A TE BOOK

GENERAL BACTERIOLOGY

WI L L I A M D O D G E F ROST
n
T F U N I VER S I T Y
g
AS SO C I A E P R O ES S O R O F B ACT E LO G Y I N T H E

OF WI SC SI N
1
o

g
NKL IN ‘
M C CA M P B E L L
OR 0 E R I O LO G Y IN T O HIO

U N I VE RS I TY
2 n
;
8

I LL US TR A TE D

N e wgo th
THE M A C M I L L A N CO M P A N Y
19 1 1
 C o wmcu r, '
19 10 ,

B Y T H E M AC M I LL AN C O M PAN Y .

S e t up l
a nd e e ctro ty p e d . P ub li h
s e d N o v e m be r, 19 10 . R e p rinte d
O cto be r, 19 11.

9m m
N e riuo nb 1
J . S C a sh i ng 00
. . B e rwi ck S m i h 00 t .

N o rwo o d , M a s s , U S A
. . . .
 T HI S B O OK

IS DE DI C AT E D

P
I N G R AT E F UL A P R EC I ATI O N

TO

B E NJ AM I N C UT LE R F R O S T

AN D

J A M E S M ccA M PB E L L

2 4 2 2 16
 PR E FA C E

ye r s o f its e xi ste nce h a s b e

BA CTE R I O L O G Y , i n th e fe w a ,

co m e ne f th e g r e t s ub j e cts o f th e d y
o o A k n wl dg ea a . o e

o f it is imp rt nt to tu d nts o f m ny diff r nt b r nch e s f

o a s e a e e a o

sci e n ce At th p r e s nt time it i u u lly t ught i n Eng l i h

. e e s s a a , s

s pe ki ng a untr i t l s t wi th
co p e i l r f re n e to me
e s a e a , s c a e e c so

p rti ul r ppli ti n s u h s m di i n
a c a a ca g r i ultur e Th e re
o , c a e c e o r a c .

i s g r wi ng d m nd f tr e ti s n th ubj t f g n r l
a o e a or a se o e s e c o e e a

b t ri l gy T h ut h r h v tt mpte d t m e t thi n d
ac e o o . e a o s a e a e o e s e e

in th p r nt v l um e
e e se o .

T h e m te r i l p r nte d h e r
a h a s b e n use d i n co ll g e
a e se e e e

cl s f
a se s numb r o f ye r with i nt e r
or a t nd p r o fit
e a s e s a .

Th e uth r wi sh to e i p
a th e i r s i nce r e
o s kno wl d gme nt t re s s ac e

fo th p r ti cul r h lp r e
r e aive d f r m th e wo rks f F i s ch r
a e ce o o e ,

L f a a r, E F S mith W ll . nd o th r s m e ntio ne d in th e b dy
.
, e s, a e o

o f th e t xt nd f th e id e s
e ,
nd i n pi r ti n impa r t e d by
a or a a s a o

th i r f rm e r te ch e r D n H L R u se ll o f th e Unive rs ity
e o a , e a . . s ,

o f Wi s c ns i n F ugg
o ti ns nd s s i stan e in th e r e vi sio n
. or s e s o a a c

o f th e m nu r ipt we e i nd bt d to P r o fe ss o r M
a sc , P R ave ar e e . .

ne l f th e U niv r s ity o f Wi co ns i n
,
o e s .

W D . .

E F M CC
Y
. . .

JUL ,
19 10 .
 T A BL E OF C O N T E N TS

PA R T I . IN T R O DUC T I O N
0

C HAPT ER I
TH E H I S T Y
OR BA T
OF C ER I O LO G Y

C HA T P ER II

S PO NTAN U EO S GE N T
ERA I O N

P AR T II . T H E M O R PH O L O GY O F B AC TER I A
C HA T P ER III

TH E F M OR AN D S T R U CT U R E OF BA TC ER I A

In tr d ti
o uc o n

H a pl o b a cte ria
iCo cc
B a illi c

S p irilla
Tri h ba t ria
c o c e

I nv l ti n f rm
o u ond p l m rph i
o s a eo o sm

T h di m
e i f ba t rial
e ns o ns o ll c e ce s

L o com ti n f ba t ria
o o o c e

R a pidity f m ti n o o o
X T AB L E OF CO NT E N T S

Br w nia m v o n o e me n t
Cap l su e s

S h ath
e s

M a gr p i g
ss ou n

Z o Ogl oe a a nd p s e ud o z o o gl oe a
P lli l e c e

C l ny o o

PT ER I V
CH A

R EP R O DU T NC IO H BA TOF T E C ERI A

In tr d ti
o uc o n

H apl o b a cte ri a
Fi i ss o n

Rat f m ltipli ati e o u c o n

S p r f r m ati
o e o on

E d pr n os o e s

Art h r p r os o e s

Tri h ba t ria
c o c e

C HA PT E R V
TH E M I N UT E S T R U CTU R E OF BA CT ER I A

Intr d ti n
o uc o

H i t l gy
s o o

C ll wall e

Pr t pla m o o s

M ta hr m ati e c o c a nd p lar granul

o e s

N l uc e us

Ch m i al tr t r
e c s uc u e

PA R T III . T HE M E T H O DS U S ED IN T HE

S T UDY O F B AC T E R I A

C HA PT ER VI

C U LTU RE ME DA I

In tr o d uc ti n o

Th e c hara t r c e o f fo o dr q rd
e ui e
 T AB LE OF CO NT ENT S

Th e co m p o siti o n a nd pre p arati o n o f m e dia

L iq id m diau e

S lid m dia
o e

L ique fi a bl e so lid m dia e

S ynth e ti c m e dia
CHA PT ER VI I

S T ER I LI ZAT I O N
Intr d ti n
o uc o

St rilizati n by p h y i al ag nt
e o s c e s

S unh gh t
Co ld
h at
Dry e
ir t a
D e c fl me
H igh dry h at e

M i t h at o s e

S tr a m i ng t a m
e s e

S t a m nd r pr
e r u e e ss u e

Di i nf ti n
s e c o

C HA PT ER V I I I
CULT U RE ME T H D A D APPA AT U
O S N R S

tr d ti n
In o uc o

M th ds f i lati n
e o o so o

Fra ti nal m th d c o e o s

Phy i l gi al m th d s o o c e o s

Dil ti n m th d u o e o s

Th e us e o f l i q ue fi ab l e so lid m diae

Su b lt r
cu u es

C HA PT ER I X
T H E M I C R O S C O P I C AL E AM NAT N
X I IO BA T C ER I A

Th e hi t ry f th m i r p
s o o e c o s co e

Th e ba t ri l gi al m i r p
c e o o c c o sco e

O il i m m r i n l e s o e ns

Abb nd n r e co e se

Th e exam i nati n f livi ng ba t ria o o c e

Th e pri n i p l f tai ni ng
c e s o s
 fi T AB L E OF CO NT ENT S

T T AX O N O M Y
“

PA R IV .

C HA PT ER X

T H E C L AS S I F AT N
IC IO OF BA
CT ER I A

Intr d ti n
o uc o

Th la i fi ati
e c ss c o n o fM ig la u

CHA PT ER XI

TH E R E LA I O T N SH I PS O F TH E BA TC E RI A

In tr d ti n
o uc o

B a t ria d fi d
c e e ne

Th p i t
e fr m bla n
o n s O t th r f rm e se ce o o e o s

Th r lati
e h i p t th p la t
e o ns s o e n s

C la i fi ati n f p la nt
ss c o o s

R lati hi p t th E m y t
e o ns s o e u ce e s

R lati hip t th Cya p hy

e o ns s o e no ce ae

R lati n hi p t th E ph y
e o s s o e u ce ae

R lati n hi p t th Myx m y t
e o s s o e o ce es

R lati hip t th Myx ba t ria

e o ns s o e o c e

l ti n h i p t a i m al
'

Th e re a o s s o n s

Pr t z a o o o

PAR T V . G EN E R AL P H Y S I O LO GY
C HA PT ER X I I
TH E R E LA I O T N OF B T A E NV NM N
AC E R I TO I RO E T

In tr d ti n
o uc o

Th f e ti f ba t ria
u nc o ns o c e

Ext rnal nditi

e co o ns

T h f d f ba t ria
e oo o c e

C h m i al m p iti n
e c co os o

Wat r e

O xyg n e

Ana ér bi lt r m th d o e cu u e e o s

T m p rat r
e e u e

T h th rm al d ath p i t
e e e o n
 T AB L E OF CO N T EN T S xfi i

Th e a ti n f h m i al
c o o c e c s o n ba t ria
c e

F rm ald hyd t
o e e , e c.

P ER X I I I C HA T

TH E R AT N BA T A E N V NM N T ( C nti n
EL IO OF C ERI To I RO E o ue d )
H FF C XT
T NA N D T N
E E E T OF E ER L CO I IO S

CHA PT E R X IV

TH E M E TA M
B O LI S OF BA T C ER I A

In tr d ti n
o uc o

Th g n ral hara t r
e e e c c e o f li f pr
e o ce s se s

Anab li m o s

C atab li m o s

R p irati n
es o

C HA PT ER X V
TH E P D UC
RO TS O F TH E M ETAB O LI S M BA T
C ERI A

In tr d ti n
o uc o

B a t rial nzym
c e d th ir m ha i m
e e s an e e c n s o f a ti
c on

A t lyti ba t rial nzym

u o c c e e es

PAR T THE BI OLO GY O F S PE C I AL I Z ED

G RO UP S O F B AC TER I A
CH A PT ER X VI

TH E P TTRO O RO P H I C BA T C ER I A

Intr d tio uc o n

N it i fyi g ba t ria
r n c e

N it g n fi i ng ba t ria
ro e -
x c e
 T AB L E OF CO N T ENT S

Su l p h r ba t ria
u c e

I r n ba t ria
o c e

CHA PT E R X V I I
TH E C H RO M G N C BAC
O E I T ERI A

Intr d ti n
o uc o

Ra g f l r
n e o co o

L ati n f p ig m nt
oc o o e s

S l bility o u

H i t ri al all i n
s o c us o s

CHAPT ER X V I I I
TH E PH T G N BAC
O O E IC T ERI A

In tr d ti n
o uc o

Di trib ti
s u o n

Fo o d r q ir m t e u e e n s

O xyg n r q ir m nt
e e u e e s

T m p rat r r q ir m nt
e e u e e u e e s

M th d
e f i lati n
o s o so o

Path g n i o e es s

C HAPT E R X I X
THE Z YM G N C BA T
O E I C ER IA

In tr d ti n
o uc o

O rga niz d nd rga niz d f n t

e a u no e e r Ie n S

L a ti a id f rm ntati n
c c c e e o

A ti a id f r m ntati n
ce c c e e o

B tyri a id f rm ntati n
u c c e e o

M tha
e f rm ntati n
ne e e o

H igh r al h l ar m ati
e d fatt y a id
co o s, o cs , a n c s

B a t rial f rm ntati n in th art

c e e e o s e s

CHA PT ER XX
T H E S AP R O GN E I C AND S AP R O P H I L I C B T AC E R I A

Intr d ti o uc on

Pr t lyti
o e o c e n zym es
 T AB L E OF CO NT EN T S XV

Th e m e c ha i m f p tr fa ti n s o u e c o n

Th e pr d t p r d d
o uc s o uce

CH A PT E R X X I
TH E PAT H N C BA TOGE I C ER I A

In tr d ti n
o uc o

Di a se i f ti d
s e s, ntagin ec o us a n co o us

L i t f p i fi i f ti di a
s o s ec c n e c o us se se s

L i t f n n p i fi i f ti di a
s o o s e c c n e c o us se se s

L i t f di a
s o f nk n w n a
se se s o u o c use

Th m a n by whi h ba t ria
e e stra n m itt d c c e a re s e

P ER XX I I C HA T

TH PATH G N BA T A ( C nti E O E IC C ERI o n ue d )

Th m d
e f a ti o f p ath g ni ba t ria
e o c o n o o e c c e

Pr f f th ti l gi al r lati n hi p
o o O e e o o c e o s

Th ff t
e e f th p ath g i ba t ria n th b dy
e c s o e o e n c c e o e o

Th p i
e pr d t f ba t ria
o s o no u s o uc s O c e

Pt m ai n o s

T xi n o s

E nd t xi n o o s

PT ER XX I I I C HA

TH PAT H G N BA T A ( C nti E O E IC C E RI o n ue d )
Th t xi ba t rial pr t i
e o c c e o e ns

Ana p hylaxi t ba t rial pr t i n s o c e o e s

B a t rial h m ly i n
c e e o s s

Antit xi n o s

A tiba t rial
n b ta
c e su s nce s

Aggl ti i n u n s

O p ni
so ns

Anti e nz ym e s
Tabl e a ntib di
Of o es

Th e fa t r whi h i fl
c o s c n ue nce a nd mo di fy i nfe c ti o ns

V ir l f ba t ria
u e nce o c e

N m b r f ba t ria
u e o c e

Av n f inf ti n
e ue o e c o
xvi T AB L E OF CO NT E N T S

bj t i f t d
Th e su e c n e c e

Imm ity t p ath g i ba t ria

un o o e n c c e

Th th ri
e f i m m nity
e o e s o u

C HA PT ER XX I V
THE BA T C E R I AL DI S EAS ES OF PLAN T S

In tr d ti n
o uc o

E arly n p ti n co ce o

Pr t e se n pti n co nce o

Th m th d f i nf ti
e e o o e c on

PA R T VI I . TH E DI S T R I BUT I O N B AC TER I A
CH A PT ER X XV

TH E BA T C ER I A O F T H E S OI L

Distrib uti o n rdi ng t a cc o o habitat

Di strib uti o n
i n th il e so

Th ep i f ba t ria
s e c e s o c e

B a t ria in th rfa lay r

c e e su ce e s

Th l g m
e e nitr g n fi ing ba t ria
u e or o e -
x c e

Th nitri fyi ng ba t ria

e c e

Th d nitri fy i ng ba t ria
e e c e

CH A PT ER XXV I
THE B T AC ER I A O F T H E AI R

Intr d ti n
o uc o

C n diti n i n th air
o o e

Th erigi n f th ba t ria f th air

o o e c e o e

Q a ntitativ di trib ti n
u e s u o

S a nal di trib ti n
e so s u o

Th p i f ba t ria in th
e s ec es o c e e ai r

C HA PT E R XX VI I
TH E BA CT E R I A WAT A D
OF ER N S EW AGE
Intr d ti n
o uc o

Th e m b r f ba t ria i n wat r
nu e O c e e

Th eigi n f th ba t ria in wat r

or o e c e e
 TAB L E OF CO NT E NT S xvfi

W at e r
a naly i s s

Th p rifi ati
e u c o n o f wat r e

S wag
e e

Th p rifi ati
e u c o n o f se wag e

C HA PT ER XX VI I I
T HE BA CT E RI A O F M I LK AN D ITS P DU T
RO C S

In tr d ti n o uc o

T h ba t ria f th
e dd r
c e O e u e

T h n m b r f ba t ria i n m ark t m ilk

e u e o c e e

Th er f ba t ria in m ilk
s o u ce o c e

Th eta m i nati n f th m ilk fr m th a ni m al

co n o o e o e

Th ntra
e e f ba t ria i t th m ilk fr m th
n ce o c e n o e o e

C l a li n
e n f m ilki g t n il
e ss o n u e s s

T h m ilk r
e e

Th ar f th m ilk
e c e o e

C HA T P ER XX I X
B T A M
TH E AC E R I OF ILK A D NP DU T IT s RO C S ( Co nti n u e d )

Th p i f ba t ria i m ilk
e s e c e s o c e n

L a ti a id ba t ria
c c c c e

S li my m ilk
B l m ilk
ue

R d m ilk
e

Th di a e pr d i g ba t ria i m ilk
se se -
o uc n c e n

Path g i ba t ria i m ilk a ft r it i draw

o en c c e n e s n

Typ h id o

A iati h l ra
s c c o e

Di p hth ria e

S arl t f v r
c e e e

CH A PT ER XXX
TH E BA CT ER I A H H U M AN B D Y
OF T E O

Intr d o uc ti n o

Th ba e t ria
c e Of th e s ki n a nd e xp o se dm uco us me m bra ne s

Th ba e t ria
c e o f th e m o u th
D ntal
e c ari e s

Th ba e t ria
c e o f th e s o t m ah c a nd i nt ti n
es es
 HI S TOR Y OF B AC TE RIO LO GY
t n
I ntro d uc i o B act e ri o l o gy is that branch o f bi o l o gical
.

sci e nce whi ch tr e at s o f bact e ri a B act e ria a e a de finit e and

. r

circum scrib e d gr o up o f mi cro Organisms lying n e ar th e base

o f th e ladd e r o f lif e The y will b e accurat e ly de scrib e d in
.

th e f o ll o wi ng chapt e rs I t is fr e qu e ntly fo und de si r abl e to

.

discuss o the r micro o rganisms with th e bact e ria b o th o n ,

acco unt o f th e fact that th e y a re cl o s e ly r e lat e d and als o b e ,

caus e th e y p ro duc e similar chang e s o r a re studie d by similar ,

me tho ds Th e mo re inclusive t e rm micr o bio l o gy which

.
,

is use d by th e F r e nch w o uld b e much b e tt e r than th e t e r m

,

“
bacte rio l o gy to de scrib e th e subj e ct matt e r di scus se d in thi s
”
-

b o o k C usto m h o w e ve r has fi e d th e w o r d ba ct e ri o lo gy
.
, ,
“
x
”

in this co untry and it will b e us e d h e re but no t infr e que ntly

, ,

in th e bro ade r s e n se While bact e ria are a sso ciat e d with

.

di se ase pro ductio n th e maj o rity o f th e m a re no t di e as e p ro

,
s -

d ucing and if th e r e w e r e no dis e as e p ro d ucm g bact e ria in th e

,
-

wo rld b a ct e ri o l o gy w o uld still b e a gr e at sci e nce I n thi s

,
.

b o o k bact e ria will b e co n side r e d fro m a bio l o gi cal st nd a

po int and th e ir structu r e and functio ns s tudi e d with o ut

,

any particul ar applicatio n in Vie w I t is th e re f o r e a ge ne ral .

T
B
 R
I NT ODUC I O T N
bact e rio l o gy and pave s th e way fo r th e app l i e d phas e s o f th e
subj e ct Th e applicati o ns o f bact e ri o l o gy a e o f particular
. r

s e rvic e in m e di cine agriculture e ith e r in th e de partm e nts o f

, ,

dairying animal husbandry ho rticultur e o r so ils ; sanitatio n

, , , ,

e ngin e e r ing h o m e e c o n o m ics and th e c o mm e rcial i ndustri e s

, ,
.

Whate ve r th e pplicatio n h o w e ve r ce rtain fu ndame ntal

a , ,

facts and m e th o ds must b e co n side re d and an att e mpt is ,

made to pre se nt th e se in thi s ge n e ral w o rk .

H i s to ry —
. Th e e xi s t e nc e o f bact e r ia o n th e gl o b e ant e

dat e s that o f man and ye t th e ir p r e se nc e was no t su sp e cte d

,

until a fe w c e nturie s ago and the ir actual e xi st e nc e was no t

de mo nstrate d until much lat e r P r o bably th e firs t auth e ntic .

o b se rvati o n o f m icro Orga ni sm s was made by Kirch e r in 165 9 ,

wh o d e mo n s trat e d minut e li Ving wo rms in putrid m e at

“ ”
,

milk vin e gar ch e e s e e tc but h e did no t de s crib e th e ir f o rm

, , ,
.
,

o r charact e r Antho ny V an Le e uw e nh o e k a Dutch natu

.
,

ralist wh o liv e d f r o m 63 to 1 7 3 disco ve r e d bact e r ia ab o ut

1 2 2 ,

1683 . Le e uwe nh o e k was a lin e n w e ave r by trhd e but a s a ,

man o f s o me l e i sure h e l e arne d to g rind l e n se s fo a pastim e r ,

and made th e m so p e rf e ctly and u se d th e m with such ke e n

ne s s that h e was abl e to se e many m i cro sc0p i c o bj e ct s fo th e r

fir st tim e and amo ng the m th e bact e ria Le e uwe nho e k no t

,
.

o nly saw th e m but h e accurat e ly d e s crib e d th e m and mad e

,

dr awi ngs o f th e m a s we ll I n a l e tt e r to th e R o yal So ci e ty

.

o f Lo ndo n in 1683 h e s aid : I saw with w o nd e r that my

“

mat e rial co ntain e d many tiny animal s whi ch mo ve d ab o ut

in a m o st amusing fa shi o n ; th e la r g e st o f th e s e (A Fig 1) ,
.
 HI S O T RY OF B ACTERI O LO GY 3

Sho we d th e live l ie s t and mo st active m o tio n mo ving thro ugh ,

th e wat e r o r saliva a s a fi sh o f pr e y dart s thr o ugh th e se a ;

th e y we re fo und e ve rywhe r e altho ugh no t in large numb e rs

, .

A se c o nd kind wa s S imilar to that mark e d B ( Fig i ) The se . .

s o m e tim e s spun a ro und in a circl e like a to p and so me tim e s ,

de scrib e d a path like that S h o wn in C D ( Fig ) th e y w e r e

— . I

p re se nt in la rge r numb e s A third kind c o ul d no t b e dis

r .

tingui sh e d so cl e arly no w th e y app e a r e d o bl o ng no w ,

quit e ro und Th e y w e r e so ve ry small that th e y di d no t s e e m

.

large r than th e b o di e s mark e d E and b e side s th e y mo ve d so

, , ,

rapi dl y that th e y w e r e co ntinually running i nto o ne an o th e r ;

th e y lo o ke d lik e a swarm
A
o f gnat s o r fli e s d nci ng
a

ab o ut to ge th e r I had
.

B t
C fi / ‘

th e imp r e ssi o n that I was

lo o king at s e ve ral tho u E Q

0
sands in a give n part o f
th e wat e r o r saliva mix e d

with a parti cle o f th e ma

t o t ial fro mth e t e e th no
F C 3 2 2

large r than a grain o f sand ,

e v e n wh e n o nly o ne part o f

th e mat e rial was add e d to

F IG L w h k fig r
. 1
— . Th eeu en oe
’
s u es . e

nin e part s o f wate o r sali va ld t k w fig r f ba t ria Aft r

'

r . o es no n u es o c e . e

F urth e r th e g re at e r part
F i h r sc e .

o f th e mat e rial c o n sist e d o f an e xtra o rdinary numb e r o f r o ds ,

o f wid e ly di ff e r e nt l e ngths but o f th e sam e diam e t e r

, So me .
4 I N T R ODUC TI O N
w e r e curv e d s o me straight as is sho wn in F th e y lay ir
, ,

re gula rly and w e r e i nt e rlace d Since I had pr e vi o usly se e n

.

living animal cul ae o f thi s s ame kind in wate r I e nde avo r e d ,

to o bs e rv e wh e th e r th e r e was lif e in the m but in n o n e di d I ,

se e th e s mall e st m o v e m e nt that might b e tak e n as a S ign o f

life . L e e uw e nh o e k s o bs e rvati o ns w e r e pur e ly o bj e ctiv e

’
,

and lacke d th e spe culative e ntire ly Oth e r writ e rs ho we ve r.

, ,

within th e n e xt ce ntury th e o rize d up o n his di sco ve ri e s and

wo rthy o f particular m e ntio n is M arcus Anto nius P l e nciz ,

a physician o f Vi e nna Pl e nci pro p o s e d a g e rm the o ry o f

. z

di se ase in 7 6 a nd taught th e e tio l o gical re latio nshi p o f

1 2 ,

Le e uw e nh o e k s a nimal culae to vari o us di s e as e s He lik e

’
.

wi se i nsi st e d o n a cau sal r e latio n ship O f th e se minut e and

unnumb e r e d f o rm s f lif e to th e pro c e ss e s o f d e co mp o siti o n
o .

I mp o rtant di sco v e r i e s r e ga r ding bact e ria w e r e mad e by

O F M ii l l e
. . r, a di stinguishe d Danish o Ol o gist M il l l e made z . r

many imp o rtant Ob se rvati o ns in r e gard to th e fo rm and struc

ture o f bacte ria and studi e d se ve r al typ e s so clo s e ly that the y
,

can b e plac e d in o ne o r an o th e r o f th e p r e se nt f o rm typ e s .

He fir s t us e d s uch t e rms as bacillu s S pi rillum and Vibri o , , ,

which a re in co mm o n use no w He als o mad e an atte mpt to

.

w o rk o ut ( 7 86) a classificatio n and in S O do ing made an

1 ,

o b se rvati o n w hi ch h as b e e n appr e ciat e d by all lat e r w o r k e r s

in this fi e ld ; name ly Th e diffi culti s that b e se t th e inve sti

,
e

ga tio n o f th e s e micr o sc o pi c a nimal s a re co mpl e x ; th e sur e

a nd d e fi nit e d e t e r mi nati o n ( o f sp e ci e s) r e qui r e s s o mu ch tim e ,

so mu ch o f acum e n o f e ye and judgm e nt so much p e r se v e r ,

 HI S O T RY OF B AC T ERI O L O GY 5

ance and practice that th e re is hardly anything e l se so diffi

,

cul t Ano the r wo rke r wh o de se rve s sp e cial m e ntio n is Ehre n

.
”

b e rg ( 7 9 5
1 I n 83 6 h e publi sh e d hi s g r e at w o rk o n
1

I nfus o ria a nd I n thi s tr e ate d o f th e bact e ria as no o ne had b e

,

fo re hi m He r e co gnize d quite a ccurate ly th e limitatio ns o f

.

thi s g ro up a nd diff e r e ntiat e d the m fro m th e true pro to zo a He .

al so re co gnize d th e principal fo r m typ e s in whi ch th e bacte ria

a re u s ually di vid e d Ano th e r w o rk e r s till sh o ul d b e m e n
.

ti ne d F e rdi na nd C o hn ( 8 8 H e was P r o f e s s o r o f
o ,
1 2

B o ta ny in th e Univ e r sity o f B r e slau Hi s w o r k s c o ntribut e d

.

gr e atly to th e advanc e o f o ur kno wle dge Of bacte ria A ne w .

e ra in th e hi st o ry o f bact e ri o l o gy b e gan with th e w o rk o f

L o uis P a s t e ur ( 8 2 2 1 P a st e ur alth o ugh by inclina

,

tio n a nd t r ai ning a ch e mi s t was e a rly l e d by hi s che mical

,

r e s e a r che s to th e study o f th e re latio n o f rnic o Organism s r

to va ri o us natu r al p r o ce s se s su ch as f e r m e ntati o n d e cay

, , ,

a nd put r e facti o n a nd lat e r to th e r e lati o n o f th e se o rganism s

,

to di se as e p r o c e sse s in man and a nimals L at e r still h e t o o k

'

up th e study o f th e sci e ntific m e tho ds o f p r e ve nting and curing

the se dise ase s Vie we d fro m any stand p o i nt hi s r e s e arch e s
.
-

a re o f th e pro fo unde st imp o rtance He intro duce d th e .

e xp e rim e ntal m e th o d and u se d it with t e lli ng e ffe ct in th e

study o f bact e ria and alli e d mic r o o rganism s F ro m an o b .

scure p o sitio n int e re sting o nly to th e p ro fe ssio nal bio l o gi s t

, ,

P a s t e ur rai se d the se m icro o rgani sms to a place o f g re ate st

inte r e s t o n acco unt o f th e i r significanc e in th e pr o ductio n o f
change s o f g re ate s t imp o rta nce to man P aste ur may w e ll .
6 I NTR ODUCTI ON

be co nside re d th e fo unde r o f bacte ri o l o gy but it r e maine d ,

fo r R o b e rt K o ch ( 1843 by his e p o ch making di sco v -

e ri e s to plac e bact e ri o l o gy in th e p o siti o n o f an i nd e p e nde nt

,

sci e nce I n 87 6 K o ch an o b scu r e G e rman physi cian pub

. 1 , , ,

l ish e d hi s articl e o n th e r e lati o n o f B act e r ium a nth r aci s to

S pl e nic f e v e r in cattl e. T h e r e lati o n wa s stat e d s o cl e arly and

th e argum e nts w e r e so c o nvi ncing that th e pap e r attract e d

wide atte nti o n and furni sh e d a m o de l fo r i nve stigati o n o f th e

r e latio n o f mi cro Orga ni sm s to dise as e pro c e sse s Si y e ars . x

late r h e de scrib e d a ne w m e th o d fo is o lating bact e ria This

r .

was by m e ans o f a cl e ve rly de vi se d l iq ue fi ab l e s o lid m e dium

a nd i s k no wn as th e g e latin plat e cultur e m e th o d T h e im .

p o r tance o f th e di sco ve ry o f thi s m e th o d to th e d e ve lo pm e nt

o f bact e ri o l o gy ca nno t b e o v e r e s timat e d Ab o ut thi s tim e
.

als o Ko ch aide d and e nco urage d th e de ve l o pm e nt o f th e

bacte rio l o gical mic ro sco p e pro vide d with a n Oil imm e r sio n
l e ns a nd an Abb e co nde nse r He al so de ve l o p e d and appli e d
.

th e use o f th e anilin dy e s fi r s t int o d iI ce d int o mi cr o s c o pical

,
r

t e chnique by We ige rt F o llo wi ng thi s wo rk th e disco ve rie s

.
,

in bacte rio l o gy have b e e n rapid a nd almo st co ntinu o us I n .

a ge ne ral way h o we ve r th e imp o rtant di sc ve ri e s have

, ,
o

b e e n gro up e d ab o ut di sco ve ri e s o r m dificatio ns f t e chnique

o o .

