o f i t h e m o d ifi ed sh e l i d an d fu r h a i r on t he i
c h n area

of O r n i thor hyn c h us . A , m o d i fi ed s h el i dh i a r ; B, f u r h a i r; 0, s ki n . X 5
.

32 G u id e fi g u re .

33 M o d i fi ed sh eli dh i a r o f c h n , at i A figu 32 ,
re . X 200 .

34 M o d i fi ed s hi ld h i e a r of c h in t B fig u e 3 2
,
a ,
r . M m ed ul l a
, . X 2 10 .

35 Mo d i fi ed s hi ld h i e a r of c h in b l w C fig u,
e o ,
re 32 . X 200 .

36 G u id e fi g u re .

37 Fu r h a i r, A — B, fi g u re 3 6 . X 450 .

38 Fu r h a i r, C D , fi g u re 3 6
—
. X 600 .

39 S i gl i
n e, so l a t e d i
c u t c u l a r sc a l e f ro m t he sh a ft of t he f ur h a i r , n e ar b ase .

X 1 2 50 .

40 Fu r h a ba i
M m d u ll ; C U u t i l ; CO
r at the t x X 650 se .
, e a , c c e ,
c or e . .

41 T an sv r t i n t h u gh t h h f t f t h f u h i m id w y b t w
e rs e sec o ro e s a o e r a r, a e ee n t he
b an d t i p M med u l l ; C U u t i c l ; CO
ase . t x
,
X 7 00 a ,
c e ,
cor e . .

42 S h f t o f t h f u h i m id w y b t w n t h b
a e n d t ip M m d ul l
r a r, a e ee e a se a .
, e a; CU ,
c u t i l e ; CO
c t ex X 7 50 , cor . .
HA I R ST R U C T U R E O F T H E MO N O T R E MAT A P LAT E 2
L EO N A U G U U HA U MAN
ST S S
 PLATE 3

E X LANA I N
P T O OF F I G UR ES

43 O u t l i n e d raw i g n of Ta c hyg l os s u s h ys t ; i i ‘

,
to sh o w t he lo c at i on of t he
v i ar o u s S pi n e an d d p i n; B h i t a r r ac t s A , sm al l s i n e s
. p of t he i
au r c u l a r e r ess o

d B u v d pi nd p i y h i f t h fl k ; C d C u v d p in nd
’ ’
an ,
c r e s n es a s n a rs o e an s an ,
c r e s es a

sp i y h i f t h f t ; D u d l p i ; E w vy h i f th u i u l d p i n ;
n a rs o e ee ,
ca a S nes ,
a a r o e a r c ar e ress o

F h ip t f t
,
f p in ; G l g
u p in d h i
s o f th ds um ; H h t p in es d ,
ar e s es an a rs o e o rs ,
s or s e s an

sp i n y h i f t h h d ; I l g p i ny d w v y h i f t h v n t
a rs o e X5 ea ,
on s an a a rs o e e er . .

44 A ng m t f pi rrad h i up n th ed m d id f T h yg l
en o s n e s an a rs o e o rs u an s es o ae os

sus ( af t er R omer ) . X 5 .

S m ll w v y h i f m t h v t
45 a Xa a r ro e en er .

46 L g w vy h i f m th v t
on X a a r ro e en er .

47 L g p i y h i f m th v t
on s X n a r ro e en e r .

48 L g pi y h i f m v ton sX n a r ro en er .

49 C v d pi y h i f m f fl k X
ur e s n a r ro o re an

50 S m l l p i e f m f fl
a k X s n ro o re an .

51 S m l l p i f m a i l d p i
s ne X ro n e ar au r c u ar e re ss o n .

52 L g pi f m d m X 5
N

ar e s ne ro o rs u . .