F o r e xampl e f o ll o wi ng th e u se o f th e a nilin dy e s a nd th e
,

p e rfe ct e d micro sco p e th e bact e ria o f a co n side r abl e numb e r o f

,

th e imp o r tant dis e as e s w e r e di s c o v e r e d a s fo i ns ta nc e th o s e

, ,
r ,

o f r e lapsi ng f e v e r p ne um o nia typh o id f v e r a nd th e p r o t o z o an

, ,
e ,

o f mala ria . F o ll o wing th e i nt r o ducti o n o f th e plat e cultur e

 T RY
HI S O OF B ACTERI OL OGY 7

a co nside rabl e numb e r o f mi cro o rgani sms r e late d to im ,

p o rta nt di s e ase s w e r e gro wn artificially and the ir re latio n

, ,

to th e di s e as e pr o ce s s d e finit e ly e s tabli sh e d s uch as th o s e o f

,

cho l e ra diphthe ria typho id f e ve r and tho se asso ciate d with

, , ,

supp urati o n M o r e r e c e ntly with the d isc o v e r y o f No vy

.
,

that ce rtain p ro t o zo a c o uld b e gr o wn artificially the r e has ,

‘

b e e n a gre a t a wa ke ni ng al o ng this li ne a nd imp o r ta nt d is

co v e ri e s hav e b e e n mad e. I n 88 P a s t e ur di sco ve r e d that h e
1 0,

co uld pr o t e ct a n a ni m al f r o m a n at tack o f a dis e a s e by ino cu

lating an atte nuat e d cultur e Th e wo rk was first do n e with

.

th e g e rm o f chi ck e n cho l e r a but lat e r with B act e rium anthra

cis and this was th e b e ginning o f th e s ci e ntific de ve l o pme nt

,

o f pr o t e ctiv e i no culati o n e mpi r ically di s c o v e r e d by J e nn e r

, ,

whe n h e de vis e d th e smallp o x vaccinatio n I n 89 5 B e h ri ng. 1 ,

and R o ux di s co ve re d th e anti to xin fo r diphthe ria Thi s

-
.

was th e b e ginni ng o f th e imp o rtant w o rk o n se rum th e rapy

which is b e ing de ve l o p e d at th e pr e se nt tim e and it was also ,

th e starting p o int fo r much o f th e imp o r tant w o r k whi ch has

b e e n do n e o n th e th e o ri e s o f im m unity .
 CHAP TER I I

S P ONTANE OUS GE NER ATI ON

n u cti o n I n e arly time s it was th e ge n e ral b e li e f
I tro d .

o f m o s t p e o pl e wh o di d no t acc e pt as lit e ral th e B iblical a c

co unt o i th e cr e ati o n o f th e wo rld that many fo rm s o f lif e

co uld ari se de no w This ide a o r th e o ry is usually sp o k e n
.

o f a s th e th e o ry o f sp o ntan e o us g e ne r ati o n o r abi o g e n e si s

,
.

E a l y P e ri o d
r . At fi r s t it was supp o s e d that ma ny high e r
fo rm s o f lif e including s o m e o f th e birds and mammals co uld
, ,

ari se in this way F o e xampl e amo ng th e e arly G re e ks we

. r ,

find that Anaximande r Of M il e tu s h e ld that a nimals w e re

fo rm e d fr o m m o i stu re and Emp e do cl e s o f Ag ige ntum b e
,
r

l ie v e d that all o f th e living b e ing s o n th e e arth ar o se thr o ugh

sp o ntan e o us g e n e ratio n : Aristo tl e whil e no t having such

,

ge n e ral b e lie f in thi s th e o ry do e s ass e rt that So me time s

,
“

animals are fo rme d in put r e fying so il so m e tim e s in plants , ,

s o m e tim e s in th e fluid s o f o th e r animals He al so state d .

”

that e ve ry dry sub stanc e which b e co m e s m o i st and e v e ry ,

m o ist b o dy whi ch b e co me s dry pro duce s living c re atu r e s

, ,

pr o vide d it is fit to n o u rish th e m L ate r w find Ovid d e

.
‘

f e nding thi s do ct rin e and in th e M iddl e Ag e s Vo n He lm o nt

,

g iv e s dir e cti o ns fo r th e artificial pr o ducti o n o f mic e which

8
 S P O NTAN E O US GE NERATI ON 9

we re s o me thi ng afte r thi s o rde r : An o l d pie ce o f cl o th and

s o m e s e e d c o rn w e r e to b e plac e d in a jar a nd aft r s tan di ng
,
e

th e pr o p e r l e ngth o f tim e w o uld sur e ly pr o duc e mic e K ir .

che r actually figure s ani mals claime d to have b e e n pro duc e d

unde r his o wn e ye .

Duri ng this p e ri o d th e th e o ry o f sp o ntan e o u s g e n e rati o n

was univ e rs ally acc e pte d . I n th e s e v e nt e e nth c e ntury

Al e xand e r R o ss c o mm e nting o n S ir Th o mas Br o wn s do ubt

’
,

as to whe the r mi ce we re bre d by put r e factio n says : TO ,

que stio n this is to qu e stio n r e a so n se nse and e xp e ri e nce

, ,
.

S e co nd P e ri o d .
— Th e fi r st e xp e rim e nts whi ch s e r i o u sly

que stio ne d thi s th e o ry w e r e th o se d e vise d by F rancisco R e di ,

in 668 R e di was a phy sician to th e G rand Duke o f Tuscany

1 . .

I n watchi ng m e at pas s fr o m th e fr e sh to th e putrid stat e h e

no tice d that b e fo re th e maggo ts app e a r e d whi ch w e re co m

,

m o ul y supp o se d to b e th e r e sult o f sp o ntane o us g e n e ratio n ,

fli e s co uld invariably b e se e n buzzing ab o ut th e m e at Th e .

ide a o ccurr e d to him that the se magg o ts might b e th e p ro g e ny

o f th e fli e s and h e put so m e m e at in a jar a nd co v e r e d th e
,

m o uth o f th e jar with a pi e ce o f pap e r a nd fo und that alth o ugh

;

th e m e at p utre fi e d magg o ts did no t app e ar L at e r th e pap e r

.

was r e place d by gauz e . M e at pro t e ct e d in thi s way w o uld

putre fy but wo ul d no t de ve l o p magg o ts Th e fli s wo uld
,
. e

de p o sit th e ir e ggs o n th e gauze whi ch w o uld no t all o w th e m

,

to r e ach th e m e at .This fi rs t se ri o us bl o w to th e th e o ry o f
sp o ntane o us g e ne ratio n was f o ll o w e d by o th e r co nclusive
e xp e rim e nts by S wamm e rdam ,
Val l isnie ri and o th e rs so
, ,
I 0 I NTR ODUC T I ON
that it was no t l o ng b e f o r e th e ide a that th e high e r fo rms o f
life aro se in thi s way was di s carde d .

Thi r d P e ri o d . Wh e n th e sci e ntists o f th e day b e gan to

study th e micro o rganism s whi ch Le e uwe nh o e k had di s
co ve r e d th e th e o ry o f sp o ntan e o us g e n e ratio n was re vive d
, ,

chi e fly as th e r e sul t o f e xp e rim e nt s by N e e dham ( 17 13

and o ne o f th e fi e rc e s t battl e s k no wn to sci e nce was f o ught
o v e r thi s th e o ry I t was v e ry difficult to u nd e r s tand h o w it
.

was that putr e scibl e flui ds w o uld sp o il aft e r th e y had b e e n

h e at e d if it we re no t tru e that th e life which app e ar e d in th e m
was Sp o ntane o usly ge ne rate d I t was g e n e rally b e li e ve d that
.

th e t e mp e ratur e O f b o iling wat e r was suffi ci e nt to kill all f o rms

o f lif e a nd th e wid e di s tributi o n o f nu cro o rgam sm S o r th e i r

‘

sp o r e s in th e air was no t g e n e rally unde rs to o d One o f th e .

first o f th e crucial e xp e rime nts whi ch p o int e d o ut th e e rro r

o f N e e d h a m s e xp e rim e nts and l e d to th e o v e rthro w o f th e
’

th e o ry o f sp o ntan e o us ge n e r atio n was mad e by Abb e L azzaro

S pallanzani ( 1 7 2 9 a fe llo w co untryman o f R e di I n
-
.

1 7 7 7 h e fill e d flask s with o rga nic s o luti o ns b o il e d th e m fo r

, ,

thr e e quarte rs o f an ho u r and th e n s e al e d the m and place d

,

th e m unde r co nditio ns favo r abl e fo th e gro wth o f mic ro r

o rgani s m s with o ut h o w e v e r havi ng th e m d e v e l o p

, , S p al l a n.

zani s critics o bj e cte d to his e xp e rim e nts o n th e gro und that

’

air had no t b e e n admitte d whi ch the y claime d was e ss e ntial

,

to th e lif e o f th e s e m ic o O gani sm s
r r F ranz S ch ul ze in 83 6
.
,
1 ,

se t asid e th e Obj e cti o ns o f th e se critics o f S pallanza ni by ar

ranging a flask with two glass tube s b e nt at right angl e s so

 S P ON TANE O U S GENERA TI ON I I

arrange d that air co uld b e drawn thro ugh th e flask Th e air .

,

ho we ve r was fi r st drawn thro ugh sulphuric acid and it was

, ,

fo und that whe n thi s was do n e sp o ntane o us ge n e ratio n di d

no t o ccur in pr e vi o usly b o il e d d e co cti o ns Fi T critics
( g h e .
,

FIG . 2 . Sc h lz Exp ri m t B l b tain lph ri a id thr g h whi h

u
’
e s e en . u s co n su u c c ou c ai r

wa s drawn b f r it w all w d t nt r th b ttl Aft r L f

e o e as o e o e e e o e . e a ar
.

h o we ve r said that th e sulphuric acid in so m e way had change d

,

th e air The o do r e S chwann th e n e xt y e ar p e rfo rm e d e xp e i

.
, ,
r

m e nts similar to tho se o f S chul ze e xce pt that h e sub stitute d ,

fo r th e sulphuric acid m e tal tub e s whi ch w e r e h e at e d re d

, ,

h o t wh e n th e air was drawn th ro ugh ( F ig P e rhaps .

S chwan n s m o s t imp o rtant Ob se rvati o n was th e fact that

’

wh e n ce rtain che mical sub stance s p o tassium a r s e niate ,

e tc
. we re adde d to th e se so lutio ns putre factio n did no t
take place S chwann may the re fo re b e r e garde d as th e
.

fo unde r o f th e scie nce o f antise ptics One o f th e m o st im .

po rtant e xp e rim e nts supp o rting th e the o ry o f bio g e n e sis was

I 2 I NT RODU CTI O N
p e rf o rm e d by S chro de r and Vo n Dusch in 185 4 Th e se wo rk .

e rs sub s titut e d c o tt o n fo r th e sulphuric acid and h e at w hi ch

had fo rm e rly b e e n us e d and sho we d that wh e n air was drawn

thr o ugh co tto n and allo w e d to co me in co ntact with p utre s

FI G 3. . hwann Exp ri m t
Sc
’
s e en . Ai r i s all w d t nt r th fla k nly aft r it
o e o e e e s o e

w h at d i th
as e e n e ho t co il d t b Aft r L f
e u e . e a ar .

cibl e mate rial that had b e e n suff cie ntly h e ate d the re was no i

putr e factio n Th e se r e sul ts we r e r e garde d by scie ntific m e n

.

as de finite ly se ttling th e que stio n .

M o d e rn P e ri o d A F r e nchman F o uch e t dir e cto r o f th e

.
, ,

N atural Hi s t o ry M us e um o f R o u e n r e o p e ne d th e qu e sti o n ,

in 85 9 up o n th e o r e tical co nsid e rati o ns

1 P ast e u r in 1860 .
, ,

e nt e r e d this fi e ld and with r e markabl e ing e nui ty and S kill

,

r e fute d th e co nclusio ns o f F o uche t and hi s fo l lo we rs Whil e .

 P N TANE O U S GENERATI O N
S O I 3

P aste ur adde d littl e that was new hi s e xp e rim e nts w e r e o f

,

such a natu r e that th e y l e ft littl e do ubt in th e minds o f th o s e

wh o b e cam e acquaint e d with the m that sp o ntan e o us g e n e ra
tio n had no fo undatio n in fact and since his time it has
, , ,

ge n e rally b e e n b e lie ve d that ho we ve r life fo rm e rly o igi r

h at e d it d o e s no t no w o rigi nat e sp o ntan e o u sly and that th e

,

l aw o f H arv e y o mne vi vum e x o vo o r i ts m o dificati o n o mne

,

vi vu m e x vi vo appli e s to th e l o w e r as w e ll as to th e high e r f o rms

o f li fe
.
 P A RT II . M O R P H O L O GY

C HAP T ER I I I

F OR M AN D S TR UC TUR E OF B AC T ER I A
In tro d ucti o n Th e fo rm and to so m e e xte nt th e struc
.
,

tur e o f bacte rial ce lls is subj e ct so so me variatio n de p e ndi ng

,

up o n th e te mp e rature at which cul tivati o n take s place and

FI G 4
. F rm typ f ba t ria P h t graph f m d l h wi g f rm d r la
. o eso c e . o o o o e s, s o n o an e

tiv iz e s M py ge var a r ;
. 1, 4 mi r
. o d rg i g fi i ;
e ne s . u e us 2 , 3, , c o co ccus un e o n ss o n

5 St
, py g ; 6 t trad ; 7 ar i a 8 B typh ; 9 B btili ;
r
. o e ne s , e s ,
s cn ,
. o sus , . su s

B a t anthra i ;
I O, c Mp. mm a ; 3 Sp csb rm i ri 1 1, 12 ,
s . co 1 , . o e e e .

th e che mical co nstituti o n o f th e m e dia o n which th e bacte ria

a re gro wn I n o rde r that accurate inf o rmatio n may b e
.

gain e d in r e gard to fo rm and s tructure it is n e ce ssary that

14
 F O RM AN D S R T U C TURE OF B ACT ERI A 1
5

th e vari o us sp e c e s o f ba cte ria b e gro wn o n standard m e dia

r .

T h e variati o n in f o r m a nd s t r u ctu r e i s o ft e n slight with so m e

S p e ci e s a nd p r o n o u nc e d i n o th e r s B act e r ia a re c o nv e ni e ntly
.

di vide d into two o rd e r s th e E ubacte ria (M igul a) o r t r ue, ,

bact e ria and th e Thi o bact ria o r sulphur bact e ria Th e

,
e , .

t r ue bacte ria with whi ch we are chi e fly co nce rn e d may b e co n

, ,

v e nie ntl y divid e d int o two sub o r d e r s a l o we r and S impl e r ,

f o r m kn o wn s th e H apl o b a cte ia ( Fi sch e r) a nd a high e r

a r ,

a nd m o r e c o mpl e x f o r m call e d th T r i ch o bact e r ia ( F is ch e r)

e .

B act e ria vary g r e atly in s hap e acc o rdi ng to th e g e n e ra and

S p e ci e s .

H apl o b a cte ri a or Lo we r r
B acte i a

Th e di ff e re nt f o rm typ e s o f th e h ap l o b acte ria a re sh o wn

in F ig .
4 .

C o cci . active o r v e ge tative stag e all th e l o w e r bac

I n th e

te ria a re co mp o se d f singl e c ll s o f minute s ize Th simpl e st

o e . e

o f th e se is a r o und ball lik e o gl o bula r f o rm in whi ch all th e

-
r

di am e t e r s a re e qual T o thi s fo rm typ e th e t e rm co ccu s has

.

b e e n giv e n Th e g ro up i subdivide d acco r di ng to th e cha r

. s

a cte ri s ti c m e th o d in whi ch th e c o cci r e p r o duc e i n di ff e r nt e

plan e ( Chap VI ) I f a c o ccus divide s so that its succe ssiv e

s . .

divi io n walls a re pa r all e l th e r e sult is a chain o f co c ci C cci

s ,
. o

whi ch r e p ro duc e in thi s mann e r are

.

S t r e p py o g e ne s) ( Fig
. T h e individ ual c o cci in th e
.

str e pto co ccus r e mai n co nne ct e d o r glue d to ge th e r in a chain

by a ge latino us se c r e tio n I f a c o ccus divide s in two d ire c
.
16 M O P HO O R L GY
tio ns at right angl e s in o ne plane th e r e sultant is a mass ,

usually irr e gular b e caus e r e pro ductio n (fi ssio n) do e s no t tak e

place in e ve ry co ccus at
th e sam e i nt e r val B a ct e r ia .

( c o cci ) whi c h r e pr o duc e in

this way are c ll e d m i c cci a cro o

( E M ic r o c cc u s py o g e ne sx . o

v a r aur e u s) I f by r e a so n . .
,

o f th e g e lati no us e nve l o p e
OOOCI
Zb
(b o
o
m m which O fte n sur ro unds bac
FIG 5
—S tr pt
. py g
e Aft r t e r ia th e cl ump O f CO CCI O f
o co ccu s o e ne s . e ,

d w“
.

S hm d t
'

an
a ce rta n sp e c e s a e h e ld to
C 1 e
l i r

g e th e r th e y a re call e d stap hyl o co cci ( Fig

,
Th e ma s f o r m d . s e

is no t unlik e a bunch o f g r ap e s I f a sp e cie s e xhibits a te nd .

e ncy fo r two c o cci to s tick t g th e r th e t e rm o e ,

dipl o cco ci ( Fig 7 ) i s appli e d ; a nd if g r o up s O f

.

1 s o

f o ur c o cci o r multipl e s o f f o ur o ccur to g e th e r 2 ,

th e t e rm te t a c cci is u se d r o .

g b
m This f o rmati o n i s al so kn o wn F 6
l
_
w
St ph y ro . , a .

as me i sm ope di a ( F ig r
o co ‘

w
.

fl Th e c o ccus may r e pro duc e in th r e e di ff e r e nt

a diam e t e rs all at right a gl a h th r
O n e s to
,
e c o e

i a nd in two d iff e r e nt plan e s a n

Di11 d a s a r e s ult
F —
IG~ 7 3 000CC
, ,
(M g . h )o no rr oe z
e .

A f t r S h m i d t pr o vidi ng divi si n tak e s pla ce at e qual in

e c o

d Wi
an
te rv al s a cub e will b e f o r m e d B a ct e ria
e s .

,
.

( co cci) r e pro duci ng in thi s charact e ri sti c fa hi o n a e kn wn s r o

as sa ci na (E S ar aurantiaca) (Fig
r x . . T h e siz e o f th e .
 F ORM AN D S TRUC T URE OF B ACTERIA 17

m e mb e rs g e nus k no wn as th e co c ci va ri e s f ro m
of th e

micro n to 2 micro n s in diam e te r M icro co ccus p o g e d ie ns . r r

( S ch rO t e ) i s said r

to b e as small as

micro n in
di am e t e r Co cci .

d o no t pro duc e

sp o r e s A f e w .

sp e cie s o f co cci are a Q

00 0
m o til e and p o sse s s

flage lla ( E M i x .

cro co ccus agil is )

B acd h I f We
P la litt rali ) Aft r
FI G 8
. .
—M e rism o
p e d ia ( no co ccus o s . e

co nce ive o f a co c S h m idt d W i c an e s .

cus b e co ming e l o ngat e d so that o ne axis is o f gre ate r le ngth

.

than th e o the rth e re re sul ts a cylindrical ro d To bacte ria o f

, .

thi s typ e th e t e rm bacil l us is appl ie d Acco r ding to M igula s .

’

clas sifi catio n th e te r m bacillus m e ans a ,

de finit e m o til e sp o r e pr o ducing ro d , ,

-

shap e d bacte rium We apply th e t e r m .

h e r e ho w e ve r in its ge ne ral se n se to l l
, ,
a

ro d shap e d bact e r ia B acilli may p o sse s s

-
.

ro u nd e d e nds ( B subtilis) ( Fig ) o r . . 10

F S ar i a
S q uare e nds ( B act anthraci
IG 9 cn ve n
s) ( )
-

F ig
. .

I I
t i l i Aft r S h m idt
. . .

rc . e c

an d W i e s Th e m e mb e rs o f this f o r m typ e vary

.

gr e atly in l e ngth Th e l e ngth vari e s f ro m to 8 mic ro ns and

. 1 0 ,

th e width f r o m to 6 micr o ns ( se e i nf a S iz e ) S p o re s may r ,

.
18 MO RPH OL O GY
be fo und in ce rtain sp e ci e s o f bacilli at de fini te p e ri o ds (se e
i nf ra Sp o r e s)
,
The se spo r e s may o ccur in vari o us p o sitio ns in
.

th e r o ds changing ,

t he ir co nto u r
s o m e what S o m e .

bacill i may also

p o ss e ss flage lla o r ,

w h i p l i k e p ro
ce sse s wh ich ,

s e rv e as a m e ans
F IG B a ill btili Aft r
10 c us su s e
O f l o co m o tio n s e e (
. . .

i nf ra , lage lla) T o th o se o d shap e d bact e ria which d o no t

F . r -

p o ss e ss flag e lla M igula appli e s th e t e rm bacte ri um e g B ac t , . . .

FI G . 11. -
B a t ri
c e um anthra i C hain f rm ati n N t th q ar
cs . o o . o e e s u e d
en s on
th e e nl arg d
e ll Aft r S hm idt nd W i
ce s . e c a e s .
 F O RM AN D S T RU CT URE OF B AC T ERI A 19

a nthracis Th e mo til e sp o re pro ducing fo rms are call e d bacilli

.
-

by M igula as n o te d ab o ve N o t infre que ntly are bacilli

,
.

surro unde d by g e latin o us capsul e s e g B caps ul atus I t is ,

. . . .

quite co mm o n fo r so me bacilli to gr o w in chains and fo rm

lo ng thre ads unde r ce rtai n co ndi tio ns 6 g B act anthraci s o r ,
. . .
,

B sub til us in b e e f b ro th ( F ig s 10
. On car e ful e xa m ina .
,

tio n it will b e s e e n that th e l o ng thre ads o r filame nts are made

up o f i ndividual bac te ria altho ugh at first sight th e thre ad ,

may app e ar to b e ho mo g e n e o us thro ugho ut .

S pi i l l a
r
— B e sid e s th e co cci and bacilli th e r e is o ne o th e r
.

co mmo n gro up o f l o we r bacte ria kno wn a s th e S pi il l a Th e se r .

bacte ria as th e ir nam e indicat e s are spiral o r c o rkscre w

, ,

shap e d Th e y are co mp o se d o f ro ds no t unlike bacilli but

.
, ,

in additio n th e y are twist e d so as to make th e m app e ar as

S pirals . Th e spiral may b e a sh o rt o ne f o urth turn o r l e ss ,

( M icro spira c o mma) ( Fig .

o r may b e l o ng and p o ss e ss

s e ve ral turns ( Spirillum ru

brum) Th e name mi c o spi a
. r r

o r vi b i o h as b e e n appli e d to
r

tho se S pirilla whi ch are ve ry

1
S h o rt and mak e but a fracti o n
,

of a turn Th e M icro spira f f

.

r
’
t

co m m a Of AS l a t Ch OICI a 15
FI G . 12 .
— M i cro s pira co m m a . Af t er

fre que ntly call e d th e Vibrio o f Sc m h idt a nd We i' '

Asiatic cho l e ra Ko ch calle d this las t m e nti o n e d o rganism th e

.
-

Co m ma bacillus o n acc o unt o f its r e s e mblance to a print

”
,
2 0 MO RP H OL O GY

FI G . 13 . pli atili s (On l ft ) Spir ha t

S p iro ch mte c . e . oc e e

o b rm i ri Aft r S hm idt nd W i
e e e . e c a e s .

’
er s co mma wh e n stain e d Lo ng cl o s e ly co il e d spirals are .
,

call e d spi ro clue te s by s o me write rs ( Fig 1 A tru e s pi il l um . r

i s rigid and sh o ws its e lf

o n a slid e a s a s e mi circl e

o f spirals ( Fig 14) a .

s pi o chae te is a c o ntinu r

o us lin e o f S piral s w hi ch

o ft e n attain gr e at l e ngth ,

a nd a mi c o sp i ra o r vi b i o r r

is a S ho rt curv e d ro d ,

si mil a r t o a c o m m a .

T h e spirillum and Vibri o

a re rigid and th e S piro ,

chacte is usually a fl e xibl e

S piral .

Th e siz e o f a spirillum

va i e s fro m micro n to r
F IG .
4 1 S p iri l l m
. a g i m Aft r
u s n u ne u . e

2 0 o r 3 0 mi cro ns i n
.

S hm id t d W i
c an e s .
 F ORM AN D S TRU CT URE OF B ACTERI A I

l e ngth and fro m

,
to 4 mi cr o ns in width acco rding to ,

th e Sp e ci e s .

Th e spirilla a e m o til e u s ually p o ss e ssing flag e lla at o ne o r

r ,

b o th p o l e s Th e co mmo n me th o d o f l o co mo ti o n thro ugh a

.
°

m e di um is by r o tatio n o n th e l o ng axis atte nde d in th e case ,

o f spir o ch a e t e s by s o m e lat e r al m o ti o n .

T h e flag e lla o r flag e llum p r e c e d e s th e c e ll

wh e n in m o tio n A fe w mo til e spir illa

.

d o no t p o ss e ss any flag e lla .

M o st spirilla d o no t pr o duc e sp o r e s .

Th e m e th o d o f r e pr o ducti o n am o ng thi s

gro up o f bact e ria is by fi ssio n b e ing simi ,

lar in this re sp e ct to th e co cci and th e

bacilli .

Tri ch o b a ct e ri a o r Hi gh e r B a cte ri a .

T h e trich o bact e ria o hi gh e r bact e ria co n

r

sist o f filam e nts o f v aryI ng l e ngth Th e .

fil am e nt s sh o w b o th tru e and fal se b r anch

ing T h e fil am e nt can by sp e cial s tains

.

b e sh o wn to b e mad e up o f indi v idual c e lls

'

s e pa r at e d f ro m e ach o the r by d e finite F C lad thrix

IG
5 . 1 . o

se pta F re que ntly all th e in di vidual c e lls di h t m a S h w c o o . o

i g fal bra h
.

“ ”
n se nc
I n th e filam e nt a e s urr o und e d by a co m
r i g typi al f tri n c o

h b i Aft r
m o n sh e ath o r m e mbran e ( F l g
a cte r a
5) (
c o . e
s e e 1
S hm idt d W i
.

c an e s .

i nfra S h e ath s)
,
. C e rtain c e ll s at o ne e nd

o f th e fil am e nt a re us e d to attach th e fil am e nt to o th e r fi l a

m e nts and to v ari o us o bj e cts At th e fre e e nd o f th e fil am e nt

.
2 2 MO RPHOL O GY
th e r e are c e rtain o the r ce lls whi ch are se t aside fo r th e purp o se
o f r e pr o ducti o n Th e se ce lls a re call e d go nidi a o r co nidia
.

( F igs 6 and . 1 The re is the re fo r e a divi sio n o f lab o r o r

, ,

a diff e r e ntiatio n in functi o n amo ng th e ce lls

o f a filam e nt in thi s gr o up .

Th e m o s t highly o rgani z e d g r o up o f th e

high e r bacte ria is r e pr e s e nte d in S t re p

to th rix actin o myc e s a f o rm unde r th e ,

C hl am yd o b acte ria ce a: ( se e F ig . 17 , A) .

actin o myce s th e filame nts d o no t

I n th e

app e ar o rdi narily to b e co mp o se d o f s e pa

rate individual s Occasio nally h o w e ve r .
, ,

th e pr o t o plasm b r e ak s up i nt o r o ds similar

to bacilli Th e r e is t r ue branchi ng similar

.

to that in th e myc e lium o f a fungus ( F ig

C lad thrix
.

FIG .6 1 . o

hyali a S h wi g
n .
Th e filam e nts o ft e n pr o duc e small
o n

g idia f rm ati
on o on
r und b o dl e s ( go nl d a) W t I n turn
.

Aft r S h m idt h
0

o ro l
e d c an p ,

Wi e s .

duce ne w individuals The r e is o fte n .

a club shap e d sw e ll ing o n th e e nd o f a fil am e nt I n all

-
.

pro bability this is a de ge n e ratio n pro duct and no t an o rgan o f

”

re pr o ductio n as was f o rme rly tho ught by so me writ e rs Th e s e

,
.

ro und b o di e s are no t similar to S p o re s a s is sho wn by th e ,

staini ng r e actio n and p o we r o f r e sistance I t h as b e e n sug .

g e ste d that th e m e mb e rs o f th e actin o myce s gro up co n stitute

a co nn e cting link b e tw e e n th e lo w e r bacte ria o n th e o ne hand
.
, ,

since th e filame nts o ccasio nally br e ak up into bacilli like r o ds -

and th e fungi o n th e o th e r since branchi ng simi lar to that in

,

th e m ce lium o f a fun g us o ccurs .

 F O RM AND S T RU C TURE OF B AC TERI A 2 3

Th e r e a e s e ve ral mo dificatio ns in fo rm and functio n amo ng

r

th e hi gh e r bact e ria Th e m e mb e rs o f o ne g r o up ( B e ggio

.

to ace aa) a re fr e e fr o m attachm e nt and u sually m o v e by

FI G . 17 .
— Th e fo m r ati
g idia (A) Str pt thrix a ti m y aft r L h
on o f on . e o c no ce s, e e

m d
ann an N m ann
( B ) S tr pt thrix h m g
eu aft r M a ( C) (D)
. e o c ro o e na, e ce .
,

Cr thrix K hni aft r K pf ( C) m i g nidi ; (D) m g i di ( E)

e no u ana, e o cro o a a cro on a .

G nidia lik b di in B a t t b r l i
o -
e o es c . u e cu o s s .

d ul a tiv em o ve m e nts o f the ir pro to plasm S o me autho rs .

have ascrib e d to the m an undulating m e mbrane wh ich during

m o til ity alte r nate ly co ntracts and r e laxe s Th e r e is no .

c o mm n she ath a ro und th e ce ll s Th e ce lls co ntain g r anul e s

o .

o f s ul phur C lo s e ly r e lat e d to th e ab o ve gro up is th e gr o up

.

Thi o thrix Thi s gro up is similar in s tructure to th e B e ggi

.

a to ace ae but o ne e nd o f th e fil ame nt is attache d to so me

,
2 4 R L GY
MO P HO O

substance and at th e o th e r go ni dia are fo rm e d Wh e n th e .

r e pro ductive c e ll s (go nidi a) burs t o ut o f th e sh e ath and a re

thro wn o ff th e y usually are e ndo we d with flage lla t e mp o ra rily
,

and swim ab o ut f r e e ly I n still an .

o th e r gro up ( S phae r o tilus o r C lad o

thrix) th e r e is de ve l o p e d what is
kno wn as fal se b r anchi ng as co n “ ”

tra ste d with th e t r u e branchi ng o f

a fungus o r w e e d Th e app e arance .

i s d ue to th e e nd c e ll o f a filam e nt

di viding twic e Th e s e c o nd divi si o n .

push e s to o ne side th e pro duct o f th e

first di vi sio n and e ach ce ll go e s o n ,

F 8 — P i illi m g la
IG 1. . en c m dividing se parate ly p ro ducing th e
u ucu .
,

S h wi g th tr
o n bra h i g
e ue nc n
so call e d bra ch
“ ”
fa f g
o f hi g h r rd r
un us o e
n
o e (.
F ig
-
.

Aft r F i h re sc e .

F u r th e r d e scripti o n o f th e Trich o

bacte ria will b e give n in co nne ctio n with th e chapte r o n

classificatio n .

I nvo l u ti o n F o rm s and P l e o m o rph i s m Am o ng th e bacilli .

,

s o me spirilla and c e rtain tricho bacte ria th e re f re qu e ntly

, ,

o ccurs pr o t o pla s mic d e g e n e rati o n a nd m o dificati o n i n f o rm in

s o m e sp e cie s This d e ge n e ratio n o f th e p r o to plasm withi n

.

th e c e ll usually di s to rts th e c o nto ur o f th e c e ll m e mb r a ne ,

pro ducing what a e kn o wn a s invo lutio n fo r m s This

“
r .
”

d e ge n e ratio n o ccu rs wh e n th e bacte rial ce ll is no t pro vide d

with suitabl e fo o d is b e i ng subj e cte d to t e mp e rature change s
, ,

o r to pr e s e nc e o f a sup e rabundanc e o f e xcr e t o ry pr o ducts .

 F O RM AN D S T RU CTURE OF BACT ERIA 2 5

Th e co cci do va y p e rce ptibly unde r any co ndi

no t s e e m to r
‘

ti o ns I t is p o ssibl e to cause a r e ge n e rati o n o f th e se inv o l u

.

tio n fo rm s in ce rtain sp e ci e s whe n favo rable co nditio ns are

pro vi de d I t i s no t p o ssibl e to cause any p e m ane nt change i n
. r

the m o p ho l o gy of a bacte ri um a nd thus o i gi na te a ne w sp e ci e s

r r

of bacte i a
r F o r e xampl e it is no t p o s sibl e to caus e a bacillus
.
,

to chang e i nt o a c o ccus and vi ce ve sa This ide a o f pl e o r .

mo rp hi s m was h e ld fo r y e ars by c e rtain bact e r i o l o gists (Z o pf .

cl Th e y e xplain e d th e o igin o f ne w sp e cie s by m e an s o f

r o

thi s phe no m e n o n P l e o mo rphism do e s no t e xi st in its true

.

s e ns e ; that is the re is no p e rmane nt change in fo rm o f a

,

d e finite sp e cie s o f bacte ria Th e bacte ria if m o di fi e d fr o m

.
-

th e ir n o rmal shap e will whe n supplie d with th e r e quisit e

,

c o ndi tio ns o f te mp e ratur e fo o d e tc as b e fo re state d r e turn

, ,
.
, ,

to th e no rmal Th e r e may b e a change in th e cha r acte ristic

.

fo rm o f gro wth o f s o me bacte ria unde r ce rtain co n di tio ns .

B acte rial ce lls may fo rm them se lve s i nto p e llicle s o Ogl oe a ,

z ,

chains fil ame nts e tc but th e typ e o f c e ll r e main s th e sam e

, ,
.
,
.

Chl o rid e o f lithium and c e rtain o th e r ch e mical s wh e n plac e d

, ,

in a m e dium co ntaining bact e ria will cause a m o di ficati o n o f ,

the ir shap e fro m th e n o rmal .

I n c e rtai n S p e ci e s o f ba ct e ria th e i ndividual ce ll s a re m o r e

pr o n e to change the ir shap e unde r varyi ng co nditio ns than

o th e rs e g B act
,
. . diphthe ri ae B act tub e rcul o sis bacte ria in
.
,
.
,

th e r o o t tub e rcl e s o f cl o v e r e tc ( F igs 19 and Vari o us . .

sp e cie s o f bact e ria pro duce invo luti o n fo rms unde r di ff e r

e nt co nditi o n s Fo r e xampl e th e ac e tic lactic butyric acid
.
, , ,
2 6 MO RPHOL O GY
pro ducing bacte ria S ho w gr e at disto rtio n in shap e wh e n th e
acids pro duce d are incr e ase d ab o ve a ce rta in p e rce ntage .

th rs
O e S o h w a chang e in shap e wh e n t e mp e ratur e is k e pt ab o v e

FI G . 19 v l ti n
—.In B btili wat r ba t ria B a t
o u o Ba t
. su s., e c e , c . c .

p t i n m ba t
as e ur a u id fr m r t n d l B a t t b r l i B a t
, c e ro s o oo o u e s, c . u e cu o s s, c .

diphth i Al l f rm h wn
era . v ry difl nt f m th n rm al f rm
o s s o are e e re ro e o o s .

Aft r F i h r
e sc e .

th e n o rmal fo r any p e ri o d o f ti m e and o the rs S ho w change in ,

shap e wh e n fo o d is lacking and e xcr e to ry pr o ducts have

incr e as e d to a high p e rce ntage To o rapid r e p ro ductio n may .

cause ab e rrant S hap e s amo ng s o m e sp e cie s Th e co n di .

tio ns whi ch pro duce invo lutio n fo rms amo ng bacte ria also
 F O RM AND S T RU CT URE OF B ACTERIA 2 7

atte nuate th e bacte ri a as far as the ir ability go e s to pro duce

fe rme nta ti o n putr e factio n di se ase e tc
, , ,
.

An o th e r th e o ry th e th e o ry o f pl e o go ny was also fo .

m e rly he ld by ce rtain bacte rio l o gists This th e o ry de alt with .

FI G . 2 0. v l ti
In o u o n fo rm
s . Ba t t b r l i
c . u e cu o s s, two m o n th ’
lt
s o l d cu ure . Af tr
e

S h m idt
c d W i an e s .

th e de ve lo pme nt o f ne w physi o l o gical charact e ristics amo ng

bacte ria Th e typical n o rmal and physio l o gically active
.
, ,

bacte rial ce ll wil l ho w e ve r alway s r e tu rn whe n th e me ce s

, ,

sary co ndi tio n s are p ro vid e d Th e passage o f so me bacte ria .

thro ugh ani n al s cau se s a re turn to th e n o r mal virul e nce and

l

typical fo rm o f th e bacte rium .

Dim e ns i o ns o f B a cte ri al C e l l s — Th e S ize vari e s with th e .

sp e cie s I n m o st instance s it is fairly co nstant withi n th e

.

li mit o f th e sam e sp e ci e s I n so me sp e cie s th e size o f th e

.
2 8 MO RPH OL O GY
bacte rial ce lls vari e s as gr e atly as do e s th e S hap e Within .

th e s am e sp e ci e s a variati o n in c e rtain c e lls o f and in o th e rs

to 3 o r 5 micr o ns in l e ngth has b e e n n o t e d Te mp e rature .
,

r e acti o n and co mp o siti o n o f th e m e di a a nd in additio n to

,
, ,

th e se th e age o f th e c e ll are r e sp o nsibl e fo r th e s e variati o ns

, .

10

FI G . 2 1
. parativ iz f ba t ria ( ) B a t infl
—
Th e co m e s e o c e . 1 c . ue nz ee .
(2 ) Ba tc .

t b r l i ( 3 ) B typh
u e cu o s s (4 ) B a t a thra i
. .
(5 ) M o sus . c . n c s . .
go no rrh ce aa . ( 6)
M py g . var a ro ( 7 ) M f d m i hii ( 8) M p
e ne s . u e us . . re u e e c . s . co mm a .
( 9 ) Sp .

und l ( ) S p b rm i ri
u a . Io . o e e e .

Th e s ize ce ll s di ffe rs wh e n intro duce d into animals and

o f th e

plants and o n th e vario us cultu r e m e dia u se d fo r th e study

o f bact e ria I t d o e s no t s e e m that b r e adth i s a ny m o r e co n
.

sta nt than l e ngth amo ng sp e cie s o f bacte ria which vary in S ize .

I t i s i nt e r e s ti ng to no t e that o ne o f th e small e s t o bs e rv e d

bacte ria P s e udo mo nas indigo fe ra ( Vo g e s) M ig is o nl y

, .
,
 F ORM AND S T RU CTURE OP B ACT ERI A 2 9

micro n in l e ngth and rm e r o n in width B acillus .

d e ntrifi cans M ig is a ro d 1 to . micr o ns in l e ngth by

to micro n in width Th e M i cro co ccus p ro g e die ns
.

“
r

( S ch rOte r) is mic r o n in di am e te r A bacte rium d e .

s crib e d by N o ca r d a nd R o uxi n pl e u ro pn e um o ni a i s s aid to

b e s til l s m all e r Witho ut do ubt th e r e a e s till small e r o rgan

. r

i sms which a e ul t am i crOS COpic S o me o f th e la rge st

r r .

bact e ria have b e e n r e p o rt e d by E rr e r a and S ch audi n Th e y .

d e sc rib e a spirillum o f th e dime ns io ns 2 3 to 8 micr o ns in 2

l e ngth by 3 to micro ns in width and also a bacillus 2 4 ,

to 8 mi cr o ns in l e ngth a nd 3 to 6 mi cr o ns wid e
0 S o m e v e ry .

la rg e co cci hav b e e n r e p o rt e d by vario us write r s

e .

F i sch e r mak e s th e e s timat e that it w o uld tak e

pus co cci to w e igh o ne g r am I n a dro p o f wat e r o ne cubic .

c e ntime te r in size pus co cci wo uld have pl e nty

o f spac e B uj wid c o u nt e d th e numb e r o f M icro co ccus
.

pyo ge ne s var aur e us ce ll s in a mi lligram o f a pure cul tur e and

.

f o und that the re we re appr o ximate ly F isch e r

state s that if an anth rax bacillus is 3 to 0 mi cr o ns l o ng and 1

1 to micro ns b r o ad that at l e ast wo uld b e

, ,

r e quir e d to e qual a mass th e size o f an o rdinary ciga r e tte .

N ag e li give s th e w e ight of an ave rage bacte rium as

a 10 000 000 000 th part o f a milligram
, , , .

Lo co m o i o t n of B a cte ri a
F l age l l a . C e r tain p e ci
s e s ofbacte ria p o sse ss th e p o we r
o f l o co mo tio n . Th e de g re e o f m o ti o n d e p e nds up o n th e a ge
3 0 M ORP H O O L GY
of th ebacte rium a nd th e t e mp e ratur e o f th e surr o unding
m e di um Th e fa cul ty o f mo vi ng ab o ut f ro m place to place
.

is p o sse ss e d by o nly a fe wvari e ti e s o f co cci e g M icro co ccus ,

. .

agi l is e tc
, but is quite co mmo n amo ng th e bacilli a nd
.
,

S pirilla M o til e bact e ria p r o gr ss by m e a ns o f th e vibrati o n

. e

o f l o ng whiplik e app e ndag s call e d flag e l la o r cilia

,
e .

Ehr e nb e rg was th e first to d e s crib e flag e lla o n bacte rial

ce lls I n 183 8 h e de sc r ib e d appe ndage s o n B t r il o cular e

.
,
. .

I n 187 0 Co hn d e scrib e d flag e lla o n S pirillum o l uta ns

,
In v .

1 87 5 Warming d e s c rib e d flag e lla o n S pirillum und ul a Sp iril

, ,

lum rugul a e tc I n 87 7 vario us s tains w e r e us e d to d e m

,
. 1 ,

o nst a te th e ir pr e se n ce
r B y m e an s o f s tains it wa s f o und
.

that a large numb e r o f bacte ria p o ss e ss e d th e se o rgans o f

l o co m o tio n B y th e ye ar 880 all bacte rio lo gis ts had ag re e d
. 1 ,

that th e re w e r e app e ndage s o f this char acte r o n ce rtain

bacte rial c e lls Th e m e tho d o f lo co mo tio n by flag e ll a is
.

use d by practically all m o til e lo w e r bact e ria Amo ng s o m e o f .

th e hi gh e r bact e ria th e e xi s t e nc e o f a n undul ati ng m e mbran e

i s claim e d in c e r tai n sp e ci e s e g B e ggiao to a I t i s s upp o s e d

,
. . .

that th e s e Sp e ci e s m o ve by th e alte rnate co ntractio n and e r

laxati o n o f th e ir u ndul ating me mbran e .

Th e flag e lla o n a bact e rium va ry in n umb e r a nd arrang e

'

m e nt acco rdi ng to th e Sp e cie s The y a e usually quit e co n . r

stant in numb e r fo r th e sp e cie s Th e flage lla may vary n . I

p o sitio n o n th e di ff e r e nt bact e ria and M e sse a in 890 , ,

1 ,

pro p o se d dividing th e fl age l l ate bact e r ia into th e fo ll o wing

class e s
 F O RM AN D S TRU CT URE OF B AC T E RI A 3 1

( ) M o1no trich o u s o ne flage llum at o ne e nd ( Ex P s e ud

.

o m o nas p yo cyane a)

( F ig .

( ) 2 ph o trich o us Lo

s e ve r al flag e lla at
o ne e nd in a bunch

( E Spirill us rugul a)
x .

( ig
F .

( 3 ) Am phit richo us
flage lla at b o th
e nd s o ne at e ach ( E
,
x .

B a a nus b utyn cus)

0
F 10 . 2 2 .
— P d se u a
o mo n s p yo cy ane ah wing m n
. S o o o

tri ch o us fl ag e lla ti o n . Aft r S hmi d t nd W i

e c a e s .

( Fig .

( 4) ritricho us flage llaPe o n all side s of th e ce ll ( Ex .

B acill u s typho sus ) ( Fig .

pr e s e nce o f amphi
Th e

tri ch o us bact e ria hav e .

b e e n que stio ne d by so me .

Th e s e inve stigat o rs ho ld
that a n amphit richo us
bacte rium is fo rm e d fr o m
a m o no tricho us bacte r
ium whi ch is acquiri ng a
ne w flag e llum pr e para

to ry to fi ssio n .

FI G . P dm a y
2 3 .
— se u o o n s s nc a y n S h wi g
ea .
I n 89 4 M igula use d
o n 1
l ph tri h fl g ll ti
o o c o us a e a on . Aft r S hm idt
e c
,

an d W i e s .
th e numb e r o f flag e lla o n
3 2 MO RP HO L O GY
a bacte rium and th e ir m o de o f attachm e nt as a basi s fo r di stin
g ui shi ng a n d s e parati ng vari o us g e ne ra F is ch e r in 895 .
,
1 ,

us e d th e flag e lla as m e an s of distingui shi ng subfamili e s .

Th e flag e lla a re l o ng

de licat e p ro ce sse s usu ,

ally o n e t o tw e n ty
tim e s l ng e r than th e o

bacte rial ce l l ( to 1

2 0 X) The y p o sse ss .

a base whi ch is slightly

large r th n th e r e s t o f a

th e pr o c e ss Th i s base .

tap e rs like a co n e with ,

a l o ng filame nto us p ro
— S pirill m r br m
F IG 4 . 2 u u u ce ss co ming o ff th e
tri h fl g l l ti Aft r S hm idt d W i
c o ns a e a on e c an e s
ap e x Th e m o de o f
. .

attachme nt o f th e flage llum to th e bact e rial c e ll is no t as ye t

de finit e ly s e ttl e d I t i pro babl e that th e pro to plasm o f th e
. s

bacte rial c e ll is co ntinu e d o ut int o th e flage lla and that th e re

is a thi n l imiti ng m e mb r a ne o n th o ut id e similar to al
.

e s , ,

th o ugh much thinn e r than th e ce ll wall p ro p e r B y th e use o f

,
.

ze rtain r e ag e nts th e pr o t o plasm o f th e flag e lla h as b e e n s e e n to

shrink away f r o m th e limiting m e mb r ane and a n attachm e nt ,

has b e e n n o te d o f th e flag e llum to th e p e riph e rial zo n e o f

pro to plasm o f th e c e ll S o me write rs h o ld th t th e flage llum
. a

is de finit e ly s e parat e d fro m th e ce ll wall and co mp o se d o f

di ffe re nt mate rial S o me have also h e ld that th e flage ll a
.
 F ORM AN D S T RU CT URE or B ACTERIA 33

a re me re ly app e ndage s o f th e ce l l wall and no t o rgans o f l o co

m o tio n I t may b e he r e state d that fl ag e lla have n e ve r b e e n
.

de mo n strat e d o n bacte ri a de vo id o f m o tility and are always

FI G . 2 5
.
— B . p ritri h
e c o us fl age ll a ti o n
. tr
Af e Sc mh idt
a nd W ie s .

fo und o n mo til e bacte ria Th e pro to plasm o f th e fl age l

.

lum may b e slightly diffe r e nt che mically f r m th e r e st o f th e o

bacte rial ce ll fo r it is a familiar fact that th e flag e lla a e ve ry

,
r

much mo re liabl e to h e at ch micals e tc than th e p r o to plasm o f

,
e ,
.
,

th e c e ll it se lf I t is p o ssibl e that th e flag e lla are co n ne ct e d with

.

th e c e ll thr o ugh p o r e s whi ch c o nn e ct th e int e r nal structur e s o f

th e two Und e r unfavo rabl e c o n di ti o n s th e flag e lla are thr o w n

.

o ff are di sint e grat e d and c o mpl e t e ly disapp e ar in a sh o rt tim e

, ,
.

D
34 MO RP HOL O GY
Th e flage lla o n a bacte rial ce ll are v e ry hard to de mo nstrate .

I t is o nly by th e m o st car e f ul t e chniqu e a nd th e use o f sp e cial

m o rdants that th e y can b e r e ve al e d Th e y canno t b e s e e n

.

de finite ly in unstai ne d pre pa r atio ns o r pre pa r atio ns stain e d

in th e o rdi na ry manne r Th e p ro to plasm o f th e flage lla do e s
.

no t r e adily e nt e r i nt o ch e mical c o mbi nati o n with th e o rdi nary

bacte rio l o gical stains Thi s p o ints to th e fact that its p ro

.

to pl asm i s di ff e r e nt fr o m that c o ntain e d within th e c e ll Th e .

flage lla may b e o fte n r e ve al e d by th e de p o sitio n o f stai ns

(AgNO e tc ) o n th e o ut side o f th e limiti ng m e mb r an e
g, . .

M any bact e ria di sca r d th e ir flag e lla wh e n th e y a re r e m o v e d

fro m th e m e dium in which th e y a e gro wing C o nse que ntly
r .

th e y a e ve ry ha r d to place intact o n a slide The y a re ve ry

r .

s e n sitive and th e flage lla are ve ry e a sily di inte grat e d during s

th e pr o c e ss o f staining . I t has b e e n no t e d that th e s taining

p o we r o f a flage llum o fte n vari e s with th e S p e ci e s o f bacte ria .

M o rdants a re u se d with th e sta ins in o rd e r that th e flag e lla

may b e de mo n strate d at all .

AS a fl age l l ate bact e r i um m o v e s it pro gr e ss e s in a d e finit e

manne r Th e m o no and l o pho tricho us bacte ria mo ve with
.

th e flag e llum o r flag e lla in fr o nt lik e c e rtain infu so ria and ,

r o tate th e s e o rgans in a ci r cular fashi o n in a di r e cti o n l o ngi

tudi nal to th e l o ng a xis o f th e c e ll Th e c e ll its e lf in c e rtain
.
,

S p e ci e s
,
ro tate s fr e e ly whe n in m o tio n Th e p e ritrich o us
.

bacte ria m o ve in a l o ngitudinal dir e ctio n as th e o th e r bact e ria

and in addi ti o n may p o sse s s an e nd o ve r e nd m o ti o n tumb
- -

ling simil ar to a s o m e rsault .

 F O RM AN D S TRUC T URE OF B ACT ERI A 35

p
R a id i ty f
o M o veVi e w e d unde r th e micro sco p e th e
m e nt .

mo tile bacte ria se e m to mo ve ve ry swiftly Thi s ide a is o f .

,

co urs e d ue to th e limite d fi e ld o f vi sio n F ische r e s timate s

,
.

th e actual av e rag e rat e o f m o v e m e nt und e r n o rmal cO ndi tio nS

o f t e mp e ratur e e tc at te n c e ntim e te r s in fifte e n minut e s o r

,
.
, ,

ab o ut millime t e r p e r se co nd I t S ho uld b e n o te d that th e.

S p e e d is c o mparativ e ly rapid taking into co n side ratio n th e

,

small S ize o f th e bact e rial ce ll Th e rapi di ty o f m o ve me nt o f a

.

m o tile bacte rium varie s with th e physio l o gical co nditio n o f

th e c e ll Th e t e mp e ratur e pr e se nce o f nutri e nt mat e rial
‘

.
, ,

pr e se nce o f e xcre to ry pro duct s and th e ge n e ral vitality o f th e

,

c e ll have much to d o with th e characte r o f th e m o ve m e nt o f

th e c e ll .

B ro wni an M o ve me nt Ve ry fre que ntly wh e n pr e parati o ns

.

o f l iv e bact e r ia are e xamin e d u nd e r th e micr o s c o pe th e y a re

'

s e e n to mo ve backwa r d and fo rward in an o s cillating fashi o n .

The y d o nOt pro gr e ss th ro ugh th e me di um Thi s mo ve m e nt .

sho ul d b e care ful ly diff e r e ntiat e d fro m true lo co mo tio n This .

phe no me no n which wa s firs t de s crib e d by B ro wn is d ue to

, ,

th e b o mbardm e nt o f m o l e cul e s in th e li q ui d m e dium i n which

th e bact e ria may b e Susp e nd e d ; P articl e s o f I ndi a ink S h o w

th e sam e so rt o f m o ve m e nt .

C aps ul e s S o m e S p e ci e s o f bact e ria s e cre t e a g e latino us

.

sub stance fro m th e o ute r laye r s o f th e ce ll p ro to plasm o r th e

c e ll wall Wh e n larg e numb e rs o f bacte ria a re fo und to ge th e r
.

se cr e ti ng th e se g e latin o u s sub s ta nc e s in gr e at am o unt th e

typical zo o gl ce a is fo rme d Ho we ve r th e bacte ria may o c

’ '

.
,
3 6 MO RP HOL OGY
cur s e parat e ly and b e co me surro und e d by a thi n she ath o f
a cle ar j e llylik e highly r e fractil e sub s tance which is usually
, , ,

FI G . 2 6 C ap lsu e s . Ba t
c .
p ne um o ni cum ( Fri dla d r)
e n e . Af t r W i h lba
e e c se um .

many tim e s br o ade r than th e bact e rial ce ll To this S he ath .

o f g e lati no us mat e rial th e nam e ca p s ul e h a s b e e n appli e d .