53 L g p i f m h ip t f t X 5
ar e S ne ro u . .

54 L g t f th p i f m ar es t h m di o li f th d m
e s A pig n es ro ne ar e e an ne o e o r su

m t d p ti
en e ; B p i gm t d p t i ; C i t hm ; D t E p t i w i t h i t h
or on . un en e or on ,
s us o ,
or on n e

f ll i l ( b l b ) X 5
o c e u . .

55 L g t f t h p i f m t h h ip t f t
ar es o f m th l t l e s d l t ft n es ro e u s, or ro e a e ra c au a u s .

A p i gm t d p t i
, ; Ben e p i gm t d p t i ; C i t hm ; D t E p t i
or on ,
un en e or on ,
s us o ,
or on

W i t h i t h f ll i l ( b l b ) X 5
n e o c e u . .

56 S m ll bl k p i f m t h m di li f th d m f m th
‘ ’
a er, ac s ne ro ne ar e e an ne o e o r su ,
or ro e

m di e d l li
an f t h t ilorsa A p i gm t d p ti ne o; B p i gm t d p t i ; e a .
,
en e or on ,
un en e or on

C i t hm,
s ; D t E p ti us w i t h i f ll i l ( b l b )
o X5 ,
or on n o c e u . .

57 S m l l t f t h pi f t h b dy
a es p i gm t d d m
o mi t th
e s nes o e o ,
un en e ,
an ore acu na e an

a ny o f t he o t h e rs , f ro m t he s id es of t he b o dy ,
t op of t h e h ea d ,
au r c u l ar i d ep r es
i
s on , o r p ip h y
er er of ta l i . B , un p i gm en t e dp o rt o n i ; C, i st h m u s ; D to E, p o rt o n i
w th i in t he X5 f o ll i c l e ( b u l b ) . .

58 F l t t d m wh t p i y h i f m
a en e t h m di
,
s li e f th d m a s n a r, ro n e ar e e an ne o e o rs u .

A t p f vi w ; B l t
, o o l vi w e X 15 ,
a er a e . .

59 S l i g h t l y fi w vy h i f m th id f t h b dy w h t h m l l
n er a a r ro n ear e s e o e o e re e s a er

Sp i b g i t pp
n es e A t p vi w; B l t
n o a l vi w ( i t
ea r iz ) .
,
0 e ,
a er a e . c r . na . s e .

60 S l d p i g m t d p i f m l t l d l t f t A p i gm t d p t i ;
en er, en e S ne, ro a era c au a u .
, en e or on

B ,
p i g m t d p t i ; C i t hm ; D t E p t i w i t h i t h f l l i l ( b l b )
un en e or on ,
s us o ,
or on n e o c e u .

X5 .

61 S l d p i gm t d p i f m l t l d l t f t B p igm t d
en er , un en e s ne, ro a e ra c au a u .
,
un en e

p t i ; C i t hm ; D t E p t i w i t h i t h f l l i l ( b l b ) X 5
or on
’
,
s us o ,
or on n e o c e u . .

62 S m ll p i f m t p a id f h d
s f m th
ne i l d p i
ro o or s es o ea ,
or ro e au r c u a r e r es s o n .

B ,
p i gm t d p t i ; C i t h m ; D p t i w i t h i t h f ll i l ( b l b ) X 2
un en e or on ,
s us ,
or on n e o c e u . .

63 64 d 65 F m
, ,
anf th p i y h i f t h h d C i t hm ; D p t i
or s o e s n a rs o e ea ,
s us ,
or on

w i t h i t h f l l i l ( b lb )
n e X 25 o c e u . .

66 W v y h i f mb a th th pi a rs d pi y h i d f m th
ro i l en e a e S n e s an s n a r s an ro e au r c u a r

d p i
e ress o nD p ti w i t h i t h f ll i l ( b l b )
.
, ( i
or t iz ) on n e o c e u . c r . na . s e .