C ap s ul e pr o ducing bact e r ia d o no t pro duc e capsul e s und e r all

-

co ndi tio ns Th e n e ce ssary and particular nutri e nt mate rial

.

must b e availabl e fo r capsul e f o rmatio n B acillu s capsulatus .

in milk pro duce s th e mo s t typical capsul e s whil e o n agar agar ,

-

it pro duce s n o n e B acte rium anthracis o n o rdi nary cultur e

.

m e dia such as ge latin agar agar be e f bro th e tc pro duc e s

,
-

, ,
.
,

no capsul e s whil e in th e bl o o d o f an animal inf e ct e d with a n

,

thrax th e capsul e s are v e ry typical S o m e time s se ve ral bac .

te rial c e ll s app e ar to b e surr o und e d by a singl e capsul e e g M ic ,

. . .

pne umo nia I n al l pr o bability thi s app e aranc e is d ue to

.

th e amalgamati o n o f th e capsul e s o f th e s e v e ral c e l ls I t is .

 F ORM AN D S TRUCT URE OF B ACT ERI A 37

usuall y n e ce ssary to use sp e cial stains to d e m o nstrate capsul e s

'

cl e arly F r e que ntly o n stain e d slide s cl e ar space s a e s e e n

. r

a r o und th e bacte rial c e ll and th e se Space s are o fte n co nfus e d

,

with capsul e s Th e spac e s a e a r tifacts d ue to th e co nt r actio n

. r

o f th e g e latin o u s cap s ul e during th e pr o c e ss o f drying M ig ul a .

says that in all pr o bability all bacte ria p o sse ss capsul e s in a

rudi m e ntary s tate .

S h e ath s . Am o ng c e rtain f o rms o f th e trich o bact e ria

( Clado thrix C r e n o thrix e tc ) th e re is d e ve l o p e d a structur e

, ,
.

whi ch h as a ve ry sim ilar o rigin to that o f a capsul e Th e .

o ut e r lay e rs o f th e c e ll wall inst e ad o f r e mai ni ng s o ft and ge

,

l atino us b e c o m e hard and fo rm what is call e d a she ath

,
Pho s .

phat e o f lime is de p o site d in th e S he aths Of so me sp e ci e s o f

bacte ria Th e sh e ath s o f bacte rial c e ll s in a ch al n o f filam e nt
.

fr e que ntly b e co m e co nn e cte d t o g e th e r making th e thr e ad o f

,

c e lls v e ry rigid Th e c e ll pro p e r co ntracts away fro m th e

.

sh e ath and is f r e e to mo ve Wh e n r e pr o ductio n tak e s place

.

th e S h e ath s br e ak o p e n and th e individual c e lls e scap e S ub .

s e que ntly th e sh e ath disi nt e grate s ( se e Fig .

M as s G r o u ping (a) Z o Ogl oe a ( b) P e ll icl e s and (c) Co l o

.
, ,

ni e s N o t infre que ntly bact e rial ce lls are fo und co lle cte d
.

t o ge th e r in aggre gate masse s Th e s e ma ss e s may have no.

de finite shap e and th e in di vidual bact e ria are h e ld to g e th e r by

,

a g e latino us se cr e tio n Th e indi vidual bacte ria may b e fo rme d

.

into de fi nit chains Th e ge latino us mate rial is e ithe r se cr e te d

e .

by th e ce ll wall o r th e o ute r lay e rs o f th e c e ll pro to plasm I n .

all pr o babili ty it o riginate s fro m th e fo rm e r Th e incre ase d .

3 8 MO RP H O L O GY
abso rptio n o f wate r by th e o ute r lay e r o f th e ce ll wall s e e ms to
b e r e sp o nsibl e fo r th e g e la tin o us s e cr e ti o n T h e t r m zo ogl aea . e

is applie d by bacte ri o l o gists to the se masse s o f bacte ria and

ge latino us mate rial e g B v ul ga tus This f o rmatio n is ,
. . . .

n o te d o n th e surface o f so lid and liqui d nutri e nt substance s .

Th e g e latin o us mat e rial s e cr e t e d i s S imila r to that whi ch

fo rms th e capsul e s o f c e rtain sp e cie s o f bacte ria ( Fig .

I t c o mp o se s a thick co v er

ing o n th e o utside o f th e
b ac t e ri al ch ai n s Th i s .

f o rmati o n is ve ry o fte n se e n
in th e pro c e sse s o f fe rm e n
t a ti o n Of S t r e p t o c o c cu s

m e s e n t e ri o i d e s in s uga r

F G 7
I . 2 Z g l a i n aq ati b t
. oo
vat
oe
s
na
O th e r clumps o f bac
u c ac e .
.

i m
r u T h i divid al
. ll e thi kly te ria which a re no t r e adi ly
n u ce s are c

aggr g at d at th p riph ry l
e e i e e e e s s so n
h ld t g th r by S o lubl e I n wa ter a nd Wt h
,

th m iddl ; th y
e e e a re e o e e

am lik r ti Aft r F i h r ari se f ro m fu si o n o r imp e r

ucus -
e se c e on . e sc e .

fe ct s e parati o n o f th e individual c e ll s a e O ft e n se e n Th e se r .

mass e s a e call e d pse ud z ogl oe a Th e y d o no t p o sse ss th e

'

r o o .

c o mpactn e s s and ge latinizatio n o f th e t r u e zo o gl oe a Z o Ogl oe a .

a nd p se ud o o Ogl oe a ma s s e s c o mp o s e th e s o ca ll e d s lim e whi ch

z -

c o lle ct o n vari o us fluids I n nutri e nt liqui ds ce rtain bacte ria .

fo rm aggre gate s pro ducing th e app e arance o f scum o n th e sur

fa ce e g B subtilis The se aggr gate s a e so m e time s call d
,
. . . . e r e

p e l l i cl e s Th e p e llicl e usually gro ws until it c o ve rs th e surface

.

o f th e l iqui d Th e y co ntain in additio n to th e bact e ria and

.
,
 F O RM AN D S T RU C TURE OF B ACT ERI A 39

th e ge latino us s e cre tio n s o m e o rgani c mate rial de rive d f ro m ,

th e m e di a Th e Sl z e o f th e p e llicl e varie s with th e S ize o f th e

.

FI G . 2 8 . Typ f ba t rial l i (A) C hl at (B li ab rm al f rm )

es o c e co o n e s. oc e e . co , no o .

(B ) C o n gl m rat (B p fi ) ( C) Am b id (B l g t ) ( D) R hiz id (B m y
o e e . zo . e o . vu a us . o .

co id e s) .
( E ) C rl d (B a t anthra i ) (F ) M y l id ( B radiat )
u e c .
(G) F ila cs . ce o . us
Aft r Fr t
.

m e nto us . e os .

v e s se l co ntai ning th e nutri e nt fluid C are S ho uld b e tak e n .

no t to c o nfu se th e s e mass e s o f bact e ria i e zo Ogl oe a ps e ud o ,

. .
,

zo Ogl oe a a nd p e llicl e s wi th th e r e gular co l o ni e s o f bact e ria

,

FI G . 2 i r pi tr t r f l i (A) C l y a wh l
9 . M c o s co c s uc u e o co o n e s . o on as o e . r
( 1) A e o
lat ( ) G m
e .
( 3 ) M r l id (4 ) C l d d ( 5 ) G yr
2 ru o se . o uo . ou e . o se . ar
( 6) M m o
r at d ( 7 ) R ti lat (B ) E d g f l i ( 8) R pa d
e . e cu e . e o co o n e s . e n .
( 9 ) L bat o e .

( ) Er
10 ( ) A i lat ( ) L a rat
o se .
( 3 ) F im bri at
11 ur cu e . 12 ce e . 1 c e .
( 4 ) C iliat
1 e .

Aft r Fr t e os .
40 M ORP H O O L GY

FI G 3 0 . . tr ak lt
( A) Typ( ) Filif m (B
es of s li) ( ) E hin lat
e cu ure s . 1 or . co . 2 c u e

(B a t a idi l ti i )
c .
( 3 ) B ad d ( S t py g n )
c -
ac c (4 ) Efl
.
(B lg t )
e e r . o e es . use . vu a us .

5 Arb r. nt ( B M y
o id )
e sce . co es .

( B ) Typ f gr wth i tab lt r N nl iq fyi g ( ) Filif rm (B li)

e s o o ns cu u es . o ue n I o . co .

( ) B ad d ( S t py g
2 e e ) (3 ) E hi at ( B a t a idi l ti i) ( 4) V ill
r. o e ne s . c n e c . c -
ac c . o us

( B a t m ri pti m ) ( 5 ) Arb r
c . u se t cu Li q fyi g ( 6) C rat ri
. o e sce n ue n : e

f rm ( B v l g ari
o 4 . ( 7 ) N p if m ( B
u s, btili 48
2 ( 8) I f di a or . su s, n un

b l if m ( B p digi
u or ) ( 9 ) S a at (M p fi nkl i )
. ro ( ) S tratif rm ( P
o sus . cc e s . er . 10 o S .

fl n )
uo re sceAft r Fr t s . e os .
 F ORM AN D S T RU CT URE OF B ACTERI A 4 1

d e ve lo p e d in nutri e nt m e di a as a r e sul t o f r e pro ductio n in a

de finite manne r Th e typ e o f co lo ny is a characteristi c fo r a
.

de fi nit e sp e cie s and is no t a n accide ntal fo rma tio n Th e typ e

.

o f c o l o ny is o f tax o no mic im p o rtanc e that is o f i m

,
p o rtance
in id e ntifying a de finit e sp e cie s ( F ig . T h e e dg e s o f th e

co l o ni e s may b e re gular smo o th o r s e rrat e d ( Fig

, ,
. Dif
fe re nt d e scriptiv e t e rm s a re appli e d to th e charact e ri s tic fash

io n in which bact e ria gro w in c o l o ni e s . Whe n bacte ria are

ino culate d o n th e surface o f agar tub e s o r by puncture into
ge latin o r agar th e vario us sp e cie s gro w in a ve ry co nstant and
,

typical mann e r ( F ig .
 C HAP T ER I V

RE PR ODUCTI ON OF B AC T ER I A
In tro d ucti o n Th e pr o ce ss o f r e pr o ductio n amo ng bacte ria
.

is e xce e dingly inte r e sting I f pro p e r fo o d and e nviro nme nt are

.

pre s e nt th e bacte rial ce ll mul tipli e s at a c e rtain stage o f its

,

e xist e nc e R e pr o ducti o n i s e ntir e ly a se xual

. Th e t e rm re .

pr o ductio n as it is u sually appli e d no t o nly include s th e

, ,

pro ductio n o f ne windi viduals but also an i ncr e ase in th e num

b e r o f individuals Th e pr o c e s s i s slightly di ff e r e nt in th e
.

l o we r and th e highe r bacte ria and co ns e que ntly re pro d uc

, , ,

ti o n will b e co nside r e d s e parate ly fo r e ach cla ss .

Re p r o d u cti o n am o n g th e H apl o b a cte ri a ( Lo we r B acte ri a )

F i ssi o n . m e tho d o f re pr o ductio n amo ng the s e fo rm s

Th e

is by binary di visi o n o r fi ssi o n Th e c e ll ab o ut to multiply

,
.

e l o ngat e s slightly and gradually b e c o m e s c o nstrict e d in th e

c e nt e r fro m all side s in transve rs e di am e te r to th e l o ng axis

o f th e c e ll ( F ig Th e c o n s tricti o n b e c o m e s d e e p e r and
.

de e p e r carrying in th e chitin o us o r ce llul o s e lik e wall and finally

,
-

th e two parts o f th e o riginal c e ll r e main attach e d by o nl y a

thin thre ad B acte rial ce ll s fo ll o w th e sam e laws o f c e ll di

.

visi o n a s m o st animal and plant ce ll s Th e ce l l usually ih .

42
 REPR ODUCTI O N OF BAC TERI A 43

cr e ase s in size to th e n o rmal adul t th e n divide s by fi ssio n as th e

,

pa re nt did N o t infr e que ntly fissio n take s place b e fo r e th e

.

Q GD00 (o)
FI G 3 1
. . iagram il l trati g fi i n f ba t ial
D us n ss o o c er ce ll (ba illi
s c and co cci ) .

Aft r N vy e o .

adult stage is r e ach e d I nvo lutio n f o rms may b e pro duc e d

.

by thi s m e ans B e fo r e any co n stricti o ns app e ar in th e ce ll

.

wall th e p ro to plasm within sho ws e vide nc e s o f b e ing c o nst ricte d

i nt o two parts So me write rs claim that a m e mbrane f o rms
.

at th e p o int o f divisio n which se parat e s th e par e nt ce ll into

two di s tinct parts Thi s in all pro bability is no t always th e
.

e xact c o nditi o n altho ugh in so m e sp e ci e s o f bact e ria an un

,

stain e d line do e s app e ar in th e pr o t o pla sm at th e p o int o f di

vi sio n I t is claim e d by s o m e that thi s dividi ng m e mbran e
.

is ce llul o se but r e ce nt r e se ar ch sho ws that no such sub

,

stance as th e ce llulo se o f high e r plant c e ll s o ccurs amo ng th e

lo we r bacte ria B acilli and spi rilla always di vide in a di e c
. r

ti o n tran sve rse to th e lo ng axi s o f th e ce l l Th e co cci di vide .

irr e gularly as re gards a de fini te dire cti o n .

R a te of M ul ti pl i ca ti o n A bact e rium may r e ach th e adult

.

fo rm and di vide in fro m twe nty minute s to an h o ur varyi ng ,

with th e S p e ci e s B act e ria a e v e ry p r o lific wh e n th e r e qui

. r

sit e co nditio ns o f fo o d te mp e ratu re and gase o us e nviro nme nt

, ,
44 MORPHOL O GY

a re at hand One bacte rium dividing o nce e ve ry h o ur in

.

twe nty fo ur ho urs w o uld p ro duce if uninhibite d

-

individual s in f o rty e ight h o urs

,
-

5 6 in d ivi duals ,

a nd in S i days th e mass pr o duce d if e ach bact e rium w e igh e d

x ,

o n th e av e rag e o f fi m o f a mil lig r am w o uld e qual ,

l

th e siz e o f th e e arth Uni nt e rrupte d divisi o n fo r any p e ri o d

‘

o f tim e b e y o nd o r e v e n to tw e nty f o ur h o urs e xc e pt in rar e

-

cas e s is an abso lute imp o s sibility Th e ge o m e trical p ro

,
.

g re S S
I O n s n o t an
I actual o ne in th e cas e o f bact e ria L o n g b e .

fo r e th e ce ll has multiplie d to th e e xte nt give n in th e ab o ve

figure s th e fo o d will have b e co m e so r e duc e d and e xcr e to ry
, ,

pro duct s will have accumul at e d to th e e xte nt o f acting as

po iso ns to th e c e ll s th e mse lve s and co nse que ntly inhi biting
fissio n so that r e pro ductio n wil l pro gr e ss much S lo w e r than
,

is p o ssibl e th e o re tically and will finally c e as e co mpl e te ly N o t .

e v e n in an inf e ct e d b o dy d o bact e ria multiply u ni nt e rrupt e dly .

I t is int e r e s ting to o bs e rv e h o w e ve r what unl imit e d p o w e rs

, ,

o f r e pr o ducti o n bact e ria hav e and that we are de aling in th e

,

cas e o f the se minute plants with a fo rce o f almo st infinite p o w e r .

Th e p o w e r o f gr o wth and r e pr o ducti o n o f th e bact e rial c e ll

is d ue in all probability to th e fact that th e f o o d o f bact e ria

, ,

is hi ghl y o rganiz e d and must b e in s o luti o n and r e ady fo r ab

so rptio n and assimilatio n Thi s o rganic fo o d is de rive d fr o m

.

th e di si nt e grati o n o f c o mpl e x animal and plant c o mp o und s .

G r e e n plants usually manufactur e th e ir f o o d fro m simpl e r

s ubstanc e s ( C0 HO nitrit e s and nitrat e s)
2 , o, Animals and .

fungi use co mpl e x o rganic co mp o unds fo r f o o d and b e fo re ,

 REPR O DU CTI ON OF B ACTERIA 45

th e y ca n b e assimilat e d the y must b e di ge s te d and bro k e n

up into s o me what simpl e r co mp o unds Th e fo o d us e d by
.

m o st bacte ria r e quir e s little o r no che mical alte ratio n b e fo r e

it can b e assimilate d by th e p ro t o plasm o f th e ce ll B acte ria .

a re no t c o mp e ll e d to s e a r ch ab o ut fo r f o o d as so m e animal s

d o but liv e natu r ally o nly in th e mid s t o f nut r i e nt mat e r ial

, .

I t sh o ul d b e n o t e d that r e p r o ducti o n in s o m e pr o t o z o a ( am oe ba)

ta k e s place fo r a time as rapidly as in th e bacte ria but o nly ,

afte r th e ce ll h as r e ce ive d and as similate d sufli cie nt fo o d .

On acc o unt o f th e i r natural c o nditi o ns o f lif e bact e ria can

gro w fas te r and r e pro duce m o r e rapidly than any o th e r animal
o r plant c e ll . Th e re is a gre at va riatio n amo ng bacte ria in
r e gard to th e rapidi ty with which the y r e pro duce S o m e .

S p e ci e s a re e xc e e di ngly rapid ( B . subtilis) and o the rs a re

,

co mparative ly sl o w (B act tub e rcul o sis)

. .

Th e flage lla o f th e m o til e bact e ria d o no t in any way

inte rfe r e with th e pro ce ss o f fi ssio n amo ng th e s e Sp e cie s .

I n th e l o ph o trich o us bact e ria th e no n fl age l l a te e nd o f th e

-

dividing ce ll fur ni sh e s flage lla fo r o ne o f th e ne w individuals .

Th e o th e r c e ll us e s th e o l d flag e llum o f th e par e nt .

Th e se o l d flage lla may b e carri e d fo ge n e r ati o ns Th e co n

r .

tigud us e nds o f th e c e lls n e v e r p r o duc e any flag e lla. I n amphi

trich o us bact e ria ne w flag e lla a re pr o vid e d by b o th ne w indi

v id ual s. The s e c e lls th e re f o r e have a ne w a nd an o ld se t o f

l o co mo tive o rgans I t is he ld by s o me that amphi t richo us bac
.

te ria d o no t e xi st and that th e bip o lar flag e lla r e sult during th e

,

pro ce ss o f fis si o n o f a mo no tricho us bacte rium th e ne windivid

,
4 6 MO RP HOL OGY
ual having pro duce d a ne wflage llum b e fo re s e parating fr o m th e
o riginal c e ll I n th e p e ritrich o us bact e ria as th e c e lls a re e l o n
.

gating pr e parato ry to fissi o n ne w flage lla a re de ve l o p e d at th e

,

p o int o f co nstrictio n b e twe e n th e Ol d flage lla On di visio n b e .

ing co mpl e t e d th e ne wflage lla co m e to l ie o n th e e nd o f th e ce ll s

,
.

S p o re F o m a ti o n
r Sp o r e f o rmati o n acco rding to vari o u s
.
,

w rit e r s is give n as a m e ans o f multiplicati o n Th e fo r mati o n

,
.

o f th e sp o r e is a m e an s o f carryi ng o u t o ne pha se o f th e p ro

c e ss o f r e pr o ducti o n Th e te rm spo re is appl ie d to th e r e st

.

ing stag e s o f plant and animal ce ll s Obvi o usly th e sp e cie s

.
,

must b e pre s e r ve d b e fo r e r e pro ductio n and multipl icatio n

can tak e place Th e spo r e has b e e n like n e d by s o m e to th e
.

s e e d o f plants Th e sp o re and th e se e d se rve to ca r ry th e

.

par e nt ce ll s thro ugh unfavo rabl e co nditio ns fo de finit e p e ri o ds r

in th e life hi sto ry o f th e o rganism S p o r e s a e pro duce d by

. r

s o me bacilli and a fe w S pi rilla T h e firs t car e ful study o f

.

S p o r e s was mad e by K o ch in 87 8 in his study o f B act e r ium

1

anthracis and by C o h n in hi s w o rk o n B acillus subtilis

,
.

S p o re f o rmati o n u s ually tak e s plac e wh e n th e bact e rial c e ll

is surro und e d by unfavo r abl e co n di tio ns su ch as e xhau stio n ,

o f th e f o o d s upply accumulatio n o f e xc re to ry pr o du ct s ab
, ,

n o rmal te mp e rature s e tc I t sh o uld b e no t e d in thiS co n

,
.

ne ctio n that c e rtain pro to zo an parasite s ( amoe ba) b e

co m e e ncyst e d whe n surr o unde d by th e ab o v e m e ntio n e d -

co n di tio n s A sp o r e will ge r minate withi n ce r tain limits o f

.

tim e whe n th e n o rmal co nditio ns a e p ro vide d Th e fir s t

,
r .

indicatio n o f sp o re fo r matio n in a bacte rial ce ll is e vide nce d

 REPR ODUCTI O N OF B AC TERI A 47

by l o s s o f th e p o w e r o f lo co mo ti o n in mo til e f o rms th e fo rmatio n ,

o f small highly r e f r actil e g r anul e s withi n th e ce ll The s e

'

, .

granul s a sum e th e app e arance Of gl o bul e s and co al e sce e ithe r

e s

in th e middl e at th e e nd o r v e ry n e ar th e e nd o f th e c e ll
, , Th e .

co al e sce d gl o b ul e thus f o rm e d i nc r e a se s in size and b e co m e s

S ph e rical o s ph e ro idal i n shap e ( F ig
r Th e sp o r e n e ve r .

b e co m e s as l o ng as th e bact e rial
c e l l b u t f r e qu e n tly b e c o m e s
,

m any tim e s b r o ad e r thu s di s t e nd ,

ing th e ce ll wall Th e sp o r e do e s .

n o t alway s di s te n d th e ro d .

Wh e n th e sp o r e is gre at e r in F 3 — S p r f rm ati Aft r IG . 2 . o e o on . e

Fi h r sc e
di am e t e r tha n th e m o th e r c e ll a nd
'

is in th e ce nte r o f th e ro d a spindl e shap e d app e arance may ,

-

b e p r o duc e d ( B b utyricus) I f it is at th e e nd o f th e c e ll th e
. .
,

c o mm o n d r um sti ck f o rm is pro duce d (B t e tani) Occasi o n . .

ally wh e n th e Sp o re is l o cat e d a slight di stance f ro m th e

,

e nd o f th e c e ll th e app e aranc e o f th e c e ll i s that o f a w e dg e

,
.

This last m e n -

tio ne d f o rmati o n

i s no t charact e r
0 .
i stic fo r any o ne

S p e ci e s but re
sul ts r ath e r as a

S p o rt
“
( Fig
”
.

FIG 33
. S p r f rm ati
. o h wi g vari p iti
e o f
on s
Wh
o
e n th e
n
sp o re
o us os o ns o

p r Ori g al s o es .
i fully f o rm e d
in .
s ,
48 MO RP HOL OGY
th e ce ll wall o f th e m o th e r c e ll ruptu r e s and lib e rate s th e
sp o r e surr o unde d by a ne wly s e cr e te d and hi ghly r e si stant
m e mbran e Th e r e st Of th e o riginal c e ll unde rgo e s di sinte
.

g ra t i o.n Wh e n a sp o r e d e v e l o ps int o a v e g e tativ e f o rm it

usually incr e as e s in size l o se s its hi gh r e fractive p o w e r
, ,

and r uptur e s Th e Sp o r e may ruptur e at th e e nd and f o rm

.

a small p o re o r o p e ning F r o m this Op e ning th e pr o to pla sm

.

gro ws o ut and f o rms th e characte ri stic bacillus o spirillum r .

Th e o l d sp o r e m e mb r a ne can b e o ft e n di s c e r n e d attach e d to

th e ne w v e g e tative f o r m O cca si o nally th e sp o r e m e mbran e

.

splits l o ngitudinal ly and s o m e tim e s transve r se ly so that th e ,

y o ung ce ll has part o f th e sp o r e o n e ithe r e nd o r o n b o th sid e s

o f it fo r a S h o rt p e ri o d a s it g r o ws in l e ngth . I n a fe w S p e ci e s

o f bact e ria th e r e i s no rupt u r e o f th e sp o r e m e mbran e it b e i ng

,

ab so rb e d by th e de ve lo ping ve ge tative c e ll I t is pro babl e

.

that th e bacte rial c e ll go e into th e sp o r e stag e as a m e a ns o f

s

carryi ng th e c e ll thro ugh unfavo rabl e co nditio ns to a tim e

wh e n no r mal co n di tio ns will b e pre se nt and th e ce l l can re
pro duc e S o m e h o ld th e Vi e w that sp o rulati o n is th e hi ghe st
.

stag e o f th e lif e histo ry o f ce rtain bact e ria I t is n e c e s sary

.

fo r th e se bact e ria to pa s s th r o ugh this s tag e in o r d e r to main

tain a vig o r o us sp e ci e s This co nditio n o f aff air s h as b e e n

.

n o te d amo ng ce rtai n o f th e alga O the r s b e li e ve that it is

.

always n e c e ssary fo r a bacte rium to b e plac e d in u nfavo rabl e

surr o undings to fo r m sp o r e s Y e t at c e rtai n te mp e rature s
.

whi ch a re favo r abl e fo r bact e rial gro wth sp o r ulatio n o ccu r s .

I t is p o ssibl e that th e e nvir o nm e nt may b e rapidly chang e d

 REPR ODU CTI ON OF B ACT ERI A 49

and no t alway s b e p e rc e ptibl e and thi s may acco unt fo th e

, r

fo rmatio n o f sp o r e s in ce rtain cas e s wh e re th e e nviro nm e nt ap

p e ars n o rmal B e lo w and ab o ve th e p o int whe r e sp o rulati o n
.

take s plac e at a t e mp e r atur e favo rabl e fo r re p ro d ubtio n no

spo r e s a e fo rme d alth o ugh fi ssio n may tak e place
r , .

C e rtai n sp e ci e s o f bact e ria may b e mad e to l o s e th e p o w e r

o f sp o r e f o r mati o n wh e n gr o wn at high t e mp e ratur e ( B act .

a nthracis) G as e o us e nviro nm e nt may al so influe nc e th e

.

p o w e r o f a sp e ci e s o f bacte ria to f o r m sp o r e s B acte rium .

anthracis in th e animal b o dy d o e s no t fo rm sp o re s but wh e n ,

give n a large am o unt o f o xyge n o n artificial me dia it r e adily

, ,

f o rms th e m .

Th e sp o r e is a v e ry r e si s tant b o dy I t is r e si s tant to h e at
.
,

s o m e sp e ci e s with standing ve ry high te mp e ratu r e s to ,

co ld and to che mi cals Sp o r e s as w e ll as ve ge tati e f o rms

,
.
,
v ,

v a ry in th e ir p o w e r s o f r e sistanc e with th e sp ci e s F o e xampl e

e . r ,

th e v e g e tative fo rm o f B act e rium anthra ci s i s kill e d by o ne

p e r c e nt ph e n o l in t w o minut e s and ,
in th e sp o r e s tag e it

r e quir e s fro m o ne to fifte e n days ( K o ch) Anth r ax sp o r e s will

.

with stand air drying fo r y e a r s The y a e th e m st r e si stant

. r o

spo r e s kno wn Th e t e st o f e fli cie ncy o f all di si nfe ctants is pa r

.

ti a l l y ba se d o n th e ir ability to d e s t r o y anthrax S p o r e s Th e .

g re t re sistance o f th e sp o r e is d ue to th e ve ry co mpact and

a

imp e rme able m e mb r ane which su r ro unds it a nd al o to th e fact ,

s

that its pro to plasm c ntains l e ss wat e r than th e ve ge tative f o rm

o .

Th e m o r e wat e r th e r e is i n a c e ll th e e a i e r it i s c o agulat e d
s .