67 L g p i y h i t h l g t t h b dy f m t h v t
on s n A t p Vi w ;
a rs , e on es on e o ,
ro e en e r .
,
o e

B l t ,
l v i w ; C i t hm
a e ra ; D p e ti w i t hi t h f l l i l ( b l b )
,
s ( i t iz )us ,
or on n e o c e u . c r . na . s e .

68 S h t p i y m wh t w vy h i
or s f m th v t
n ,
A t p vi w ; B
so e a a a rs , ro e en er .
,
o e ,

l t a era l v i w ; C i t hm ; D p
e ti w ithi t h f ll i l (b lb )
,
s X 15us ,
or on n e o c e u .
,

69 L g w vy h i th l g t th b dy f m v t
on a Xa rs , e on es on e o ro en er .

70 G u id e fi g u re .

71 T ip of fl a t t e n ed s pi y h i f n a r ro m n e a r t he m e di an l i ne of t he d o r su m , f ro m
A — B, fi g u r e 7 0 X 170 .
 PLATE 4

E X LANA I N P T O OF F I G UR ES

74 S i
ec t o n of t he me h a
sa i b r, D E , fi g u re 7 0
e t w ee n M , m e ul l a 1 20 —
. d . X .

75 S ti
ec on of t he s ame h a i b r, e t w ee n E— F , fi g u re 7 0 M , m e ull a 1 20 . d X .

76 C u t i ul c a r s c a l es o f t he s am e h a r , m i
w a b e t w ee n t h e b a se an d t i p id y . X
1 75 .

77 G u id e fi g u re .

78 S ec t o n of i t he s p i y fl tt
n a en e d ha i r of t he t op of t he hea d b ,
e t w e en B C -
,

fi g u re 7 7 . M m e d ull a
, . X 20 .

79 S tiec on of t he s am e ha i b r, e t w en C— D , fi g u re 7 7 . M m e d u ll a
,
. X 22 .

80 B lbu of t he sam e ha i r, at F , fig u re 7 7 . M m ed ul l a ; S l ev el
, ,
of t he ep i

de mi r s ( m o u t h o f t he f ll i
o c le ) . X 32 .

81 C u t i c u l ar sc al e s o f t h e s am e h a i r a t p i o nt D , fi g u re 7 7 . X 20 .

82 S ec t o n o f i t he s am e ha i
D , fi g u rs 7 7 ,
r, n e a r sh ow n i g t he i sol ate dg ro u p of

a dv enti t i u m d u ll y
o s e ar c el l s so m e t m e s r e sen t i n i p di ff er e n t p o rt o n s o f i the sh a ft .

M me ,
d u l l y el l X 27
ar c s . .

83 G u id fi g u e re .

84 T ip f t h w v y h i f
o e a a r ro m t he i
au r c u l a r d p e ress o n , a t i t he p i gm en t e dp or

t i on , A — B, fi g u r e 8 3 M , me . d ul l a . X 500 .

85 S i
ec t o n o f t h e s a m e h a i b r, e t w ee n B C, fi g u r e 8 3
—
’
. M m e d u ll a ; C U
, ,
c ut cl e ; i
CO , c o rt e x X . 450 .

86 S ec t o n o f i t he sam e ha i r, at D , fi g u re 8 3 . M m e d u ll a ; C U
, ,
c ut c l e ; i CO ,
c o rt e x X
. 450 .

87 C u t i c u l a r s c al es o f t he s am e ha i r, m id w y b a e t w een C D , fi g u re 8 3
-
. X 500 .

Sh ft f th
88 a o e fl at t en e d s pi y h n i
a r of t he v en t e r, m idw y b a e t w e en t he b a se

an d t ip X 18 . .

89 S h ft f t h a o e sam e ha i r, n ea r t he b a se . M m e d ull a
,
. X 18 .

90 S h f t f t h a o e lon g w vy h i a a r of t he v en t e r , m id w y b a e t w een t he b ase a n d

t ip X 40. .

9 1 Sh ft f th a o e s am e ha i r n ear t he b a se M .
,
me d ul l a . X 40 .