Th e S p o r e is c o mp o s e d o f c o nd e n se d p ro t o pl sm a O r dinary.
5 0 MO RPH OL O GY
bacte ri o l o gical stains ( anilin dye s) do no t stain sp o re s I t .
‘

re quire s sp e cial m e th o d s to stain th e se high ly r e fractil e b o di e s .

Arth o sp o re s
r . I n c e rtain v e g e tativ e bact e rial c e lls C lado ,

thrix Thio thrix and p e rhap s a fe wmicro c o cci which gro w in a

, ,

filame nto us fo rm it is claim e d by s o me write rs ( Hue pp e e t al )

, ,
.

that s o m e o f th e c e ll s incr e as e in size b e co me r e fractil e but , ,

sta in e asily and d e ve l o p a hi gh e r p o we r o f r e si stanc e than th e

o th e r m e mb e r s o f th e sam e gr o up o f individuals Th e y d e .

tach th e ms e lve s fro m th e thre ad and swim ab o ut in the m e

d ium surr o un di ng th e m The s e c e lls a e calle d a rth spo re s
. r ro .
l

I t is claim e d that wh e n r e p r o ducti o n tak e s plac e th e se c e lls

‘

are th e o nl y o n e s c o nc e r ne d in th e p r o c e ss The r e is no ne w .

fo rmatio n Withi n th e pro to plasm o f th e v e g e tative c e ll as in ,

m o st Sp o re fo rming bact e ria but th e wh o l e o f th e pr o t o plasm

-

o f th e c e ll e nt e r s int o th e f o rmati o n o f th e b o dy kn o wn as th e

arthro sp o r e Th e y d o no t p o sse ss any such incre ase d p o w e rs

.
‘

o f r e sistanc e as d o e ndo sp o r e s Th e maj o rity o f b acte rio l o

.

gists at th e pre se nt d o no t re co gnize any such b o dy a s th e

arthro spo r e ( s e e Fig .

R e pro d ucti o n am o ng th e Tri ch o b acte i a r ( H igh e r B acte i a ) r

Th e high e r bact e ria a re usually filam e nto u s fo rms and in a ,

large numb e r o f sp e ci e s p o sse ss she aths Th e filame nts a e m de . r a

up o f individuals whi ch a re se pa r ate but co nne cte d t rminally e

with e ach o th e r One e nd o f th e filame nt is usually attach e d

.

to s o m e Obj e ct Aft e r r e achi ng a c e rtain l e ngth th e c e lls o n

.

1
No t use d in th e se nse pr p d by D B ary
o o se e .
 REPRODU C T I O N OF B AC TERI A 5 1

th e fre e e nd o f th e filame nt divide in o ne two o r thre e plan e s

, ,
‘

a nd f o rm r e pr o ductiv e b o di e o f a sp e cializ e d typ e kn o wn as

s
go nidia The se go nidia are cast o ff acquire flag e lla in so m e
.
,

S p e ci e s and fo r a p e ri o d may b e m o til e

, , ,
. The y th e n atta ch
to s o m e o bj e ct ( I r e pr o duc e a ne w filam e nt . G o nidia are
fr e qu e ntly th am e shap e as th e o rigi nal ce ll altho ugh th e y
,

o ft e n as s um e a rath e r sph e r ical shap e in c e rtain sp e ci e s . At

o ne tim e g o ni di a w e r e tho ught by so me to b e S p o r e s but ,

th e y are no n r e si stant and d o no t give any o f th e staining

-

r e actio ns fo r sp o r e s .
 C HAP T ER V

M I NUTE STR UC TUR E OF T HE B AC TE R I A L C ELL

I nt r o d u cti o n No twith standing th e fact that th e bact e ria
.

a re am o ng th e small e st o rganic b o di e s k no wn many facts hav e ,

b e e n de t e rmi ne d in r e gard to th e ir mi nut e structur e Up to .

th e pr e se nt tim e h o w e ve r o ur kn o wl e dg e is v e ry e l e m e nta r y
, ,
.

Littl e ca n b e l e arn e d r e gardi ng th e minut e a nat o my o f th e

bact e rial c e ll wh e n in a Viabl e s tat e o r in a stai n e d pr e pa r a

ti o n und e r th e o rdinary micro sco p e e ve n whe n magnifi e d ,

10 00 to 2
0 00 diam e te rs M uch m o re mu st b e l e arn e d in
.

r e ga r d to th e anato my and che mistry o f th e bact e ria b e

f o r e we can have a c o mpl e t e unde r standi ng o f b a cte rio l o g
ical pro c e ss e s in g e ne ral I t is pro babl e that th e ne w ult r a
.

vio l e t micro sco p e wh e n p e rf e ct e d will b e o f valu in w o rking e

o ut th e structur e o f ba ct e ria I t h as s e e m e d advisabl e to co n

.

sid e r the s ubj e ct matt e r o f this chapt e r und e r two se parat e

-

h e ad s th e hi sto l o gy and th e ch e mical structur e

, .

H i s to l o gy o f th e B a cte r i al C e l l Th e c e ll s o f all S p e ci e s
.

o f bact e ria a re c o mp o se d o f that c o mm o n o r ga nic s ubsta nce

t e rm e d p ro t o plasm c e ll sap and a limiting m e mbran e similar

, ,

to so m e plant a n d so m e a nimal c e lls .

C e l l Wal l . Th e pro t o plasm is said to be nak e d in a gr e at

52
 MI NU T E S TRUCTURE OF THE B ACTERIAL CELL 53

many animal c e lls I t may b e state d that the r e is usually

.

a se mblance o f a c e ll wall in the se case s I n a fe w in stance s .

th e c e ll wall o f ani mal c e lls appr o ximat e s th e c e ll wall o f th e

,

/
plant i n st re ngth Th e bacte ria in s tructu r e stand b e tw e e n
.

th e pla nt and th e a ni mal F o r e xampl e th e c e ll wall is o f

.
,

a highe r rd e r than th e animal c e ll wall but it is no t so highly

o ,

de ve l o p e d as th e c e ll wall o f true plants I t h as b e e n p o sitive ly

.

pro ve d that th e bact e rial c e ll p o ss e ss a limiting ce ll wall

simi lar to plant ce lls in so me case s a nd a nimal c e ll s in o th e rs .

M a ny o th e r p o i nt s go to S h o w that th e bact e ria b e l o ng to an

int e r m e diat e class as far as structure is co nce rne d Th e c e ll .

d o e s no t p o sse ss any cl a rly d e fin e d nucl e us o r nucl e o lus and

e ,

in thi s pa r ticular diff e rs fro m th e maj o rity o f animal a nd

plant c e ll s S tain e d by th e o rdinary bact e rio l o gical stains
.

( anilin dy e s) no d e finit e s tru ctur e can b e d e mo nst r at e d o n

,

ac co unt o f th e int e n sity o f th e stain B act e ria stain m o r e

.

de e ply with th e anilin dy e s than plant ce ll s .

P r t pl as m
o o . Acc o r ding to so m e auth o riti e s th e pr o t o plasm
o f th e bact e rial c e ll is no t di s t ribut e d thr o ugh o ut th e wh o l e

c e ll but co mp o s e s a lay r ne xt to th e ce ll wall Th e ce ntral

,
e .

part o f th e c e ll in co mm o n with pl nt ce ll s co ntains th e c e ll

,
a ,

sap co mp o s e d la rg e ly o f wate r and o rganic and in o rganic

,

substan ce s in so lutio n Th e e xact co nditio n o f affai r s in all

.
,

p r o bability is that th e re is a so me what d n se r lay e r o f pro to

,
e

plasm call e d e cto pl asm ne xt to th e c ll wall This laye r is

, ,
e .

no t d e finite and s e parat e fr o m th e i nt e ri o r o f th e c e ll thu s ,

cre ating a ce ntral cavity as so m e autho rs state M any .

54 MO RP HO L O GY
thr e ads Of pro to plasmic mate rial run fro m this o ut e r lay e r
thro ugh th e inte ri o r o f th e ce ll fo rming as it w e re a n e two rk
,

o f pr o t o plasmic thr e ads in th e c e nt e r Th e int e r s tic e s o f

.

thi s n e two rk co ntai n th e wate r i no rganic and o rganic sub

, ,

stance s which co mp o se th e so call e d ce ll sap Th e sub

-
.

stan ce s withi n th e e cto plasm co mp o se th e nto pl m Th e e as .

thickn e ss o f th e o ute r pr o to plasmic lay e r o r e cto pl m va ri e s

,
as ,

i n th e s am e and di ff e r e nt S p e ci e s On th e ave r ag e it is thi ck e r

.

in th e Ol d ce lls .

B y th e w e ll kn o wn phy sical ph e n o m e n o n o f pl as m o lysi s

-

s o m e ve ry int e r e s ting p o ints can b e l e arn e d in r e gard to th e

structur e o f bacte ria F o r i ns tance if a live ba cte rial c e ll
.
,

is place d in a so luti o n o f hi gh e r d e n sity (hyp e rto nic s o luti o n)

than its c e ll sap th e s o lv e nts ( wate r e tc ) c o ntaine d th e r e in
, ,
.

will pas s thro ugh th e laye r o f p r o to plasm n e xt to th e ce ll wall ,

thro ugh th e ce ll wall to j o in th e liqui d o n th e o utside and thus ,

s e rve to dilut e th e salts pr e se nt in th e so lutio n Th e lay e r .

o f pr o t o pla s m ( e ct o pla s m) c o ns titut e s a se mip e rm e abl e m e m

bran e Th e ce ll wall is p e rm e abl e to m o st so lute s such as

.
,

salts ur e a suga r s e tc As a r e sult Of th e l o s s o f wat e r by th e

, , ,
.

bacte rial c e ll th e pr o t o pla sm r e tracts fr o m th e c e ll wall and

f o rms its e lf into i r r e gul ar masse s withi n th e c e ll I n ce rtain .

o f th e infus o ria th e p r o t o plasmi clay e r i s attach e d to th e c e ll

wall and th e who l e c e ll assume s an i r r e gular shap e This

,
.

e xp e rim e nt sh o ws that th e r e i s a d e finit e c e ll wall a nd that th e

pr o to plasm o f th e c e ll and th e ce ll wall are di stinct fro m e ach

o th e r . I t als o s e rv e s to e xplain c e rtain o th e r b o di e s such as ,
 MI NUT E S T RUC TURE OF TH E B AC T ERI AL C ELL 55

m e ta chro matic and p o lar granul e s whi ch will b e r e fe rre d to

,

late r.

The r e is a g re at di ff e r e nc e in th e way th e va rio us sp e ci e s o f

bacte ria are aff e cte d by s o lutio ns o f highe r and l o we d e nsi ty r
‘

o r o s m o tic pr e ssu r e Th e re is with o ut do ubt a gr e at diff e re nc e

’

ch e mically and anato mi cally b e twe e n th e p ro to plasm Of

di ff e r e nt sp e ci e s o f bacte ria and thi s acco u nts fo r th e inab il
,

ity o f so me sp e cie s to plasm o lyze r e adi ly The r e is a diff e re nc e

.

in th e p e rme ability o f th e e cto plasm to th e so lve nts Th . e

e ct o pla s m in s o m e fe w sp e ci e s d o e s no t pr e v e nt th e di ffu si n o

i n and o ut o f th e c e ll o f c e rtain pa r ti cular so lut e s whil e in ,

o th e r bact e ria th e s am e s o lut e is unabl e to pas s thr o ugh th e

e ct o pla s mi c lay e r . Thi s is what co n stitut e s a s mip e rm e abl e e

m e mbran e ,
a m e mbrane p e rm e abl e to s o lv nts but no t to e

s o lute s Ol d bact e ria plasm o lyz e mo r e e a ily than y o ung

. s

bact e ria and thi s is pro bably d ue to th e f o r m e r havi ng a

,

thicke r laye r o f pro to plasm o r s e mip e r me abl e m e mb r ane .

Th e c e ll sap co ntai ns so lut e s d e rive d fr o m e xt e r nal fluids

a s w ll as so m e g e ne rat e d withi n
e . Th e p r e se nce o f th e se so

lute s withi n p ro duce s a g re at o smo tic p re ssur e again s t th e

p ro to plasmic lay e r and th e ce ll wall and the r e is a co nsta nt
,

t e nd e ncy fo r th e ce ll s to b e co m e di st e nde d d ue to th pas sage

,
e

i nto th e ce ll o f so lve nt s such as wate r fr o m th e e xte ri o r T h e

, ,
.

pr e ssur e inside a bact e rial ce ll may e qual two o th r e e atmo s r

p h e
r e s
. I t i s d ue to th e r igidity o f th e c e ll wall that th e

bacte rial c e ll is abl e to with tand an int e rnal pr e ssur e whi ch

s

wo ul d e asily rupture th e pro to plasmic laye r whi ch r e ally

5 6 M ORP H O O L GY
de te rmin e s th e o sm o tic p r e ssur e Osmo tic pr e ssur e it
.
,

may b e said in passing d e p e nd s up o n th e am o unt o f cry s

,

tal l o i d s ( s o lut e s) in so luti o n P r e ssu r e is p r o duc e d by

.

m o l e cul e s a nd i o ns p o undi ng agai nst th e s e mip e r m e abl e m e m

bran e Th e r e is a gr e at diffe re nce in th e way in which vari o u s
.
'

substance s diff use thr o ugh this o r any se mip e rm e abl e m e m

brane Th e o sm o tic p r e ssure will b e di r e ctly pro p r tio nat e
. o

to th e numb e r o f m o l e cul e s o f no ne l e ct r o lyt e s ( uga r s a nd s

o th e r ca r b o hydrat e s ,
ur e a ,
I n th e ca se o f th e e l c e

tro l yte s ( s alts) th e i o n s in s o luti o n p r o du ce th e o sm o ti c

pr e ssur e .

I f bact e ria a re accust o m e d gradually to th e so luti n s o f o

high o sm o tic pr e ssu r (hyp e rto ni c s o luti o n s) by pa ssag e

e

thr o ugh fl uids o f i nc r e asi ng de nsity th r will b e no plas

,
e e

m o l ysis . Thi s sh o w s that th e ce ll is so c o nstitute d that th e

s o lut e s su ch a s salts a nd c ystal o id al no ne l e ct r o lyt e s diff u se
r

sl o wly th r o ugh th e e cto pla sm a nd b e co m e i nco rp o r at e d in th e

c e ll sap o r e nto plasm Th e e cto plasm is e asily p e r m e at e d by
.

s o lve nts a nd in time is p e r m e abl e to ce rt in so l ute s a .

I t i s p o ssibl e to chang e th e co nt e nt s o f a bact e rial c e ll by

placing it in a hyp o t o nic s o luti o n that is a s luti o n o f l e ss

, ,
o

d e nsity and co n se qu e ntly l o w e r o smo tic p r e ssu r e than th e

c ll sap
e . T h e classical e xp e rim e nt is to plac e ba ct e ria whi ch

have b e e n accus to m e d to g ro wing o n o r in cultur e m e di a co n

taini ng s alt s i nto a so l utio n o f di still e d wat e r Th e di still e d
.

wate r pa s re adily thro ugh th e ce ll wall and th e se m ip e rm e

sse

abl e e cto plasm Th e ce ll incr e ase s in siz e co ns e que nt to th e

.
 MI NUT E S TRU CT URE OF TH E B AC T ERI A L C ELL 57

abso rptio n o f wate r a nd if k e pt und r such co nditio ns fo r

,
e

a ny l e ngth o f tim e th e c e ll wall wi l l r uptu r e ( r h e xi s) a nd th e

p r o to plasmi c a nd the r co nte nts e scap e i nto th e sur r o unding

o

liquid (pla m ptysis) A six p e r c e nt s lutio n o f cane s ug r

s o . o a

e x e rt s a pr e ssur e o f 3 7 5 mm o f m e r cury at 0 .
4 de g re e C 1 s .

( S ixty p o u n d s to th e s quar e i nch ) J u s t S O u nd e r natu r.al co n ,

di tio ns th e fo r m a nd s t r u ctu r e o f a ba ct e rial c e ll may b e al

,

t e d ac co r di ng to th e cha r a ct e r o f th e f o o d a nd th e d e nsity o f
er

th e s u rr o u ndi ng m e dium Th e -ph e no m na o f pla sm o ly is

. e s

and pla sm o pty sis d ubtle ss acco unt fo r th e invo lutio n fo rms
o

amo ng ce r tain sp e ci e s and th e so call e d pl o mo rphi sm o f -

e

s o m e ce lls I t may b e said in thi s co nne cti o n that it is im

.

p o s sibl e to subj e ct all sp e cie s o f ba cte ria to th e ph e n o m e no n

o f pla s m o pty si s d ue to th e fact that th e ir c ll wall a nd
,
e

e ct o pla s m a re so c o nstitut e d that salts may di ffu se o ut m r e o

re adily than in o the rs .

M e tach o m a ti c and P o l a G a nul s

r Und e r c e rtai n co n
r r e .

di tio ns in s o m e sp e ci e s O f bact ria th e c e ll S h o w a p e c uliar

e s

g r anular app e arance wh e n staine d with th e o r dina ry anilin

dye s Th e g r anul e s are irr e gular in siz e d o no t o ccur in any
.
,

de fi nite ar r ange me nt a nd stain da rk e r than o at va rian ce

,
r

with th e su r o undi ng p ro to plasm (L o e ffl r s m e thyl e ne bl ue

r e
’

stain o n B act diphth e ria ) Th e se granul e s a e k no wn a s

. . r

m tach o m a ti c g a nul e s
e r r Th e p r o t o plasm in th e se ca se s se e ms
.

to hav e a b e ad e d app e a r an ce Th e r e have b e e n va rio us

.

staini ng me tho d de vi se d to de mo nstrat the se g ranul e s I n

s e .

o th e r sp e ci e s o f bact e ria th e r e i s a c o nc e ntrati o n o f th e ro

p
5 8 MO RP HOL O GY
tOp l as m at th e p o l e s o f th e ce ll Wh e n stain e d by th e vari o us
.

bacte rio lo gical stains th e s e granul e s o f co nde nse d pro to plasm

,

stain d e e ply and are w e ll diff e re ntiate d fro m th e r e st o f th e

c e ll The s e granul e s are kno wn as p o l a g anul e s ( me thyl e n e
. r r

blue stain o n B p e stis) F r e qu e ntly d ue to th e accumulati o n

. .

o f th e granul e s in c e rtai n parts o f th e c e ll th e r e maining S lightly

s tai ne d pr o t o plasm giv e s th e app e aranc e o f vacu o l e s The se

.

vacuo l e s app e ar to co ntain a cl e ar substance in all pro bability

,

c e ll sap As to th e significance o f b o th m e tachro matic a nd

.

p o lar granul e s the re is still do ubt Obvi o usly it is n e ce ssary

.
,

to d e t e rmin e wh e th e r th e r e is a ny c o n si s t e ncy in th e ir d e v e l o p

m e nt in c e r tain sp e ci e s o f bacte ria unde r any and all co nditi o ns .

I t may b e stat e d that c e r tai n bact e ria S h o w a slight granular

c o nditio n o f the ir pro to plasm at all time s (B act diphth e ri a ) .

,

whil e o the rs do no t S ho w such structure s with co nsiste ncy .

I t is fr e qu e ntly Ob se rv e d wh e n th e bact e ria a re gr o w n u nd e r

unsati sfacto ry co nditio ns o f te mp e rature m o i sture o r f o o d
, ,

supply I t has b e e n state d that thi s granular fo rmati o n

.

withi n s o m e bacte rial ce ll s p r e ce d e s r e pro ductio n Th e y w e r e

.

supp o s e d by so m e to b e similar to go nidia Th e y a e al so . r

kn o wn as B ab e s Ernst granul e s I t h as no t b e e n p o sitive ly

-
.

pro ve d that th e se granule s have anythi ng what e v e r to d o

with th e p r o ce ss o f r e pro ductio n So me write rs have claim e d
.

that the y w e r e fo o d pro ducts but thi s is an e rr o ne o us co n

,

ce p tio n I n all pr o babili ty th e y a re pr o duc e d arti ficially in

.

th e c e ll by plasm o lysis . AS m e nti o n e d ab o v e wh e n bact e ria

,

a re plac e d in a m e dia o f gr e at e r d e nsity ( hyp e rt o nic) than

 MI N UT E S T RUC T UR E OF THE BAC TERI AL CELL 59

that o f th e ir e nto plasm th e s o lve nts within th e ce ll di ff us e

,

o ut thr o ugh th e s e m ip e rm e abl e e ct o pla s m a nd th e r e main

,

ing p ro to plasm in th e ce ll r e tracts a nd co ll e cts in i r r e gular

particle s in vari o us p o rti o ns o f th e c e ll I t is plausible that
.

thi s co nde ns e d pro to plasm co mp o se s what are call e d me ta

chro matic and po la r g r anul e s S o m e autho rs have ascrib e d
.

to bact e ria th e p o w e r o f sp o nta ne o u s pla s m o ly si s to e xplain

th e i rr e gular staining o f such bact e r ia as B act e rium diphth e r i a ,

B act e rium tub e rcul o si s and th e p o l a r stai ning o f B acillus

,
o

p e stis (bub o nic p lague ) I t sho uld no t b e o ve rl o o k e d

.

that wh e n staini ng a pr e pa r atio n o f bact e ria all th e p re re qui

sit e s fo r plasmo ly si s a e p re se nt F o r e xampl e th e he ating
r .
,

o f th e p r e pa r ati o n b e f o r e th e s tai n i s appli e d driv e s th e wat e r

o ut o f th e c e ll and c o nc e ntrat e s th e s o lut e s a nd c o ns e qu e ntly

,

caus e s a r e tracti o n o f th e c e ll pro to plasm fro m th e ce ll wall ,

wh ich may co ll e ct in granul e s I t is p ro bable that th e fo r

.

matio n o f p o lar g r anul e s to g e the r with th e fo rmatio n o f

,

vacuo l e s a re e ithe r artificially pr o duce d o r that th e y are

, ,

de ge n e rative change s in th e ba cte rial ce ll De fi nit e g ranule s

.

o f sulphu rhav e b e e n n o t e d i n c e r tain o f th e hi gh e r bact e ria

( gg
B e i a to a ce a) .

N ucl e us . In r e gard to th e p e rpl e xing que stio n as to

wh e th e r o r no t a nucl e us e xi sts in th e bacte rial c e ll th e re h as ,

b e e n a g re at de al o f inve stigatio n AS s tate d b e fo re th e r e is

.
,

no d e fi nit e and circum s crib e d nucl e u s in th e bact e rial c e ll

'

Wh e n staine d wi th th e o r di nary nu cl e a r stains (h e mato xyli n ,

basic anili n dye s ,

th e mat e rial ( chr o matin ) whi ch tak e s
6O MO RP HOL OGY
th e stain is di stribute d thr o ugho ut th e ce ll The r e are also .

p o rti o n s o f th e c e ll whi ch in so m e sp e ci e s d o no t stain ( ach r o

matic) S o m e write rs ho ld th e Vi e w that th e chr o matic
.

sub stance is nucl e ar mat e rial Th e nucle ar mate rial h as

.
,

th e r e fo re a g e ne ral di stributio n thro ugho ut th e ce ll Thi s is

,
.

undo ubte dly tru e in th e light o f pre se nt r e se arch S o m e bac .

t e ria fo r e xampl e c e rtain Vibri o s co nsist almo s t e ntir e ly o f

, ,

chro matin Th e am o unt o f ch ro mati c mate rial vari e s a gre at

.

de al e ve n in th e sam e sp e cie s u nde r diffe r e nt co nditio ns I t .

h a s b e e n sugg e s t e d that th e r e as o n th e nucl e ar mat e rial is no t

aggr e gat e d in th e r e gula r fo rm o f th e nucl e us may b e ascrib e d
to th e fact that th e bact e ria a re l o w i n th e s tag e o f e vo luti o n ,

and that th e y divide so r api dl y that th e nucl e ar mate rial do e s

no t hav e sufli cie nt tim e to f o rm its e lf int o th e r e s ting s tag e whi ch

th e n o r mal nucl e u s r e pr e s e nts B li ts chl i s tudi e d c e r tain la r g e

.

fo rms o f bact e ria which co ntain sulphur g r anul e s ( B e g

g i a to a ce a ) and ,
h e c o nclud e d that th e larg e part o f th e c e ll
was nucl e us surro und e d by a thi n lay e r o f pr o to plasm F i sch e r .

i s inclin e d to b e li e v e that B il ts ch l i s pr e parati o ns w e r e d ue to

‘
’

plasm o lysis N akanishi claim s h e h as de mo n s trate d a d e fi

.

nit e nucl e us in th e c e ll s o f s o me S p e ci e s o f bacte ria Thi s .

O bs e rvati o n has no t b e e n c o nfi r m e d .

Ch e m i cal S tru ctu e o f th e B a cte ri al C e l l

r Our kno wl e dg e
.

o f th e e xact and co mpl e t e ch e mical c o n s tituti o n o f th e bact e rial

ce ll is at p r e s e nt quit e as limite d as o ur kn o wl e dge o f th e hi s

to l o gi cal structur e I t may b e said that bact e ria a e e xc e e d
. r

ingl y labil e and th e r e fo r e th e o rdi nary a nalytical m e th o ds

,
 MI NUT E S T RU CTURE OF THE B ACTERIAL CELL 61

canno t b e satisfacto rily appli e d to th e m I t is w e ll kno wn .

that th e ch e mi cal co n stitutio n o f th e bact e rial ce ll va ri e s

gr e atly in di ff e r e nt sp e ci e s and in th e s am e sp e ci e s whe n th e
,

bact e ria are gro wn o n diff e r e nt vari e tie s o f cultur e m e di a ,

a nd o n th e s am e m e dia Th e che mical c o nstitutio n al so

.

va rie s with th e age o f th e bacte ria Th e fo llo wing fro m We ll s .

,

C h e mical P ath o l o gy quo ting C ram e r give s a v e ry go o d

”
, ,

state me nt o f th e ir c o mp o sitio n Cho l e ra Vibri o s gro wn o n b o uil

.

l o n o n analy si s sh o w e d p e r c e n t pr o t e id a nd p e r

c e nt ash Wh e n gro wn o n m e dia f r e e fro m pro te in (Uschi nsky s

.
’

m e di um) th e y sho w e d 3 p e r c e nt pr o t e in a n d p e r c e nt
a sh . I n th e sam e m e dium p e r c e nt and p e r c e nt
pr o t e in w e r e no te d in two st r ains o f ch o l e ra vibrio s I t can .

no t b e s aid that any o ne sp e ci e s o f bact e ria has a d e fi nit e

ch e mical st r uctur e Ho we ve r th e re are s o m e ch e mical sub

.
,

stance s which are fo und in th e same Sp e ci e s o f bact e ria co n

tinuo usl y as fo r e xampl e th e fats in acid fa s t bact e ria
, , ,
Th e -
.

variatio n in structur e is m o re quantitative than qual itative .

Th e o ld e r analys e s o f which th e r e a re ma ny a re o n th e
, ,

wh o l e inco rr e ct in th e light o f mo d e rn m e th o d s o f r e s e arch .