92 T v ran s e rse s ec t o n o f i t he s pi y n fl at t ened h a i f

r o t he v t en e r , m idw y a be
t w ee n t h e b a se a n d t ip . CU, c ut c l e i ; CO , c o r t e x X . 20 .

93 T v r an s erse sec t o n o f i t he s am e h a r n ear t h e i b ase . CU , cut cle ; i CO , cor

t ex . X 20 .

94 M e di an lon g i t di u n a l sec t o n i t h ro u h l a r g g pi e S ne of t he d o r su m m ,
id w y a

b et w ee n t he b a se an d t ip . M m e d u ll a ; C U
, ,
c u t c le ; i CO , c o rt e x X . 4 .

95 M e di an l o n g i t u di n al s ec t o n i t h ro u h t h e t i p g of t he sa m pi
e s ne . M m ed ,

ul la ; C U, cut cle ; i CO , x X4
co rte . .

96 T r an s v e r s e s ec ti th
ongh m p i rou sa e S n e, m idw y b a e t w ee n t he b a se a n d t ip .

M m e d u ll a ; C U
, ,
cu t i l ; CO
c e t x X4 ,
c or e . .

97 Sh wi g th
o n i
e c u t c u l a r sc a l es n ea r t he b as e o f t he sa me s pi ne . B, p ort o n i
w th i i n t h f ll i e o c le ( b ul b ) . X 4 .

98 St e re o g ra m of p o rt o n o fi d o r sal s pi n e , se c t on e i d ,
an d i
w t h the e c t al e n d

b ok r en . T h e m et ho d of f rac t u r n i gm ay be i di
n c at iv e of t he f o rm o f t he h o r n y ,

c o a l es c e d fu if
,
s o rm c o r t i c u l a r c e l l s wh i ch com p o se t he c o rt e x of t he s pi ne . M ,

me d ul l a; CU , i
c ut c l e ; CO , c o rt e x ; B C0 b k .
’
,
ro en c o rt e x X 4 .
H AI R ST RU C T U R E o r TH E ATA
MO N O T R E M P L AT E 4
L EON A U G U U HA U
ST S S MAN

C O CU
498 IN D E X
N FE R IO R 1 2-m m p ig em by ELSO N \ I T A N D D W M BA UM
G A R NE R E A D v l p m t f t h
i n a r o , a sso oc x
l d d i g p , .
.

e at e w i t h t he ra n a e o f t h e o rt a l T e e o en o e
y di y i gl d
.

s s t e m m t o t he c ar na l s s te m Ah t m
mb
u er ne an s in an
v v t l d v l p
.

s e nc e o f t h e e na c a a 3 95 Nu f O
er o th p t o va n os n a a
p
e e e o
I n fla m m a t i o n ) i
R e ac t o n s o f c e ll s i n t he t a l i m t f th v y ( l bi
en t) w t ho e o ar a n o ra
ph ibi
1 es e
v d f
,

of A m a n l a r ae t o i n j ec t e c ro t o n e i a l re e r e n c e t o t he
0 1 1 ( ase t c pi
I n t e r c a la t e d l S CS dS t u es o n s t r pe m us c l e di i d G A W A C H IK A N O S U K E C o n t n b ut l o n t o
s t r uc t u r e VI T h e c o m p ara t w e b i s t o l t h e h x s t o l o g y o f t he res p l ra t o ry s a ce s o f p
o g y o f t h e l eg a n d w m g m u sc le o f t h e as , w p t h e e rt e r a t e l u n g s v b
p i f
w 1 t h s ec a l re ere n c e t o t h e e no m e n o n ph O GA A
W C H IK A N O S U K E T h e fi n e r ra m l fi c a
of str ip v
e r e e rs a l d i g c o n t ra c t o n a n d i t ro n s o f t h e
,

u m a n l un g h
b w
ur n
g
t o t h e e ne t c r el a t o n i
et ee n c o n t ra c i 0 1 1 ( a se pt l c m fi a m m a t x o n ) R eac t l o n s o f
b d v
.