Th e maj o r ity o f bact e ria c o ntain liquid s p ro t e in s wat e r a nd , , ,

salts such as th o s e o f s o di um p o tassium magne sium and ph o s

, , , ,

p h o ru s P h
. o s ph o rus in all pr o bability o ccur s in th e la r g e s t

amo unts Acco rding to a r e ce nt inve s tigati n o f I wan o ff th e

. o

principal c nstitu e nt o f th e bacte rial c e ll is a ge nuine nucl e o

o

p ro t e in co ntaining trace s o f iro n and sulphur An th e r . o

write r N i shi mura has r e p o rte d th e finding O f ce r tain purin

, ,
62 MO RP HOL O GY
base s th e nucl e o pr o te in e nd pro ducts such as guanin ade nin
,
- -

, , ,

and xanthi n in bacte rial c e ll s We lls says that th e nu cl e o .

pro t e ids o f bacte rial ce ll s have b e e n kno wn to split Off p e n

t o se s similar to high e r nucl e o pro t e ins C e rtain bacte ria have -
.

b e e n analyze d and fo und to b e co mp o s e d o f gl yco nucl e o

pro t e ins and mucin (e g B co l i) (Le ach) N e ncki iso late d a
. . . .

nitro g e n o us b o dy fro m putre factive bact e ria which is S imilar ,

in co mp o siti o n to p e pto n e Glo bulins and nucl e o album e ns .

-

hav e b e e n r e p o rte d in th e pr o to plasm o f bacte ria Th e .

g e lati no us mat e r ial s e c r e t e d by th e bact e ria in z o ogl oe al and

p s e u d o ozOg l oe l f o m a tio ns is v e r y similar i n ch e mical c o n s ti
a r

tuti o n to mu ci n .

F ats hav e b e e n f o u nd in th e pr o to pla s m o f a larg e n umb e r

o f sp e ci e s o f bact e ria T h e fat h a s b e e n discl o s e d by staining

.

r e actio ns and by th e e xtractio n o f th e ce ll s with th e vari o us

fat so lve nts such as h o t alco ho l chl o ro fo rm e the r e tc
, , , ,
.

L e cithi n a ph o sph o riz e d fat ch o l e s te rin a m o nat o mic alc o h o l

, ,

d e rive d fro m fatty acids ; simpl e and sp e cial bact e rial fats ,

have b e e n iso lat e d S o m e lip o chr o me s ( co l o ring matt e r o f

.

fats) are also pr e se nt Th e pr o p o r tio n o f mate rial so lubl e in

.

fat so lve nts in B act e rium tub e r cul o sis and o th e r acid fast bac -

t e ria is ab o ut 2 to 4 p e r ce nt o f th e w e ight Of th e bact e ria

0 0 .

With S udan I I I whi ch is a characte ri stic s tain fo r fat it

, ,

h as b e e n p o ssibl e to d e m o nstrat e fat o nly in B act e ria anthracis ,

S taphyl o co ccus py o g e n e s var aur e u s B act m uco sis and in all .

,
.
,

th e acid fa s t bact e ria i e

-
B act tub e rc ul o si s B act l e pr a
,
. .
,
.
,
.
,

B act sm e gmatis B act m oe ll e ri (grass bacillus) B act butyri

.
,
.
,
.
 MI NU T E S T RU CT URE OF THE BACT ERI AL CELL 63

(butt e r bacillus ) and o th e rs I n o th e

,
r sp e ci e s no fat was d e .

m o nstra te d by S udan I I I B acte ria fo rm th e m o s t o f th e ir

~ .

fat wh e n gro wn o n p o tat o o r glyce rine agar Th e amo unt .

o f th e fat in a bact e rial c e ll may b e in c e as e d by g bw ing th e r r

bacte ria o n a fat rich me dium -

.

K re sling giv e s th e f o ll o wing analy sis o f th e mat e rial aft e r

1

e xtracting B act e rium tub e rcul o sis with chl o r o f o rm .

Fr fatt y a id
ee ,
c s pe r ce nt

N utral fat nd fatt y a id t r

e s a c es e s pe r ce n t
Al h l
co btai n d fr m fatt y a id t r
o s o e o c es e s pe r ce nt

L ithin
ec pe r ce nt

S ub t n s a l ubl in wat r
ce s so e e pe r ce n t .

fo r e go ing analysis sho ws th e fatty de rivative s which

Th e

may b e pr e se nt in o ne Sp e ci e s o f bacte ria .

I t h a s b e e n n o t e d by s o m e ch e mists that th e e th e rial

e xtracts d o no t c o ntai n th e acid—fast s ub s tanc e s I t has .

b e e n claim e d that th e so call e d acid fas t sub s tance is an - -

alco h o l havi ng s o m e o f th e pro p e rti e s o f a wa This sub x .

tance was so lubl e in h o t ab so lute alco ho l o r e th e r and in so lubl e

in c o ld alc h o l and e th e r thu s s ugge sting i ts similarity to a
o ,

wa Ol e c i so ce tinic and nysistini c a nd so m e sap o nifi e d

''
‘
x . l , , ,

lau r ic acids a re pre s e nt and it is fro m th e m that th e fats are

,

f o rme d .

I n all pr o babil ity th e in so lubl e c e ll wall o f bact e ria is

no t c o mp o se d o f pu r e c e l lul s e Th e id e a that th e wall o r o .

limiting me mbrane o f th e ce l l was co mp o se d o f ce llulo s e

1
Q uo t dfe ro m W ll e s
’
C h m i al P ath l gy
e c o o .
64 MO RPHOL O GY
or h e mi c e l lul o s e like m o st plant ce lls has b e e n he ld fo r
-

, ,

y e ars by a larg e numb e r o f bact e rio lo gi s ts R e ce nt r e s e arch

.

has sho w n that th e c e ll wall in th e maj o rity o f sp e ci e s is

m o re lik e that o f c e rtain animal c e ll s than o f plants Th e .

c e ll wall is like chitin I t is i nso lubl e and splits into 80 p e r

.

c e nt to 90 p e r c e nt Of a nitro ge n o us ca rb o hyd rat e gl yco sa m in

( W e ll.s) C hi tin is an amin o co mp o und o f a ca rb o hydrat e
-
.

I t 15 a p o lym e r o f a simpl e animal carb o hydrat e as ce l l u

l o se is a p o lyme r o f a simpl e plant carb o hydrate ( We ll s) .

C hitin is f o u nd in qui t e a fe w plac e s such as o n th e wings

,

o f c e rtai n i nse ct s o n th e c o v e r ing s o f b e e tl e s and i n th e sh e lls

, ,

o f c e rtain cru s tac e a ns s u ch as th e crab and l o bst e r

,
.

N o de finit e carb o hyd r at e s hav e b e e n r e p o rte d in th e bac

te rial c e ll and pr o v e n abs o lut e ly to b e pr e s e nt
,
C e rtain sub
.

stance s similar to glyco g e n have b e e n re p o rte d in B acte rium

tub e rcul o sis (Le ve ne ) I n s o me fe w bacte ria a cco rdi ng to
.
,

C ram e r wh e n tr e at e d with i o di n e a sub s tanc e giving th e blu e

, ,

co l o r re acti o n fo r starch is n o te d .

C hl o r o phyll h as b e e n r e p o rt e d in s o m e bact e ria On inv e s

.

tiga tio n it has b e e n f o und that it is no t c o rr e ct to class th e s e

o rganisms as bact e ria a nd that th e y sh o uld b e plac e d am o ng

th e alg a o r S chi o p h yce a

z . S ulphur granul e s hav e b e e n n o t e d

in ce rtain sp e cie s o f bact e ria in th e cla s B e ggia to a ce a

s .

P igm e nt gra nul e s a re n o t e d am o ng so m e sp e ci e s o f bact e ria

( chr o m o g e nic ) wh e n c e rtain d e finit e co nditi o n s a e pr o vid e d r .

Th e chr o m o g e nic bact e ria and th e ch e mi s try o f pigm e nts will

b e co nsid e r e d in a f o ll o wi ng chapt e r .
 MI NUT E S T RUCT URE O F THE B AC T ERI AL CELL 65

So me bacte ria fo rm withi n th e ir p ro to pla sm co mpl e x che m

ical sub stance s kno wn as t o xins e ndo to xin s pto mains and
, , ,

e nzym e s Th e ch e mistry o f th e se co mp o und s is kno wn o nly to

.

a limit e d e xte nt The y wil l b e co nside r e d mo re in de tail in

.

sub se que nt chapte rs .

Ch e mi cal S tru ctu r e o f S p o r e s and F l ge l l a aSp o r e s a re

.

co mp o se d o f co nde ns e d pro to plasm Th e y co ntain th e same

.

che mical substance s as th e bact e rial ce ll fro m which th e y we re

fo rme d with th e e xce ptio n that th e Co nte nt o f wate r is l o w e re d
,

much b e lo w that o f th e o ri ginal ce ll Sp o r e s are co mp o se d

.

o f 6 p e r c e nt dri e d substa nc e ( Drym o nt)

0 Th e wat e r which
.

is p r e s e nt is h e ld in a way to r e sist drying t e mp e rature s b e

l o w 00 C Th e S p o r e wall is chiti no us in s tructur e rath e r

°
1 .

than ce ll ulo se l ike and co ntains in additio n so m e v e ry

-

,
”

hygro sco pic e xtractive matte r (We lls) .

F l age ll a are c o mp o s e d o f c o nd e n se d pr o t e in Aside fr o m .

thi s as far as is kn o wn th e ir ch e mical co nstitue nts a e no t

, ,
r

unl ik e th o s e in th e v e g e tativ e f o rm .
 P A RT III . M ET H O DS US E D I N
THE S T U DY OF BACT E R I A

CHAP T ER VI

CULTUR E M EDIA

n u cti o n Ve ry littl e can b e de te rmin e d ab o ut th e

I tro d .

bacte ria by micr o sc o pical e xaminatio n al o n e I t is n e c e ssary .

to cultivat e th e m artificially in o rd e r to study th e m satis

facto il y
r . Th e r e as o n fo r thi s will b e s e e n wh e n o ne r e aliz e s

that upwards o f two th o usand di ff e r e nt sp e ci e s o f bacte ria have

b e e n de scrib e d s o m e o f th e m man s w o r s t e n e mi e s and all
,
’
,

o f th e m b e l o nging to th e thr e e s impl e f o rm typ e s which hav e

b e e n pr e vi o usly de scrib e d B y micro sco pical e xaminatio n we

.

can l e ar n s o m e thi ng o f th e ir di s tributi o n so me thi ng o f th e ir

,

f o rm and structur e and s o m e thi ng o f th e ir r e lati o n to o r at

, ,

l e ast th e ir as so ciatio n with dis e a se s and such natu ral pr o c e s se s

, ,

as fe rm e ntatio n putr e factio n e tc but n o thi ng m o re

, ,
.
,
.

Th e artificial cultivati o n o f bact e ria h a s b e e n d e v e l o p e d

e ntir e ly withi n th e la s t h alf c e ntury Th e fir s t att e mpts w e r e

.

crude and unsatisfact o ry as Vi e we d fro m o ur pr e se nt s tand

,

p o int but since th e time o f P aste ur and K o ch gre at pr o gr e s s

,

has b e e n made Th e sci e nce o f bacte rio l o gy b egins with this

.

w o rk Th e unde rlying principl e s o f th e artifi cial cultivati o n

.

66
 C UL T URE E
M DI A 67

of bacte ria include th e se le ctio n o f f o o d me di a th e se curing ,

and mainta ining its ste rility and th e me tho ds o f iso latio n
,

and subculturing .

Ch aracte r o f F o o d re quire d b y B a cte ri a —

Th e kind o f
.

fo o d r e quire d by bacte ria vari e s gr e atly with th e diffe r e nt

fo rms So me r e quire fo r in s tance living o rgani c matte r ;
.
, ,

o th e rs r e quir e and live o nly o n d e ad o rganic matt e r ; whil e

s till o the rs live o nly in th e abs e nce o f o rganic matte r .

Att e mpts hav e b e e n mad e to suppl y a unive rsal m e di um .

This is e vide n tly an imp o ssibility M e di a have b e e n de vis e d

.
,

ho we ve r whi ch furnish a g o o d pabulum fo r a large numb e r

,

o f bact e ria . V ari o us de co cti o ns have b e e n use d in th e past ,

such as tho s e fro m fruit wh e at p e a straw manure e tc ;

, , , ,
.

but th e m e dium mo st uni ve rsally use d at th e pr e se nt time

is that sugge ste d by Lo e fll e r which is a b e e f bro th o r b o uillo n
,
.

Up o n thi s m e di um m o st o f th e bact e ria can gro w wh e n ap

pro pri e te additio ns have b e e n made with th e e xce ptio n o f th o se
,

bacte ria whi ch r e quir e living o rgani c matte r ; that is th e o bli ,

gate parasite s and th e bacte ria whi ch live o nly o n in o rganic

fo o d substance s .

C o m po s i ti o n and P re p arati o n o f Va ri o us Cul ture M e d i a .

A larg e n umb e r o f diff e r e nt c ul tur e m e di a hav e b e e n d e vis e d

and are in co nstant use but an atte mpt will b e made he re to
,

di scu ss o nly th o s e that a re m o st bo m m o nl y us e d and such ,

di scussio ns will b e o nly o f a g e n e ral nature I t will b e .

n e ce ssary to r e fe r to lab o rato ry guide s fo r th e e xact fo r

mul a and m e th o ds o f pr e para ti o n C ultur e me di a may b e
.
68 M E T HODS U S E D IN THE S TUDY OF B ACT ERIA
r o ughly di vide d into tho s e co ntaining albumin o us matte r and ,

the s e may b e e ith e r liquid s o lid o r l iq ue fi ab l e so lid m e di a

, , ,

a nd no nalb um ino us o r synth e tic m e dia .

Al b um i no us or P ro te in M e di a

Li qui d M e di a

B o ui l l o n Thi s m e di um is th e b asI S o f m o st o f th e o rdinary

.

m e dia us e d in th e lab o rato ry such as g e latin and agar and is

, ,

also larg e ly us e d a s a cul ture me di um its e lf I t has th e fo l .

l o wing ingre di e nts Th e e xtractio ns o f 5 00 grams o f l e an

.

b e e f 0 gram s o f p e pto n e 5 grams o f s o dium chl o ride ne u

,
1 , ,

tral i e d a nd dis so lv e d in o ne lit e r o f di s til l e d wat e r

z Th e nutri .

tive value o f th e vario u s ingr e die nts is a subj e ct that sh o uld b e

bri e fly co nside r e d at this p o int Th e nutritive value o f th e
.

m e at de p e nds almo st e ntir e ly up o n th e albumin o us sub

stance s Of whi ch it is co mp o s e d but mo s t o f th e s e are inso lubl e
,

and m o st o f th e m that d o go int o so lutio n are pr e cipitate d o n

he ati ng so that v e ry little pro te in mat e rial is furni sh e d by
,

th e m e at I ts purp o s e in th e m e dium is larg e ly to furnish e x

‘

tractive s chi e fly cr e atin and xanthi n b o die s whi ch are use ful
, ,

to m o st bact e ria T h e p e pt o n e whi ch is a s o lubl e albumi n

.
, ,

fo rm e d fro m pro te ins by th e actio n Of th e gastric juic e is ,

th e principal s o urce o f nitr o g e n fo r th e bact e r ia Th e so .

dium chl o rid e was o riginally us e d to aid in th e s o luti o n o f

th e p e pt o n e but it is no w no t n e e d e d fo r that purp o s e and in

,

s o m e lab o rato rie s its use is b e ing discarde d .

 C UL TURE M DI A E 69

I n th e pr e paratio n o f b o uill o n o ne o f th e mo s t imp o rtant

,

things to b e c o nsid e r e d is its n e utralizati o n AS a ge ne ral rul e .

th e bact e ria r e quir e an alkalin e rath e r than an acid r e acti o n ,

and th e m e di um mu st the r e fo r e b e made e ith e r ne utral o r

, ,

ve ry slightly alkal in e Th e indicato rs sugge ste d in th e

.

vari o us te xtb o o ks have b e e n re d and blue litmus pap e r tu ,

m e ric pap e r and ph e n o lphthal e in S o m e o rganic substance s

,
.

a re b o th acid and al kalin e to c e rtain indi cat o rs fr e sh milk ,

fo r instanc e will turn re d litmus p ap e r blu e and blu e litmus

,

pap e r re d Oth e r substance s particularly th e pho sphat e s

.
, ,

may b e n e utral to o ne in di cato r and acid o r alkalin e to an o th e r .

The se substance s thus give a do ubl e o r a mp ho te ri c r e acti o n .

I t has b e e n d e t e rmin e d by S chultz and o th e rs lat e r that

, ,

ph e no lphthal e in is l e ast lik e ly to give ambigu o us re sults and ,

it has larg e ly r e place d all o th e r indicato rs in thi s wo rk I t .

mus t b e u nde rs to o d ho w e ve r that it r e acts to th e carb o nate s

, , ,

such as carb o n di o xide and me di a sh o uld b e n e utraliz e d with

,

thi s indicat o r o nly afte r b e ing b o il e d and whil e still h o t I t is .

also tru e that th e alkaline p o int with ph e no lphthal e in is re

m o ve d so m e dis tance fr o m th e n e utral p o int o f litmus so that ,

wh e n a m e di um is n e utralize d by m e ans o f phe no lphthal e in it

is m o r e alkalin e than th e bacte ria n e e d and it is n e c e ssary o n
,

that acco unt wh e n m e dia is n e utralize d in th is way to add a

, ,

ce rtain amo unt o f n o rmal acid to th e me dium to bring it back

to a p o int m o st suitabl e to th e bact e ria Wh e n a me dium is
.

just S lightly alkalin e to phe no lphthal e in it is re pr e se nte d as ,

ha ving a r e actio n o f 0 Any alkalin e adde d in addi ti o n to this

.
7 0 M E TH ODS U S E D IN THE S TUDY OF B AC TERI A
w o uld b e de signate d by th e pe r ce nt r e pre se nte d in fi gure s
pr e ce de d by th e minus S ign ; and me dia whi ch is acid to
phe no lphthal e in is r e pre s e nte d by th e p e ce nt o f acid p re r

c e de d by th e plus S ign M o st o f th e bacte ria fo r in stance re

.
, ,

quir e a me dium fro m 0 to 1 ; whil e c e rtain bact e ria s uch ,

a s th o se c o mm o nl y f o u nd in wat e r r e ach th e ir maximum o f

,

de ve l o pm e nt wh e n th e me dium is 1 5 I n th e pr e paratio n . .

o f b o ui ll o n as w e ll as that o f ma ny o th e r m e dia it is n e ce s sary

, ,

to cl e ar it and this is d o n e by addi ng a n e gg o r s o m e si m ilar

, ,

albumino us mate rial whi l e th e m e dium is co o l at le ast

, ,

b e lo w th e co agul atio n p o int o f th e pro te in s i e 60 C . .

°
.
,

the n h e at e d sufli cie ntl y to th o ro ugh ly co agulate th e albu

mino us sub s tance Thi s draws th e fi ne particl e s Of th e p re
.

cip ita te int o la r g e r clump s o ma ss e s which wil l b e h e ld back

,
r ,

by th e filt e r mate rial and th e m e di um which go e s thro ugh th e

,

fil t e r u nd e r th e se c o nditi o ns will b e p e r f e ctly cl e ar .

B o uill o n may b e m o difi e d in vari o u s way s by th e additi o n

o f glyc e r in e ( 6 p e r c e nt) fo r th e B a ct tub e r cul o si s and B act. .

mall e i by th e addi tio n o f d e xtro se o r lacto se ( p e r c e nt) fo r 1

f e rme ntatio n te sts ; carb o lic acid 5 p e r c e nt ) fo r B ty .

p h o sus, e tc .

S ugar -
F re e B o ui l l o n ( S mith)
ce rtain w o rk such
. Fo r ,

as th e de te ctio n o f indo l a sp e cial me dium is us e d fro m which

,

th e muscl e sugar h as b e e n r e m o ve d S inc e mu s cl e always .

co ntains a small am o unt o f sugar (in o sit) it is n e ce ssa ry to

re mo ve thi s and it is do n e at S m ith s sugge sti o n by ino cul at
,
’

ing a bacte rium capabl e o f splitting up thi s sugar and do ing

 CUL T URE E
M DI A 7 1

it whil e th e m e at is b e ing di ge ste d I n thi s way th e sugar is .

go tte n rid o f and th e pre paratio n o f th e me dium go e s o n fro m

,

thi s p o int as in th e cas e o f o rdi nary b o uillo n .

Dunham s S o l uti o n ’
Dunham s s o luti o n i s use d at ti ne s
.
’
r

fo r th e study o f bacte ria I t simply c o nsists o f a 1 p e r c e nt

.

p e pto ne s o lutio n .

M il k .Th e mi lk o f c o co anuts has b e e n us e d in sp e cial

instance s but co w s mi lk is ve ry fr e qu e ntly us e d fo r gro wing

,
’

bacte ria F re sh milk sho ul d b e s e cure d and th e fat s e parat e d

.
,

pr e fe rably by ce ntrifugali zati o n M ilk is so m e tim e s quite .

difli cul t to st e rilize o n acc o unt o f th e v e ry r e sista nt sp o r e s

whi ch it may co ntain a nd ste aming o n f o ur o r fi v e co ns e cutive
,

days may b e n e ce ssary Litmus is fr e qu e ntly adde d to milk

.
,

whi ch incr e as e s th e us e fuln e ss o f thi s ve ry us e ful m e dium .

Am o ng o the r liquid m e di a us e d a re wh e y urin e and b e e r , ,

w o rt Th e latte r is us e d almo st e xclusive ly fo r th e gro wth o f

.

ye ast .

S o lid M e di a

P o ta to e s . P o tat o e s w e re firs t us e d by S chro e d e r . He

b o il e d the m with th e ir jacke ts o n and th e n cut the m in two
with a s te ril e kni f e place d th e m in a co ve re d di sh and gre w
, ,

bacte ria o n th e cut surface Esma r ch late r us e d Sli ce s o f .

p o tato e s in sp e cial glass di sh e s whi ch h e had made ; but at th e

pre se nt time th e y are us e d almo st e ntire ly in t e s t tub e s and
P e tri di sh e s C ylinde rs o f th e p o tato with a mping fro nt
.
,

surface are put in te st tub e s which are st e rilize d and us e d as

,

o th e r te st tub e m e di a I t is d e sirabl e that th e c o l o r

.
7 2 M E THODS U S E D IN THE S T UDY OF B AC TERIA
of th e p o tato sh o uld no t b e chang e d and to s e cur e this ,

it is b e st to e ith e r b o il th e p o tato o r p e rhaps what is b e tt e r , ,

to k e e p th e cut p o tat o e s i n ru nning wat e r fo r tw e lve h o urs o r

m o r e b e fo re th e cylinde rs a e cut o ut r .

Othe Ve ge ta bl e s
r C arr o ts turnips and o th e r v e g e tabl e s
.
, ,

a re s o m e tim e s us e d fo r sp e ci al bact e ria in th e sam e way that

p o tato e s are us e d .

B l o o d S e rum . Thi s mate rial furnish e s a sp e cially valuabl e

m e dium fo r c e rtain di se ase pro ducing bact e ria B e e f ho rse
-
.
, ,

a nd human s e rum a re th e o n e s c o mm o nly us e d I n a ny ca se .

,

th e bl o o d is c o ll e ct e d all o w e d to cl o t and th e n th e cl e ar s e rum

, ,

is drawn Off Thi s is put into th e appr o priate ve ss e l and

.

ste ril ize d Th e t e mp e ratur e o f s te rilizatio n is usual ly l o w

.

in o rde r to pre se rv e as much as p o ssibl e th e t r anspa re ncy

, ,

o f th e m e di um B y ste rilizati o n b e l o w th e co agulati o n

.

p o int o r by m e ans o f th e addi tio n o f wate r which rai se s

, ,

th e c o agul ati o n p o int it is p o s sibl e to st e riliz e this m e dium in

,

a liquid c o n di tio n but it is usually s o lidifi e d o r inspissate d A

,
.

S p e cial mixtur e kn o wn as Lo e ffl e r s bl o o d s e rum i s mad e by

’

addi ng o ne part o f d e xtr o se b o uillo n to thr e e parts o f b e e f

bl o o d s e rum I t is almo st unive rsally us e d fo r gro wi ng th e
.

B act di phth e ri a in diagn o stic lab o rato ri e s

. .

E ggs . Eggs a re us e d at th e pr e s e nt tim e larg e ly fo r th e

gro wth o f B act tub e rcul o sis F r e sh he n s e ggs a e bro k e n

. .
’
r

with gre at e s t care th e co nte nts put in s te ril e ve s se l s S hak e n

, ,

up and run i nto st e rile t e st tub e s wh e r e it is h e ate d and co agu

late d Eggs may also b e use d fo r o th e r wo rk
. .
 C UL TURE E
M DI A 73

Li qu e fi ab l e S o l i d M e di a

Ge l ati n . G e latin cultur e m e di um is mad e fro m b o uillo n

by th e additio n o f ge latin Ge latin is made fro m co nn e ctive
.

tissue e xtracte d e ithe r at o r di nary pre ssure o r in th e dige s

,

to r ( aut o clave ) . I ts physical charact e ristics that a re o f

sp e cial i nt e r e st to th e bacte ri o l o gist are its m e lting and s o lidi

fyi ng p o i nts .I t is n e c e s sary that a g e latin sh o uld b e us e d

whi ch in o rdi nary p e rce ntage s will no t inte rfe r e with bact e rial
gr o wth and wil l r e main so lid at th e t e mp e rature o f a warm
,

ro o m C h e mi cally its mo st imp o rtant f e ature is its acidity

.
,

and th e high e r grade s which have a l o w acidity co mpara ,

ti v e l y sh o uld b e us e d
,
. F r o m 10 to 18 p e r c e nt o f g e latin

is adde d to b o uillo n in th e pr e paratio n o f thi s m e di um Th e .

pr e parati o n o f g e latin Off e rs no difli cul tie s b e y o nd that o f

b o uill o n e xce pt that it is ne c e ssary to e xe r ci se g r e at car e in
n e utralizatio n and to fil te r it whil e h o t F o r th e filtratio n
.

o f g e latin and th e o th e r l i q ue fi ab l e s o lid m e dia we t ab so rb

,

e nt c o tt o n can b e us e d to advantag e altho ugh it is p o ssibl e

,

to fil t e r th e pr o p e rly pr e par e d m e dium thro ugh filte r pap e r .

I t is b e st st e riliz e d in str e aming s t e am o n thre e c o ns e cutive

days ; but it may b e ste rilize d in an auto clave at a pre ssure no t

to e xc e e d te n p o und s fo r fift e e n minut e s A fact that must
.

always b e b o rn e in m ind in pre paring ge latin is that its ge

latinizing p o we r is injur e d by pro l o nge d he ating duri ng th e
pro c e s so f pre paratio n o r ste rili zatio n and is l o st imm e diate ly
,

whe n h e ate d to 140 C in th e auto clave

°
. .
74 M E TH ODS U S E D IN THE S T UDY OF B ACTERI A
G e latin is fre qu e ntly m o difi e d by th e addi ti o n o f such sub
stance s a s de xtro se lacto se ; and a m e dium fo rm e rly e x te n
,
é

S iv e l y us e d fo r th e is o lati o n o f B typho sus wa s a p o tat o

.

wat e r ge latin co ntaini ng p o tassium io dide ( Elsn e r )

Agar . Agar is a c ul ture m e dium mad e by adding a s mall

p e r c e n t o f th e v e g e tabl e agar agar to

-
b o ui ll o n Agar agar .
-

is a s e aw e e d that g ro ws in th e P acific O ce an and it is kn o wn ,

as B e ngal isinglass o r C e ylo n mo ss I t o ccu rs in th e marke t

,
.

as transpar e nt strips 01 sticks and as a whi te p o wde r I t

, ,
.

diff e rs fr o m g e latin in b e ing a carb o hydrate simply and no t a ,

pro te in I t is ne utral in r e acti o n and me lts at ab o ut 99 C

.
°
.

and so li difi e s at ab o ut 40 C I ts chi e f value in bacte ri o l o gy

°
.

co m e s fr o m its hi gh m e lting p o int and al so fro m th e fact that

,

it is no t acte d up o n by f e rm e nts which liqu e fy ge latin so that ,

it may b e use d to gro w bacte ria whi ch n e e d a high e r te m

p e ra tu re than g e lati n w o uld r e main s o l id at and to gr o w
bacte ria which wo uld lique fy ge latin F ro m 1 to 2 p e r c e nt .

o f th e agar agar is us e d
-
Th e pr e parati o n o f agar o fl e rs s o m e
.

difli cul ty to th e b e ginn e r o n acc o unt o f its difficult s o lubility

and its h igh s o lidifying p o int whi ch make s it di ffi cult to filte r ;
,

but if th e agar is go tt e n into a s o lutio n firs t in th e wate r and ,

th e o th e r ingr e di e nts add e d lat e r it may b e pr e par e d ve ry

,

r e adily and e ve n fil t e r e d thro ugh filte r pap e r at th e o rdi nary

ro o m t e mp e rature Fo r filtrati o n ho w e ve r abs o rb e nt co tto n
.
, ,

is fre que ntly us e d and th e pro ce ss is s o me tim e s haste ne d by

th e use o i air s ucti o n
-

Agar is vari o usly m o difi e d by th e addi ti o n o f sugars glyc ,

 C UL T URE E
M DI A 75

e rine e tc A ve ry co mm o n mo dificatio nis that o f Wurtz

,
.
,

wh e r e b o th lacto s e and litmus are adde d .

N o nal b um ino us ( S yn th e si z e d ) M e di a Th e album i no us .

-

m e di a have a ve ry inde fi ni te co mp o sitio n and it is quite

.

imp o ssibl e to make two batche s o f m e di a e xactly alike Thi s .

is ve ry unfo rtunate b e cause ce rtain change s in th e characte r

is tics o f bact e r ia a re caus e d by e v e n slight cha ng e s in th e

co mp o sitio n o f th e m e di um ; and it is so me time s d ifli cul t to

d e te rmi n e whe th e r slight diffe re nce s that a e re co gnize d in r

cultur e s are d ue to i nh e re nt di ff e re nce s in th e o rgani sm o r to

diffe re n ce s in th e cultur e m e dia I t w o uld b e v e ry de si r abl e
.
,

the re fo re to have a cultu r e m e dium o f de fi nite co mp o sitio n

,

which co ul d b e e xactly r e pro duce d fr o m tim e to time At .

t e mpts have b e e n made to pr o duce such m e di a by th e use o f

che m ically pu re substanc e s in whi ch th e nitro ge n wa s fur
ni sh e d in a c o mparative ly l o w f o rm as fo r instanc e s o dium
, , ,

a sp a ra gina te Dujardi n S h o w e d that fu ngi c o uld gr o w in

.

no nal b um ino us substanc e s as e arly as 84 L at e r P ast e ur

1 1 .

u se d th e m in hi s ye as t wo rk ; and in 893 Usch insky sh o we d 1

that m o st patho ge ni c bact e ria w o uld gro w o n such m e di a .

Uschi nsky s m e di um w hi ch may b e give n as an illus trati o n

’
,

o f th e s e m e dia had th e fo ll o wing co mp o si tio n :

,

W at e r
G l y ce ri ne

S o di um hl rid
c o e

C al i um hl rid
c c o e

M agn i um ul p hat
es s e
7 6 M E THODS U S E D IN THE S TUDY OF B AC T ERIA
B ip o tassium p h o sp hat e 2 to

Am m o n um i la tat
c e 6 to 7

So di um a sp aragina te

This m e dium h as b e e n use d fo r sp e cial purp o s s but fo r e ,

ma ny fo rms it h as no t b e e n fo und as valuabl e as Uschinsky

tho ught B ri e ge r and o th e r s have f o und it use ful in study
.

ing th e pro ductio n o f pto main s and t o xins On th e wh o l e .

,

th e use o f synth e tic m e di a in th e past has b e e n di s app o inting .

I t is to b e h o p e d that in s o m e o f Emil F i s ch e r s p o l ype p tie d s

’
,

substance s will b e fo und which will m e e t th e de mands o f th e

bacte ria .

A s o li d synth e siz e d m e dium has b e e n us e d to a limi te d

,

e xt e nt .I t c o nsists o f a nutri e nt s o luti o n to whi ch wat e r

glass o r a s o dium silicate j e lly is adde d ; in this way a so lid ,

transpar e nt m e di um is fo rm e d and up o n thi s th e nitrifying

,

bact e r ia which de ve lo p o nly o n no nal b um ino us m e dia have

, ,

b e e n gro wn .
 CHAP TER VI I

S TER I L I Z ATI ON

t
I ntro d uc i o n . B y st e rilizati o n is m e ant th e pr o ce ss by

whi ch th e vitality o f all th e m icro Organi sm s in o r up o n an

o bj e ct a re d e s tr o y e d
,
and an Obj e ct which is e ntire ly fre e
fr o m all living micro o rgani sms and the ir sp o r e s is de s crib e d
a s st e ril e
.

S t e rilizati o n h as co m e to b e c o nfo und e d with disinf e cti o n .

B y di si nfe cti o n i s me ant a pr o c e ss which insur e s th e d e struc

ti o n o f tho se livi ng f o rm s which have th e p o we r Of pr o ducing

an i nfe cti o n o r dis e as e and may o r may no t b e co mple te in
, ,

th e s e n se o f s t e rilizati o n .

I n th e lab o r at o ry it is cust o mary to sp e ak o f s t e rilizati o n as

th e pr o c e ss whi ch fr e e s a substanc e fr o m bact e ria but d o e s

,

no t chang e its c o mp o siti o n o r i nt e r f e r e with i ts nut r itiv e

quality That is it r e nde rs it g e rm fr e e but no t barr e n This

.
,
-
.

is usually acco mpli sh e d by a physical ag e nt Di si nfe cti o n has

.

co me to imply a change in co mp o sitio n d ue to th e fact that

,

ch e mical s a e usually e mpl o ye d in th e pro ce ss Whil e th e

r .

ab o ve t e rms a re no t strictly co rr e ct the ir use is co nve ni e nt

,
.

B o th ste ril izati o n and di si nfe ctio n may b e acco mpli she d by
e ith e r physical o r ch e mical ag e nts .

77
7 8 M E TH ODS U S E D IN THE S T UDY OF BACT ERI A

r za ti o n
S te il i b y Ph ysi cal Age
'
nts
F il tra ti o n p o ssibl e to ste rilize fluids by me ans o f
. I t is

filtrati o n B y thi s m e ans th e bacte ria are actually r e mo ve d

. .

P ractically th e o nly m e ans o f pr o ducing a g e rm fr e e filtrat e -

is by m e an s o f unglaze d p o rce lain Thi s has b e e n us e d e sp e .

cial l y wh e r e it i s d e si r e d to s t e ril i z e sub sta II Ce S that w o uld b e

’

change d wh e n subj e cte d to th e acti o n o f h e at o r ch e micals .

F iltrati o n is u s e d la rge ly to fr e e th e bact e r ia f r o m th e ir so lubl e

pro ducts o f gro wth e g th e ir to xins Th e r e a e two syste ms

,
. . . r

o f p o rc e lain filt e r s o ne i s th e P a s t e ur C hamb e rlain th e o th e r

,
-

th e B e rk e f e ld syst e m Th e P a s t e ur sy s t e m is th e m o s t
.

r e liabl e but it is ve ry S l o w Th e B e rke fe ld is much m o r e

, .

rapid but it is co nse qu e ntly no t so r e liabl e With e ithe r

, , ,
.

syste m g re at car e is n e c e s sary with th e ir use and fr e que nt

st e rilizati o n o f th e m is n e ce ssarysince th e bacte ria fr e que ntly
.

gr o w thro ugh th e ir walls afte r th e y have b e e n in use fo r a

sh o rt tim e .

S unli ght has g o o d di sinfe cting o r st e rilizing pr o p e rti e s

whi ch w e r e first call e d att e nti o n to by Do wns and B lunt in

187 7 and sinc e th e n many e xp e r im e nts hav e b e e n p e rf o rm e d
,
.

Whil e it has th e p o w e r o f de stro yi ng bacte ria wh e n th e y are

e xp o s e d to it fo r a fe w mi nut e s o r a fe w h o urs its p o w e r ,

o f d e t r o ying th e m cann o t b e d e p e nd e d up o n and it is o f

s ,

no p r actical imp o rtanc e in thi s c o nn e cti o n .

C ld
o C o ld is o r di narily th o ught to b e v e ry e ffe ctiv e in
.

de str o ying bact e ria but in re ality it is o f ve ry littl e valu e

,
.
 S TE R I L I zAT I O N 79

F ri sch in ,
xp o se d b o th bacilli and m icro co cci to minus
187 7 , e

8 7 C pr o du ce d by th e vap r ati o n o f li q uid carb o n di o xide

°
.
,
e o .

Al l such bact e ria g r e w wh e n br o ught und e r favo rabl e c o ndi

tio ns .

A numb e r o f lat e r e xp e rim e nts sh o w e d that whil e fr e e zi ng

kill s a ce r tain numb e r o f th e bacte ria it do e s no t kill all O f
the m by any m e ans and many fo rms m ay r e main alive fo r
,

m o nth s n fro ze n wate r Thi s is true al so o f di se as e pro duc

I .
-

ing bact e ria .

D y H at
r
— I n 85 4 S ch ul tz and S chwann fi r st sh o w e d
e . 1 ,

th e r e al valu e o f d r y h e at in d e s tr o yi ng bact e ria in th e ir e x

p er i m e nts up o n s p o ntan e o us g e n e r ati o n ( s e e C hap I I ) . .

3

Di ct F l am e
re Thi s is c o n stantly u se d in th e lab o rato ry
.

fo r st e rilizing th e platinum n e e dl e s but its use i s alm o st ,

e ntir e ly limit e d to thi s and to th e d e s tructi o n o f w o rthl e s s

but i nf e cti o us mate rial such as th e carcasse s o f infe cte d,

..

ani mal s I n thi s case a sp e cial apparatus is n e ce ssary

.
, ,

kn o wn as th e incine rato r .

Hi gh D y H t Dry h e at at a t e mp e ratur e o f 5 0 C
°
r ea . 1 .

is co nstantly us ed in th e lab o r ato ry fo r th e ste rilizatio n o f

such mate rial s as gla sswar e co tto n batting e tc I t has b e e n , ,
.

sh o wn that thi s t e mp e r ature is sufli cie nt to kill all fo rms o f

life a nd may b e de p e nde d up o n as a m e ans o f p ro ducing p e r
,

fe ct st e rilizati o n pr o viding th e h e at ca n re ad ily r e ach all

,

pa r ts o f th e o bj e ct s to b e s te rilize d Th e he at is supplie d in .

an o v n o e a h t air ste ril ize r Thi s is so m e time s pro vide d

r o -
.

with a th e rm o m e t e r but usual ly this is no t ne c e s sary since

, ,
80 M E THODS U S E D IN THE S TUDY OF B ACTERI A ‘

incipie nt charring o f co tt o n o ccu r s at ab o ut thi s te mp e ratur e ;

and it is thus o nly n e ce s sary to run th e st e rilize r until co tto n
in th e flask o r t e st tub e s o r co tto n place d in fo r this sp e cial
,

purp o s e is slightly bro wne d

,
.

Thi s m e th o d o f ste rilizatio n has a ve ry limite d use since it ,

canno t e vide ntly b e use d fo r s te rilizi ng liquids ; and b e caus e

, ,

o f th e charring o f v e g e tabl e and ani mal matt e r at th e ab o v e

t e mp e rature thi s m e tho d canno t b e u se d to ste rilize sub

,

stanc e s made o f the s e and similar mate rials .

M o i s t He a t S t e am fo r th e purp o s e o f st e rilizati o n has

.
-

b e e n use d since th e tim e o f S chultz and S chwann s e p e ri ’

x

m e nt and is applie d in th e fo rm o f e ith e r stre aming ste am o r

,

as st e am unde r pre ssure .

S tre ami ng S te a m — Th e t e mp e ratur e o f stre a ming st e am

.

de p e nds o f co urse up o n th e baro me tric pre ssure but app o xi

, , ,
r

mat e s 00 C I t is much mo re e ffi ci e nt than dry he at at th e

1
°
.

sam e t e mp e ratur e Thi s is pro bably d ue to th e fact that th e

.

walls o r m e mbrane s o f th e b acte ria b e co m e m o i st e n e d and

, ,

swe ll e d by th e ste am and thus p e rmit th e m o r e r e ady e ntrance

o f th e h e at to th e int e ri o r o f th e c e ll B y m e a ns o f s tr e ami ng
.

st e am mate rials may b e ste riliz e d witho ut i njury whi ch wo uld

no t s ta nd d r y h e at I t is h o w e v e r practically imp o ssibl e to
.
, ,

kill bacte ria whe n th e y are in th e sp o r e co nditio n by thi s

m e ans fo r it has b e e n fo und that an e xp os u re o f sixte e n h o urs
,

is n e ce ssary to kill c e rtain sp o r e bact e ria fo und in th e so il A .

m e tho d whi ch o ve rco me s th e difli cul ty o f l o ng e xp o su re was

sugge ste d by Tyndall and is kn o wn as th e inte r mitte nt m e tho d
 S TERI LI Z ATI ON 81

of ste rilizati o n as th e di sco n tinuo us m e th o d and s o me time s

, ,

as Tyndall izatio n .

I n this m e th o d c ul ture m e di a a e s ubj e ct e d to a ste ril i a

r z

ti o n o f fro m te n to twe nty minute s o n o ne day al l o w e d ’

to sta nd twe nty fOurh o u r s and h e ate d again th e n all o w e d

-

to stand an o th e r tw e nty fo ur h o u r s a nd h e at e d a thi r d tim e

-

, .

Thi s me tho d almo s t i nvariably pro duce s co mpl e te ste ril i a z

tio n pro viding o f co urs e th e mate rial has r e ach e d th e p ro p e r

, , ,

te mp e rature thro ugho ut I ts e fli cie ncy de p e nds up o n two

.

thi ngs : fi r st th e fact that st re ami ng ste am applied fo r te n to

,

twe nty m inli te s kil l s al l ge rmlife not in th e sp o re s tage ; and

’

w
,

s e co nd s tanding twe nty fo ur ho u rs all o s the sp o re s which

,
-

we re no t kille d o n th e first applicatio n o f s te am Opp o rtunity ,

fo r g e rmi nati o n S O that o n a s e c o nd e xp o s u r e th e y o ung

,

bacte ria a e kille d Th e third e xp o sure fo r te n to twe nty

r .

minute s make s it almo st ce rtain that all sp o re s have ge rmi

na te d and h ave b e e n kill e d Evid e ntl y th is m e tho d is no t
.

applicable to all mate rials si nce an e ss e ntial fe atu re is th e

,

Op p o rtlini ty fo r th e g e rmi nati on O f th e sp o r e s I t w o uld b e

l

im p o ssibl é to st e ril ize e mpty gl assware i nstrum e nt s ban , ,

dage s e tc by thi s me tho d since the re is no o ppo rtunity o n

,
.
, ,

the s e mate rial s fo r th e gro wth o f the bacte ria in th e i nte rvals
b e twe e n he atings .

Thi s me th o d o f ste rilizati o n is car r ie d o ut in a ste ame r o f ,

whi ch the re are a numb e r o f di ffe re nt typ e s Al l that is .

n e ce ssary is a ve s se l which will ho ld wat e r whi ch has a sh e lf ,

a b o ve th e wat e r a nd a fairly tight c o v e r

,
An e xc e ll e nt fo rm
.
,

G
82 M E TH ODS U S E D IN THE S T UDY OF B ACT ERIA
alm o t unive rsally us e d in thi co untry is th e Arno ld ste am
s s ,

ste ril i ze r ( Fig Th is h as th e advantag e o f b e ing p rac

.

tically auto matic sinc e th e s te am which is pro duce d is COn

,

d e nse d a nd r e h e at e d .

S te m und e P ssu e
a S t e am und e r pr e ss ur e has b e e n
r re r .

u se d by th e F r e nch s ch o o l fo r a go o d whil e and is no w co ming

into g e ne ral use He r e a t e mp e ratu re o f 0 5 to . C is 1 12 0
°
.

o maintaine d fo r fift e e n mi nute s .. ,

th e latt e r t e mp e r atu r e i s d e

st r uctive o f all f o rms o f ge rm

life and may b e de p e nd e d up o n
a s a c e rtain m e an s o f s te ril i a z

ti o n Thi s me th o d o f st e rili .

b za tio n i s made p o ssibl e by th e .

FIG —A i m pl t am t riliz r e
4 3 s s e s e e

i ti g f a g alva iz d ir pail use O f a Sp e Cl al p l e CC Of appa

. .

'

co ns s n o n e on .

a 18th v r d b fal b m m ratu s kn o wn as the auto clave

e co e an a se o '

whi ch is e sse ntially a co pp e r v e ss e l that can b e h e rm e tically

s e al e d pr o vid e d with a th e rm o m e t e r and st e am gaug e
,
In .

p r actic e it is no t f o und ne c e s sary to use a th e r m o m e t e r si nc e ,

th e t e mp e r atur e h a s a d e finit e rati o to th e p r e s s u r e p ro v i d ,

i ng h o w e ve r that th e atm o sph e r e withi n th e aut o clav e is

, ,

saturat e d with wat e r and this can b e acc o mpli she d by ,

l e avi ng a p e t co ck o p e n until s t e am e scap e s f re e ly

-
.

On acco u nt o f th e q uickne ss with w hi ch mat e rial s can b e

ste r ilize d in th e aut o clave thi s m e tho d is b e co mi ng d e s e rve dly ,

p o pular At fi rs t it was th o ught that th e high t e mp e ratur e

.

inju re d th e cultur e m e dia but it has b e e n fo und that the ,

 S TERI LI ZATI O N 83

o rdi nary culture me dia a e uninjure d by th e e xp o sur e ne ce s

r

sary fo r ste rili zatio n Wh e n sugar m e dia a e h e ate d in th e

. r

aut o clave th e y a e fr e qu e ntly dark e ne d b e cause o f a change

,
r

p ro duce d in th e sugar by th e
high h e at Thi s change is
.

k no wn as ca r ame lizatio n A .

c o mmi tte e o f th e Ame ri can

P ublic He alth As s o ciati o n
h as h o w e ve r r e c e ntly ag r e e d
, ,

that th e auto clave may b e

us e d fo th e st e ril izati o n o f
r

th e s m e dia pro vi di ng th e
e ,

t e mp e ratu re do e s no t ge t
ab o ve 1 C st e am p r e s sure
2 0
°
.
,

fifte e n p o unds and th e e ,

x

p o s u r e i s n o t l o ng e r than ‘

fifte e n m inut e s .

An e xc e e di ngly co nve ni e nt
f o rm o f au to clave is that
manufacture d by B ramhall
a nd De a n a nd sh o wn in
,

F igure 3 5 .

Th e i nte rmitt e nt m e th o d
F 35 A t lav f with t am
IG . . u oc e o r us e s e

o f s te rl l l a tio n h a s b e e n u se d
z d r Pr r un e e ssu e

fo r s t e rilizing bl o o d a nd th e r pro t e i n mat e rial by using a

o

t e mp e ratu r e b e l o w th e co agul ti o n p o i nt o f th e mate rial to b e

a

s t e rili ze d e g 5 5 to 60 C I t is n e ce ssary h o w e ve r to mak e

°
. .
, .
, ,
84 M E TH ODS U S E D IN THE S T UDY OF B AC T ERI A
m o re than thre e e xp o sur e s and to maintain th e te mp e ratu re
,

fo a p e ri o d o f tim e c o nsid e r ably ab o v e tw e nty mi nut e s (o ne

r

h o ur o r m o r e )
.

Di s i nf e cti o n
. Ch e mi cal sub s tan ce s a e u s e d in th e
r

lab o rato ry fo r th e purp o se o f d e s t ro yi ng o l d cultur e s fo r ,

washi ng th e d e sks a nd co ve ri ng up inf e cti o us matt e r acci

,

d e ntally spill e d washi ng th e hands e tc Th e ch e micals

, ,
.

u sually u se d a e ca rb o lic acid ( 3 to 5 p e r ce nt so lutio n)

r ,

c o rr o sive sil bl im ate ( to

I Th e valu e o f th e s e i s d is

cuss e d in a lat e r chapte r (se e Chap X I ) . .

A fe w di si nf e ctants hav e b e e n us e d fo r st e rilizing cultur e

m e dia su ch as ch l o ro fo r m and hydr ge n p e ro xid e Th e s e
,
o .

ch e mical sub sta nce s a e lat e r e ith e r drive n O ff by h e at o r a e

r ,
r

r e nde re d ha r ml e ss by th e natural change s which go o n so that ,

th e nutritive pr o p e rti e s o f th e substanc e s a e no t d e str o y e d

r .
 CHAP TER VI I I

C UL TUR E M E TH ODS AN D APPAR ATUS

In tro d ucti o n Th e artificial cultivatio n o f bacte ria marks

.

th e b e ginning o f th e sci e nc e o f bact e ri o l o gy L ittl e was d o n e

.

with th e bacte ria o kno wn ab o ut the m until it was p o ssibl e

r

to cultivat e th e m a r tificially . Tlfe m e th o d s o f cultivati o n

w e re diffi cul t and rathe r slo wly d e ve l o p e d Si nce bact e ria

.

o ccur alway s in natur e in a mi x e d cul tur e p r actically no

,

pro gre ss was made until m e tho ds o f is o latio n w e re de vi se d .

Th e picking up o f th e S i ngl e o r ganism wa s so difficult a ta sk

that it was fir st do n e by B arb e r within th e las t fe w y e ars A .

c o n side rabl e numb e r o f di ffe re nt p e o pl e have take n part in

th e d e ve l o pm e nt o f o ur pr e se nt m e tho d s but it is r e ally to
,

R o b e rt K o ch that chi e f cr e dit is d ue , a nd hi s plat e cultur e

m e tho d o f i so latio n r e ally marks th e b e ginning o f accurate

‘

wo rk in thi s li ne Wh e n a si ngl e o rganism is plant e d o n a

.

s o lid culture m e dium th e r e sulting mas s o f gro wth is calle d

,

a co lo ny and is e vide ntly made up o f th e de sce ndants o f a

singl e c e ll
. Wh e n a co lo ny is o btaine d we have a mate rial
,

fo r making subcultur e s micro sc o pical e xaminati o ns animal

, ,

i no culatio ns e tc; Cul ture s made fro m a S ingle co lo ny are

,

call e d pure culture s and in stud yl ng any o rganism th e first

,

r e qui site is to ge t a pure cultur e Wh e n this is o btain e d the

.
,

85
86 M E TH ODS U S E D IN THE S T UDY OF B ACTERIA
study o f it can pr o ce e d with ce rtainty In th e e arl y d ays o f .
'

bacte ri o lo gy mixe d culture s we re fr e que ntly co nfus e d with

,

pur e culture s and curi o us and ve ry e rro n e o us ide as w e re

,

o btain e d which it has tak e n many y e ars to c o rr e ct .

M e th o d s of I s o l ati o n

F racti o nal M e tho ds . P ast e ur andw o rking o n th e Co hn ,

pro bl e m o f artificial cul tivatio n w o rk e d o n th e supp o siti o n
,

that vario us sp e ci e s o f mi cro Organism s o ccurring in a mix ,

tur e will multiply un e qually in any give n cultur e me di um a c

,

co rding to th e ir di ff e r e nt natur e s Acco rdi ng to this m e th o d

.
,

o ne sp e ci e s will attain th e asc e nd e ncy a nd can b e re m o v e d

and o btain e d in pur e cultur e Whi l e thi s ide a is partially

.

true it is almo st always th e ca s e that th e o th e r kinds o f m icro

,

o rgani s ms in th e mixtur e a re no t d e ad but simply i nactive ,

and are like ly to b e transfe rr e d with th e o th e rs and und e r

di ffe re nt co nditio ns may gai n th e a sce nd e ncy K l e b s use d .

much th e sam e m e tho d t r an sfe r ri ng th e m o st luxuriant

,

gro wths fr o m culture to cultur e until th e pur e cul tu re was

o btain e d Th e s e m e th o ds a re u se d o nly to a ve ry limit e d
.

e xt e nt at th e pr e s e nt tim e and if we had to d e p e nd up o n

,

th e m o ur kno wl e dge o f bact e ria w o uld b e limi te d and o ur

,

ide a o f th e ir li f e hi sto ry e rro ne o us .

P hysi o l o gi ca l M e tho d s Th e phy si o l o gical di ffe r e nc e s in

.
-

bact e ria have b e e n made use o f in a numb e r o f sp e cial m e th o d s .

R o b e r ts i n 187 4 and C o hn in 187 6 i so lat e d c e rtain S p o r e ;

, , , ,
 C UL TURE M E THODS AN D A P PARAT U S 87

b e aring bacilli by h e ating th e culture b e fo re incubating it to

n e arly o r quite th e b o iling p o int o f wate r B y this m e th o d .

it is p o ssibl e to is o lat e in pure cultur e such bact e ria as th e

hay ba cillus a nd th e p o tato bacillus G ayo n and Dup e tit .

d e vi se d a cultur e tub e to take advantage o f th e fact that

c e r tain bacte ria w e re much mo re m o til e than o th e r s The se .

tub e s w e r e pro vid e d with a l o ng glass spiral thro ugh whi ch

th e bact e ria w e r e o blig e d to trav e l Th e m o st m o til e fo rms
.

wo ul d ge t thro ugh th e tub e first and co uld b e r e mo ve d b e fo r e

th e sl o w e r f o rms arrive d P ari e tti us e d a disinf e ctant to
.

i nhibit th e gro wth o f ce rtain bact e ria witho ut inte rf e ring

with th e gro wth o f o the rs a nd by m e ans o f a w e ak s o luti o n
,

o f carb o lic a cid and hydr o chl o ric acid in wat e r h e f o u nd it

p o ssible to se parate B co li fr o m o th e r bact e ria and h e e ve n

.
,

claime d that h e co uld se parate thi s bacillus fro m B typh o sus . .

Th e se me th o ds are o ccasio nally us e d at th e pr e s e nt time but ,

th e difli cul tie s a re th e sam e as th o s e i ndicat e d u nd e r th e

h e ad o f F ractio nal M e th o ds i e it is n e c e s sary to start fro m

,
. .

a n u nkn o wn mixtur e and it is th e r e f o r e imp o s sibl e to kn o w

, , ,

th e r e s ult s
. Th e w e ak e r sp e ci e s may no t b e d e s tr o y e d so ,

that aft e r th e stro nge r sp e ci e s b e gin to d e ge n e rate th e o th e rs

g e t a chanc e to multiply ; and th e n again two o r
,
m o r e,
sp e ci e s
may thrive e qually w e ll .

Th Dil uti o n M tho ds

e e I n th e s e m e th o ds th e pri ncipl e
.

i nvo lve d is to di lute mate rial with a ste rile liquid as wat r ,
e ,

to such an e xt e nt that a singl e u nit o f th e mixtu r e c o ntai ns

no t m o r e than o ne bact e r ial c e ll List e r in 87 8 Obtain e d a

.
,
1 ,
88 M E TH ODS U S E D IN TH E S TUDY OF B ACTERI A
pur e cultur e o f th e lactic acid bact e ria Thi s h e di d by co unt
.

ing und e r th e mi cr o s co p e th e numb e r o f bact e ria in a dr o p Of

milk He th e n adde d suffici e nt s te ril e wate r so that any

.

particular dro p o f th e mixtur e w o uld co ntain no t o ve r a S i ngl e

o rgani s m He th e n add e d a dr o p o f thi s mi xtur e to e ach o f
.

fi v e st e ril e t e st tub e s O f milk Only o ne O f th e s e tub e s co agu

.

late d and h e naturally infe rre d that hi s o bj e ct was attain e d

,

a nd that h e had in that o ne tub e o f milk a pur e c ul tur e o f th e

lactic acid bacte ria N ae ge li and F ritz aft e rward s u se d thi s

.

m e tho d P ast e ur appli e d th e s e principl e s to th e study o f

.

y e ast but h e di lut e d his y e a st with air rath e r than with wat e r
,
.