t1on a n s and c e l ls m t h e t e n] o f A m p h i b l a n l a r ae t o
In t race ll u l a r h e m o c y t og e m c a c t 1 v 1 t y o n t h e m j e c t e d c ro t o n
p a rt o f t h e gi
a n t - c e lls , a n d t o t h e s 1 g n 1 fi O va O n t he os t n a t a l e el o m e n t o f t h e p d v p
d a f
.

c a n c e o f t h e s o- c a ll e m 1 t o t 1 c fi g u r es i n o v r y ( a lb m o ra t ) w 1 t h es p ec x a l re ere n ce
h
t es e c e l l s u rt er st u F
e s o n re d h di b t o t he n um e r o f b
m ar ro w
I E x e r me nt a l II p i
o ne
to Cy a
O v r y ( a l b m o ra t ) w 1 t h e s ec x al r e e r e nc e t o f
gi p i f d b p
. ,

l o c , w i t h s ec a l re e r e n c e t o t h e a t a t he n u m er o f o v a n t he os t n a t a l
su gg i g
est n . d v
e el o men t o f t h e p
ORD AN H E F urt h er s t u d 1es o n re d
AFhEZ J AM ES W H ear t m u sc u l at u re
b n
.

w of
.
, .
,
o e -
m a rro E x p e n m e n t al I II t e atrxa
Cyt gi f
.

olo c . w 1 t h s p ec ml re e r e n c e t o t h e
d t
a a su es t n gg i g
1 n t r a c el l u l a r h e m o c y t o P1g em by ma t e d w 1 t h t h e r a i n a d g
g i
e n c ac t t ivi y
o n t he a rt o f t h e a nt p gi t he
r
p
o , asso
o rt al s y s t e m m t o t h e c ar d m a l
e of

c e ll s ,
so—c a l l e
a nd t o t h e s 1g n 1 fi ea n ee o f t h e
d i i
m t o t c fi g u re s i n t ese c e lls h t em
a A b . sen c e o f t he v en a c a v a m f er x o r
sys

J O R D AN H E di pd
S t u es o n s t r1 e m u sc l e
in 2 l
y
P o r t a s st e m m t o t h e ca r d m a ] s st e m
l Ah y
aa a
, .

V I T h e c o m p ara t w e h i s t o l v
.

s t r uc t u re s e n ce o f t h e 1 n f er1 0 r m l 2- m m
Wi g w en a c v
.

by
,

o g y o f t h e l eg a n d n m us c l e o f t h e
p l g e m r o , assoc mt e d w 1 t h t h e d ra m ag e
as p .
p i f ph
w i t h s e c a l re ere n c e t o t h e e n o me no n
f h
of strip v d g
e r e er sa l c o n t r ac t 1o n a n d
o t e
P u l m o n a r y e o l u t i o n m t h e m a m m ah a v
b w
uu n
A
t o t heg i e n e t c re l a t 1 o n et ee n c o n t r ae
i b d
c r1 t 1 q u e o f t h e t h eo r l es o f
t on i
a n s a nd dd
n t e r c al a t e 1 sc s

I D N E Y t b ul A t u dy w h p
AT ) , i t es ec i a l re ere n c e t o t he n u m er f b
f th u e s o e 11 p 0 1 d
of o va O u t h e o s t n a t al e el o m e n t p d v p
( o n t e nt of t he
o f t h e o v ry ( al b m o
.

a
A R\ i
A E t o n j e c t e c ro t o n 0 1 1 ( as e t c l n d pi R eac t l o n s o f cell s i n t h e t a l o f A m p h l b x a n l a r l
fl am ma t l o n) R ea c t o n s o f c e l l s i n t he i v ae t o m j ec t ed c r o t o n 0 1 1 ( ase p t l c l n fi a m
i
t a l o f A m p h 1 b 1a n mat l o n)
L eg a n d w m g m us c l e o f t h e as , t s ec a l w p Wi h p i R ed o n e- m a r r o b I E x p e n me n t a l II w
f ph ip f
.

re e re n c e t o t he e n o m e n o n o f st r e re C y t o l og l c , w 1 t h s p e mal re ere n c e t o t h e
ve rs a l uu n d g
c o n t rac t o n a n d t o t h e g e i d
a t a s ug g es t m g m t ra cell ul ar h e m oc y t o

i
n e t c re l at 1 o n et b w
ee n c o n t ra c t o n an s i b d g
em e a c t 1 v 1 t y o n t h e art o f t h e g l a n t -cell s , p
a n d 1 n t e rc a la t e d S t u es o n s t r1 ed di pd a n d t o t h e s x g m fi ca n ce o f t h e so -c all e d
h h
1 sc s
m usc l e s t r uc t ure V I T h e c o m a ra t e p iv m 1 t o t x c fig u res m t es e c el l s u rt e r F
hi gy
.

st o l o o f t he 1 s t u d 1 es
L p id
i o c o n t e n t o f t he k ne t u ule A st u id y b dy R es p l rat ory s ac es o f t h e e rt e rat e l u n g s p v b
o f t he 69 C o n t rl b ut l o ns t o t h e h l st o l og y o f t h e
L g
un T h e fi n e i ra m i c a t o ns o f t he u m a n 3 1 5 if i h
L ung s C o n t n b u t l o n s t o t he s t o lo o f t he hi gy MI T H CH RI ST A N N A I A st u dy of t he
p v b
,

b
.

res p l ra t o ry s a c es o f t h e ert e ra t e h p o nd c o n t en t o f t he k x d n ey t u ule

AMM A h S p ac e s of
t he v e rt eb x a t e l u n s C o n t rl b u g
f p m
I L A A
y
c ri t ue o f t he t eo n e s
t l o n s t o t h e h l s t o l o g y o f t h e r es ra t o r pi y
Mar ro w
o ul o nar

E x e r m e nt al
I p i
e v o l ut 1 o n l n

II C t o l o i c
t he
y g S t rl p e d m us c le s t r uc t u re V I T he comp r
t l v e h l s t o l o g y o f t h e l e g a n d w m g m u sc l e
333
. aa
f d gg
,

w i t h s p e ma l re er e n c e t o t h e a t a s u
i
es t
ivi y o f t h e w asp , w 1 t h s p e c x a l re ere nc e t o t h e f
i ng
o n t he
n t rac e ll u la r h e m o c y t o g e n i c a c t
p
a rt o f t h e
t
1 a n t -c e ll s a n d t o t he g ph e n o m e n o n o f s t r1 p e re ersa l d u r m g c o n v
t rac t x o h a n d t o t h e g e net l c re l a t x o n b e
d i i w b d
.

S i g n i fica nc e o f t h e s o -c a l le m t o t c fi g ures t ee n c o n t ra c t l o n a n s a n d m t ercal a t e d

in h
t es e c e lls u r t er s t u F
es o n re d h di d l S CS S t u es o n di
S t r uc t u re o f t h e Mo n o t re m a t a
Mi t o t i c
fi g u res m t e s e c e lls h
u rt er s t u F h di c a l i n v es t i g at x o n o f t he h a l r
A nu c ro l og l

on l ed o ne- m a rro w b I E x er m e n t a l
es
p i S t r u c t ur e V I T h e c o m p a ra t nv e h ns t o log y o f
Cy gi p i a
. .

II t o lo c , W l t h s ec a l re fe l e n c e t o t he t he l e g a n d w m g m u s c l e o f t he w sp , w 1 t h
d i
.

a t a s ug g es t m g n t ra c e ll u la r h e m o c y t o
p gi S p ec x a l re ere n c e t o t h e f
e no m e n o n o f ph
g e m c ae t 1 v 1 t y o n t he a rt o f t h e a n t - c e ll s
d st u p v
e r e ers a l d un n g c o n t ra c t no n a n d t o
b w
.

a nd t o t he S i g n i fi c a n c e o f t h e s o -c a lle t he g e n et x c x e la t x o n et
Mo n o t re ma t a A m 1 oro lo g c a l 1 n vc s t 1g a t 1 o n i ee n c o n t rac t no n
b a n d s a n d i n t el c a l a t e d d x s c s S t u d l es o n
o f t h e h a 1 r s t r uc t u re o f t he st u p dm e us c l e
Musc le s t r uc t ure V I T h e c o m p ara t w e h i s S y st e m A b se n c e o f t he v e na c a v m fer l o r
p ig
.
a
t o lo g y o f t h e l eg a n d m m; m usc l e o f t he by
.

m 1 2- m m e m r o . a s so ma t ed w 1 t h
w p w h p i
as , i t s e c a l re ere n c e t o t h e p h e f a
t h e d ra m a g e o f t h e o r t a l s s t e m m t o t h e p y
n o m e no n o f s t r 1 p e re ers a l u r n c o n t ra e v d i g c a rd m a l
i
t o n a n d t o t he g e net c rel a t o n et ee n i i b w y
S s t em m t o t h e c a r d m a l s s t e m A se n ce o f y b
c o n t r ac t 1 o n b d
a n s a n d n t e rc a l a t e d IS CS i d v v
.

di
S t u es o n st r 1 e pd t h e e n a c a a m f e x no r m a 1 2- m m p x g e m
b r y o , assoc na t e d 1 t h t he d ra m a g e o f t he “
Mus c u l at u re o f t he a t u a ea rt H p 0 1 tal 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
 IN D E X 49 9

A IL v
A m p h x b i a n l ar a e t o i nj e c t e c ro
of d A S P w1t h f
s p ec 1 a1 re ere n ce t o t he p h e
VV v
.

t o n 0 1 1 ( a sep t l c 1 n fi a m m a t l o n ) R ea c h o m e n o m o f s t r 1 p e re ers al d uu n g c o n
t l o ns o f c e l l e m t h e g
t ra c t mn a n d t o t h e e n e t i c r ela t 1 o u b e
Tu bu le A st u dyo f t h e 11 p 0 1 d c o n t e n t o f t h e tw a
ee n co n t ra c t 1 o n b n s a n d i n t er cal a t e
d
pd
d
d y
.

k i ne d 1 scs S t u d x es o n s t r1 e m u sc l e s t r u c
t ur e VI T h e c o m p a r a t w e h 1 s t o l og y o f
D e v el o p m e n t
.

T E R IN E g l an d s m m a n t h e l eg a n d w m g m u s c l e o f t h e
w p w h f
.

of t he n m u sc l e o f t h e as , i t s p ec 1al re er
e n ce t o t h e ph v
e n o m e n o n o f s t r 1 e re ersa l
EN A a ai f
n eri or i n a 1 2—mm p l g e m r o by h
d u rm g c o n t rac t 1 o n a n d t o t e g e n et 1 c
b w b d
c v
as soc x a t e dt p
h t he d ram ag e o f t he o rt al
.

rel a t i o n et ee n c o n t rac t 1 o n an s a n d
s y s t em m t o y
t h e c a r d m a l s s t em.Ab 1 n t er ca1a t ed d 1 sc s
. S t u d l es o n s t r1 e pd
sen c e o f t h e m u sc l e s t ruc t u re V I T h e c o mp arat w e
V b
er t e r a t e l u ng s C o n t r 1 b ut 1 0 n s t o t he h l S
.

p
h gy
i st o l o
.

o f t h e l eg a n d
t ol og y of t h e reS p l rat o ry s ac es o f t h e