Dry y e ast was mix e d with p o wd e r e d gyp sum and thi s mix ,

ture was th r o wn int o th e air As th e fi ne dust o f th e mixtu re

.

s e ttl e d it f e ll into flasks ab o ut th e ro o m c o ntaining a sui tabl e

,

cultur e m e dium and fro m which th e co tto n plug s had b e e n

r e mo ve d Hanso n dilute d ye ast with wat e r and di st ribute d
.

th e mixtur e i n flasks He t o o k a st e p f o rward h o w e ve r

.
, ,

by allo wi ng th e m to r e mai n u ndi sturb e d fo r a fe w days ,

and th e n r e j e cte d a s impur e tho se having mo re than a singl e

sp e ck in a b o ttl e Thi s m e tho d o f dilutio n was carri e d to th e
.

e xtr e m e by th e F r e nch sch o o l Whi l e it is p o ssibl e to ge t a

.

pur e culture in thi s way it is quit e imp o s sibl e to is o late all

,

th e bact e ria o ccurri ng in a mi xtur e wh o s e c o nt e nt is k no wn .

Use of Li quefi a bl e S o li d M e di a — Thi s was sugg e ste d by

.

th e use O f p o tat o as a cultur e m e di um S chro e d e r in 87 2

.
,
1 ,

n o tice d that th e gro wth whi ch o ccurs o n a co o k e d p o tato wh e n

it is e xp o se d to th e air is d ue to bact e ria He also use d .
 C UL TURE M E TH ODS AN D A P PARATU S 89

po tato e s to cul tivate th e co l o r p ro d ucingtb acte ria Ko ch

-
.

late r use d thi s m e dium fo r iso lating b acte ria and it was this ,

wo r k that sugge ste d to him hi s g e latin plate m e th o d whi ch

made hi m a s w e ll a s th e scie nce which h e fo u nde d famo us
, ,
.

He tri e d ni n e di ff e re nt sub stanc e s b e f o r e h e hit up o n th e use

o f g e latin . Ge latin is use d simply to give so lidity and no t ,

fo r any nutritiv e valu e Kl e b s and B re ff e l d had pr e vi o usly

.

us e d it in o bj e ct glass pr e paratio n s to pre ve nt e vap o ratio n

whil e studying th e fungi but th e y di d no t use it as an is o lat
,

ing me dium Th e me th o d is as fo llo ws Th e s te r ilize d g e la

. .

tin is me lt e d and whil e it is still warm and in a fluid state it

,

is i no culate d with th e mixture Of bacte ria pr o p e rly dilut e d ,

.

Th e m e di um i s t h e n p o ur e d o n to a glas s plat e pr e vi o usly ,

ste rilize d and pl atte d in a di sh o r unde r a b e ll jar to pro t e ct it

,

fro m co ntam inatio n Th e s e gla ss plate s w e r e fr e que ntly

.

place d o ne ab o ve th e o the r o n glass b e nch e s and sto re d in a

m o ist chamb e r F o r a go o d many y e ars th e o riginal glass
.

plate o f Ko ch has no t b e e n u se d but in its place th e Pe tri,

dish whi ch co nsists o f a fl at b o tt o m e d glass dish with a similar

,
-

glass co ve r (Fig E smarch sugg e st e d th e use o f what is

.

kn o wn as Esmarch 1011 I n this cas e th e ge latin inst e ad o f

.

b e ing p o u r e d o ut o f th e tub e is r o ll e d in a thin lay e r o n th e

inte ri o r o f th e tub e F o r de tail e d de scriptio n s o f th e s e p ro
.

ce sse s th e r e ade r is r e fe r r e d to lab o rato ry gui de s .

Th e g e latin plat e has a limit e d use o n acco unt o f th e fact

that many o f th e di se as e pro ducing bacte ria gro w r e adily

-

o nl y at t e mp e ratur e s ab o v e that at which ge latin will r e main

9 0 M E TH ODS U S E D IN TH E S T UDY OF B AC T ERI A

so lid and also b e cause many bact e ria dige st th e ge latin and
, .

th e c o l o ni e s run t o g e th e r I t wa s th e r e fo r e a co n sid e r bl e
.
, ,
a

s t e p in advanc e wh e n agar agar r e plac e d g e latin

-
Th e hi gh .

Plat e cu lt r
u e . L ig ht ar a r pr t l i fr m whi h p r
e s e e se n co o n e s o c u e cul

t
ure s can b
e o btai d Aft r L ipm a
ne . e n.

m e lti ng p o i nt o f this mat e rial make s it p o ibl e to use it at ss

a ny t e mp e r atur e suitabl e f th e g r o wth O f ba ct e r ia

or I t h as .
,

ho we v r ce r tain di sadvantage s amo ng whi ch sh o uld b e m e n

e , ,

tio ne d th e fa ct that ce rtain ba ct e r ia gro w much l e ss chara ct e r

istica l l y o n it than o n g e latin and al so b e caus e th e wat e r o f

,
 C UL TURE ME T H O DS AN D A PPARATU S 9 1

co nde n satio n whi ch gath e r s o n th e m e dium p e rmits ce rtain

bact e ria to o ve rg ro w th e o th e r s a nd cause co nfusio n .

I n th e plat e m e th o d th e si ngl e bact e ria a e s e pa rat e d and r

‘

imb e dde d in th e so lid m e dium which is suitabl e fo r the ir

gro wth and the y s o o n pr o duce as a r e sult a gro wth o r
, , ,

co l o ny varyi ng in siz e fro m a pinh e ad to a ce ntime t e r ,

ac co rding to th e natu r e o f th e bacte rium a nd th e co nditio ns

und r whi ch it is g ro wn I t is take n fo r g r a nt e d in this
e .

m e th o d that e ach co lo ny gro w s f r o m a s i ngl e i ndividual ,

o riginall y imb e dd e d in th e m e dium Thi s may o may no t

. r

b e tr ue Am o ng th e ba ct e r ia it g e ne rally is tru e but se ve ral

.
,

y e asts may cling to ge the r a nd f o r m a co lo ny th e i ndividual s ,

o f w hi ch a e o f m o r e than o ne sp e ci e s o r vari e ty
r To o ve r .

c o m e thi s diffi culty Hans o n intro duce d hi s pure culture

m e tho d whi ch c o n sis ts o f a ge lati n o r agar plat e mad e o n a
,

co ve r glass which can b e pla ce d un de r th e mi cro sc o p e and

-

e xami ne d Th e c o v e r gla s i s ma r k e d o ff int o s qua r e s whi ch

.
-
s

a e numb e r e d a nd a r e co rd i s m ad e o f th e c e ll s o ccu rr i ng in
r ,

th e vari o us squar e s a nd o nly th o se co l o ni e s are us e d fo r

,

subculturing which a re de rive d fr o m a singl e c e ll .

S u b ul tu e s
c Te st tub e cultur e are u u lly use d in mak
r . s s a

ing sub cultu r e s b e cause o f th e c o nve ni nce in handling and e

also b e cau se o f th e slight dange r f r o m co ntami natio n Th e .

g e latin stab culture is f re que ntly u e d I n thi s th e platinum

s .
,

n e e dl e whi ch h as b e e n cha rge d with th e pa r ticula r bact e ria is

plunge d to th e de pth o f th e g latin in th te st tub e so that
e e ,

th e bact e ia a e l e ft all al o ng th e line o f th e s tab

r r Th e se cul .
9 2 M E THODS U S E D IN TH E S T UDY OF B ACT ERI A
ture s are o f th e gre ate st value since minut e di ff e r e nce s in
cul tur e charact e rs a re r e ve al e d Agar p o tato blo o d s e rum
.
, , ,

and o cca si o nally g e latin a e p r e pa r e d with a slo p e d surface

, ,
r ,

and cultur e s a re made o n th e surface o f the s e kno wn as st r e ak

cultur e s in whi ch th e bact e ria are so wn o n a str e ak made by
,

th e platinum n e e dl e by drawing it th e full l e ngth o f th e sl o p e d

( se e Fig 3 0) s urfac e F luid cul tur e s in b o uill o n Dunh am s

’
. .
,

s o lutio n milk e tc a re fr e que ntly us d B acte ria are s o me

, ,
.
,
e .

tim e s gro wn in mass cul tu r e s fo r sp e cial purp o se s particula rly ,

in b o uillo n fo r th e p ro ductio n o f th e t o xin s A gre at many .

atte mpt s have b e e n made to make p e rma ne nt pr e pa r atio ns

o f ba ct e r ial cultu r e s T h e att e mpt s h o w e v e r hav e b e e n
.
, ,

larg e ly u nsu cce ssful sin ce th e cultur e s fade I n co lo r and

o th e r wi s e chang e in a r e lativ e ly sh o r t tim e .
 C HAP TER IX

M I CR O S C O PI CAL EXA MI NATI O N OF B AC TE R I A

Hi s to ry o f th e M i cr o s co p e . principl e o f th e l e ns has
Th e

b e e n unde rs to o d f ro m th e e a rli st time M agnifyi ng glass e s

e .

o f r o ck cry s tal hav e b e n f o u nd in th e r ui ns o f N im r o d

e .

Arist o phan e s 400 B C sp e ak s o f bu r ni ng sph e r e s whi ch

, ,

w e r e s o ld in th e sh o p s o f Ath e n s ; and S e n e ca in th e mi ddl e ,

o f th e first c e ntury sp e aks o f th e magni fyi ng p o w e r o f gla s s

,

glo b e s fi l l e d with wat e r I t was no t h o we ve r until th e s e ve n

.
, ,

te e nth c e ntu r y that p o w e r ful mag ni fying l e nse s w e r e u se d

fo r sci e ntific purp o se s L e e uw e nh o e k s w o r k with th e simpl e

’
.

l e ns h as alre ady b e e n r e fe r re d to Th e first co mpo und mic ro

.

sco p e was pro bably made by Hans and Z acharias J ansse n o f

M iddl e burg Ho lla nd in 5 90 I n th e c o mp o und micro s c o p e
, ,
1 .
,

as is w e ll kno wn we o b se rve no t th e o bj e ct but a n image o f th e

,

o bj e ct
. S uch a micro sc o p e c o nsists o f two sy s t e m s o f l e nse s ,

o ne with a sh o r t a nd th e o th e r with a l o ng f o cu s Th e se a e . r

th e e sse ntial pa r t s o f th e mi cr o s c o p e Th e r e a e ce rtain m e

. r

ch a ni c l applia nc e s ne c e s s ary to k e e p th e l e nse s in p o siti o n

a ,

to a ff o r d s t e adi n e ss p r o p e r illumi nati o n e tc

,
T h e inst rum e nt
,
.

with o ut th e ptical pa r ts is call e d th e stand Th e l e ns

o .

n e a r e s t th e o bj ect is kno wn as th e o bj e ctive whi le th e l e ns ,

94 M E TH ODS U S E D IN THE S T UDY OF B AC T ERI A

n e ar e s t th e o b se rv e r s e ye is kn o wn as th e e ye pi e c e o r Ocul ar
’
-
.

Two o f th e m o s t se rio us de fe cts o f l e ns e s a e what are kno wn r

as sph e ri cal and chro matic ab e rrati o n An o il imme rsi o n .

-

l e ns and an Abb é sub s ta ge c o nd e ns e r a re e sse ntial parts o f

a bacte ri o l o gical micr o sco p e .

Th e Exam i na ti o n o f B a cte ri a B acte ria may b e e xamin e d

.

unde r th e micro sco p e in a livi ng co n di tio n o r staine d The y a re .

e xami ne d in a living c o nditi o n i n what a re kn o wn as hanging

dro p pr e paratio ns Th e se a re p r e pa re d by taking a d I Op o f th e

.

fluid culture o r an e mul si o n o f th e bacte ria and putting it o n a

, ,

c o v e r glass This c o ve r glass is th e n inve r te d o ve r th e cavity

-
.
-

in a h o ll o w gro und glass slide Th e e dge o f this cavity is o il e d

.

so as to pr e v e nt e vap o r ati o n Th e bact e ria in th e ha ngi ng

.

d rOp are thus fu r ni sh e d with o pp o rtu nity to m o v e ab o ut ,

o r to r e main in a natu r al c o nditi o n Thi s pre paratio n e n

.

abl e s th e o b se rve r to d e t e rmi n e wh e the r o r no t th e re is Vital

mo v e me nt and als o to de te ct o th e r subtle p e culiariti e s so that
, ,

thi s pre parati o n is o ne o f g re at imp o r tance Th e b e ginn e r

,
.

fre que ntly finds diff culty in maki ng it b e caus e o f th e fact

i

that th e bact e ria may b e s o m e distanc e fr o m th e to p o f th e

c o ve r glass and thus th e wo rking di s ta nc e o f th e l e ns is ve ry
-

slight als o b e caus e o f th e diffi culty in r e gulating th e amo unt

,

o f light sinc e th e s e bact e ria are s o n e arly transpar e nt that a

,

little to o much light r e nde rs th e m invi sibl e .

B acte ria are als o e xami ne d in th e stai ne d co ndi ti o n by what is

kno wn as a co ve r glass pre paratio n S tai ni ng m e tho ds w e r e no t
.

int ro duce d until th e latte r part o f th e la s t ce ntury but sinc e the ir ,

 R P
MI C O S CO I CA L E XAM I NATI O N OF B AC TERI A 95

intro ductio n th e y have s e rve d a ve ry us e ful purp o s e Amo ng .

th e adva ntag e s to b e gain e d by stai ning ba ct e ria a re : it brings

o ut th e mi nut e va riatio ns in f o rm and size that w o uld

e s cap e un no ti ce d i n th e un s tai ne d p r e parati o n ; it fhcfl i ta te s

th e diff e r e ntiati o n o f ba ct e ria f r o m th e ir surr o unding s e g ,

. .

bacte ria fro m tissue ; and in so me cas e s ce rtain stains act as

di ff e r e nti al ag e nt s aidi ng in th e ide ntificati o n o f sp e ci e s
,
.

C a r mi n e fro m th e ca rdi nal ins e ct was int r o duc e d in 85 0

, ,
1 .

He mat xyli n ( camp e achy o l o gw o o d ) was us e d with gr e at

o ,
r

succ e ss in hi s to lo gy but ne ithe r o f th e ab o ve s tains are appli

,

cabl e to bacte ria since th e y give with th e se fo rms o nly a faint

tinge Th e y a re chi e fly us e d i n bact e ri o l o gy no w as co nt r ast
.

stains We ige rt in 87 7 intro duc e d th e anil in dy e s The s e

.
,
1 .

a re d e r ivativ e o f th e c o al ta r pr o duct a ni lin C H N H

-
Eh r , e s o .

lich first divide d th e s e anili n dye s i nto two g ro ups acco rdi ng as ,

th e staini ng acti o n d e p e nd e d o n th e bas ic o r acid p o r ti o n o f th e

m o l e cul e Fo r e xampl e fu chsi n th e a ce tat e o f ro sani lin

.
, ,

( t i am id o t ip h e nyl ca b ino l CmH N C H O ) d e riv e s i ts stain

r r r , 19 3 2 4 o

ing acti o n fr o m r o s anilin and is th e r e fo r e basic Amm o ni um .

picrate d e rl v e s its staining acti o n fro m th e pi ri c a cid pa r t o f c

th e m o l e cul e a nd is th e r e f o r e acid B a sic stai ns e sp e cially

.
,

active to ward nucl e ar chro mati n are th e o nly o ne s use d fo r th e

,

s taini ng o f bact e r ia Th e acid s tai ns a re e sp e cially valuabl e

.

fo r th e p r o t o pla s m o f th e a nimal o r plant c e ll Th e a nili n .

dye s have all th e co lo rs o f th e rainb o w So me o f th e m o st .

imp o rtant are

9 6 M E THODS U S E D IN THE S T UDY OF BAC TERI A
Vio l e t
M e th y l l t (H ff m an Vi l t d hlia )
-
v io e o
’
s o e , a

Ge ntian vi l t ( b nzy l i l t Py kt nin)

-
o e e -
v o e , o a

Bl ue

Me th y l n bl u (p h ny l
e e e e e ne b lu )e

V i t ria bl u
c o e

T hi ni n bl u
o e

Re d

B a i f h i n ( ba i r bin m g nt )
s c uc s s c u , a e a

S a fra nin ( f h i n G i fl ) uc s ,
ro e

B r wn
o

B i m ar k br wn ( v vin p h ny l n br wn)
s c o e su ,
e e e o

The s e dye s are usually us e d in a w e ak aqu e o us so luti o n ,

and in pr e paring th e bacte ria fo r micro sco pical e xami natio n

the y are spr e ad in a thi n film o n a gr e as e f r e e co ve r glass - -

fixe d by h e at staine d wa she d in wate r and mo unte d in C anada

, , ,

bal sam th e e xact d e tail s fo r which mus t b e go tt e n f ro m a lab

,

o rat o ry gui d e I nst e ad o f spre a di ng th e bact e ria p ro mi scu

.

o usl y o v e r th e surfac e o f a c o v e r glass a c o v e r glas s may b e - -

lo w
,

e r e d o n to th e c ul tur e and an impr e ssi o n pr e parati o n mad e ,

whi ch is fixe d and stain e d as usual Thi s is a diffi t pr e para .

tio n to mak e but wh e n pr o p e rly mad e give s th e r e lati o n whi ch

,

individuals b e ar to o ne an o th e r in th e co lo ny I n staining .

it is n e ce s sary fo r ce rtain purp o s e s to add substance s that are

kno wn as mo rdants Anilin o il w e ak alkali e s ca rb o lic acid
.
, , ,

and tannic acid are th e mo rdants mo st fre qu e ntly us e d .

Th e se m o rdants haste n th e actio n o f th e dye s and are n o ce s ,

sary with ce rtain bacte ria and are fre que ntly us e d e ve n fo r ,
 MI C RO SCO P I CAL E X AMI NATI ON OF B ACT ERI A 97

g e n e ral purp o s e s Sp e cial stains have b e e n de vis e d fo r sp e cial

.

purp o se s Lo e ffle r s alkaline m e thyl e n e blue is use d fo r stai n

.
’

ing th e B act diphthe ri a Zi e hl s carb o l fuchsin fo r th e B act

.
,
’
.

tub e rcul o sis a nd many o the r sp e cial stains are us e d fo r sp e cial

,

p urp o s e s P r o
. c e s s e s fo r d o ubl e staining hav e b e e n d e vi s e d ,

e g th o se fo r di ffe r e ntiating a S p o r e
. . f ro m th e mo the r ce ll ,

th e m e tachr o matic b o di e s fro m th e r e s t o f th e pr o t o pla s m th e ,

cap sul e fro m th e ce ll e tc Ce rtain s e l e ctive stains a re also

,
.

u se d I n particular G ram s stain w

. hich co nsi s ts e s se ntially
,
’
, , ,

o f tr e ating th e bact e ria with a particular d ye i e a nilin o il ,

. .

g e n tian Vio l e t ; with G ram s i o di n e s o lutio n (io di ne and p o tas ’

sium io dide ) ; th e n washi ng I n alco ho l With s o m e bacte ria .

th e d ye a nd th e i o di ne s o luti o n f o rm a c o mp o u nd with th e

pro to plasm o f th e ce l l which is in so lubl e in al co ho l S uch .

bacte ria whe n put thr o ugh th is pro ce ss re tain th e vio le t

, ,

co lo r Oth e r bacte ria wh e n tr e ate d with th e stain and th e

.
,

io di n e so lutio n fail to e ff e ct a co mp o und with the p ro to plasm

,

inso luble in alco ho l (i o di ne p ara ro sanil in pro te in) S uch - -

.

bacte ria whe n tre ate d by this pro ce ss a e co l o rl e ss ; o r if a

, ,
r ,

co unte r sta in h as b e e n use d a re o f an e nti re ly di ff e re nt co lo r ,

.

B y m e a ns o f G ram s stain it is p o ssibl e to di ff e r e ntiate bac

’

te ria that a re mo rph o l o gically similar and thi s s tain is o ne o f ,

th e bact e ri o l o gists m o st valuabl e a s s e t s

’
.

M o st path o g e nic mi c o co cci r e tain th e Vi o l e t c o l o r wh e n

r

tre ate d by thi s me tho d and are th e re fo r e said to b e p o sitive , ,

to G ram M a ny o f th e dis e as e pro ducing baci l l i d o no t tak e

.
-

th e G ram stain are n e gative to it whil e o th e rs a re p o sitive

, , .

H
98 M E TH ODS U S E D IN THE S T UDY OF B ACT ERI A

Am o ng th ewe ll kn o w di s e as e pro ducing bact e ria that a re

-
n -

p o sitive to G ram sho uld b e m e ntio n e d B act anthracis B act .

, .

p ne um o ni a B act tub e rcul o si s B act diphth e ria and B acil lus

,
.
,
.
,

t e tanus Amo ng th e co mmo n di se as e pro ducing bacte ria ne g

.
-

ative to G ram sho uld b e m e nti o n e d B typho su s B fe se i .

,
. r

( malig na nt oe d e ma) B act mall e i (gla nd e r s) M icr o co ccus

,
.
,

go no rrh e a and S pi rillum r e curr e n s

,
.

S o m e sp e ci e s o f bact e r ia b e a r th e nam e o f acid fas t -

bacte ria (bact e ria o f tub e rcul o sis l e p ro sy Th e pr o t o

, ,

plasm Of th e se bact e ria ho ld s th e d ye e ve n wh e n tre at e d with

stro ng d e co l o rizing age nts such as mi ne ral acid s and alc o h o ls
, .

I t r e quir e s s o m e tim e fo th e p r o t o plasm f th e bact e ria to tak e

r o

up th e stain but o nc e co mbi ne d a s with th e sp o r e d e co l o ri a

, , ,
z

tio n is difficult I t is claim e d that th e high c o nt e nt o f fat

.

acco unts fo r th e staining p r o p e r ti e s o f th e se o rga ni sms Th e .

acid fast ne ss o f B act e rium tub e rculo si s is d ue to a hi gh m o

-

l o cular alco h o l which has no t b e e n wo rk e d o ut th o ro ughly .

I t is v e ry similar to a wax acc o rdi ng to so m e writ e r s I t is .

said to b e s o luble in b o iling ab so lut e alco ho l S o m e bact e ria .

no t n o r mally acid fa s t may b e mad e S O by gr o wi ng th e m o n

-

m e dia c o ntai ni ng a larg e amo unt o f fat .

A sp e cial stai n h a s b e e n d e vi se d fo r th e flag e lla In a .

co nside rabl e numb e r o f such stai ns th e mo rdant is tannic acid ,

and an iro n salt Th e pro ce ss o f staini ng flage lla is o ne o f th e

.

m o st diffi cult staini ng pr o ce s se s that th e bacte ri o l o gi s t h as to

ma ste r a nd altho ugh th e me tho ds have b e e n g re atly simplifie d
,

in re ce nt ye ars the y still call fo r gr e at skill and p e rsi st e nce .

 R
MI C O S CO I C A P L E XAM I NATI O N OF B AC TERI A 99

Th e P i r n cipl e s i nvo l ve d in S taini ng Th e d e ve lo pm e nt o f .

th e t e chniqu e o f s taini ng has b e e n la r g e ly e mpi ri cal a nd at th e ,

p re se nt time o nly th e m o st g e n e ral o f th e pri ncipl e s invo lve d a e r

unde r sto o d I n fact it is no t k no wn whe th r th e the o ry o f

.
,
e

staini ng r e sts up o n a ch e mical o r a phy sical basi s Th e r e is .

so me e vid e nc e as wil l b e se e n to Sh o w that th e pri ncipl e

, ,

invo lve d in s taining is m e re ly a phy sical o ne Th e r e is o th e r .

e vid e nc e that at l e a s t in c e rtain cas e s th e r e acti o n b e tw e e n

-
, ,

th e d ye and pr o to pla sm may b e a ch e m ical o ne I n a g e ne ral .

way it s e e ms tru e that th e stai ni ng o f th e pr o t o pla sm i re s

late d to th e qu e stio n o f di ff usi o n and th o se co nditi o ns which

,

aid di ffusi o n aid als o in stai ni ng I n o rdi nary r o utine w o r k

.
,

th e dri e d bact e rial c e ll s are stain e d with an aqu e o u s so luti o n o f

th e d ye and c o n ce nt r at e d alc o h o lic s o luti o ns a e no t sa ti sfa c

,
r

to ry fo r staini ng d ri e d bacte ria Th e re is di ff usio n b e twe e n th e

.

dri e d bacte rial ce ll and th e wate ry so luti o n and no ne in th e ,

case o f drie d bact e ria and alco h o l I f th e bacte ria a e fi r s t . r

we t th e n th e y wil l stain with th e alc o h o lic so luti o n s Th e

,
.

maj o rity o f bacte ria stain r e adi ly with th e o rdinary aqu e o us

s o luti o n o f th e dye s To stain ce rtain o f the m ho we ve r it is
.
, ,

n e ce s sary to r e so r t to S p e cial me ans S o me o f th e bact e ria .

,

as we have se e n have c e rtain fatty o r waxy sub s tanc e s in the ir

,

c e ll walls whi ch re nde r the ir s ta i ni ng by wat ry s lutio n o f th e

,
e o

dye s difli cul t o r we ll nigh imp o s ibl e Wh e n th e y a re o nce

-
s .

staine d ho we ve r it is d ifli cul t to d e stai n the m F o r thi s

, ,
.

p pu r o se eo
v n the e mi n e r al acid s may b e u s e d Th se bacil l i a re . e

kno wn a s th e acid fast bacilli I n o r de r to stain th e m it is

-
.
I OO M E T HODS U S E D IN THE S TUDY OF B ACTERI A
n e ce ssary to add m o rdants to th e stain Th e p rincipl e l n .

vo lve d h e re is th e same as that use d in dye i ng cl o th F o r .

staining purp o se s th e m o rdants u sually u se d a e th e iro n salts r ,

w e ak alkali e s carb o lic acid anilin Oil a nd tannin Th e se

, , ,
.
'

m o rdants e nabl e o ne t stain e ve n th e m o st r e sis tant bacte ria

o .

Whe n the y are o nce staine d it is p o s ibl e to r e mo ve th e s tain s

fr o m o the r o bj e cts in th e sp e cim e n l e ss acid fast than th e m ,

-

s e lve s by m e ans o f s o me d e co l o rizing ag e nt such as th e m ine al

, ,
r .

acids o rgani c acids alco ho ls e tc I n thi s way it is po ssibl e

, , ,
.

to s e cur e d o ubl e stains in whi ch o ne ki nd o f bact e ria i s s tain e d

,

o ne c o l o r and o th e r bact e ria an o th e r c o l o r S uch a d o ubl e .

stain is us e d fo r th e diffe re ntial diagn o sis o f ce rtain bacte ria .

Th e ba cillus o f tub e rcul o si s fo r in s tanc e is s tain e d in sputum

, ,

by u sing a do ubl e stain by m e ans o f which th e B act tub e rcu

,
.

l o sis is c o l o r e d a d e e p crims o n whil e th e r e s t o f th e pre parati o n

,

is blue o r gre e n in co l o r Th e B act tub e rcul o si s has thi s acid

. .

fast pro p e r ty in co mmo n with a numb e r o f o th e r bacte ria but ,

it is u sually p o ssibl e by m e ans o f slight variati o ns in th e

,

t e chnique o r by s o me o the r m e ans to di ffe r e ntiat e th is g e rm

, ,

f ro m th e o the r acid fa st bacill i De p e n di ng up o n principl e s

-
.

simi lar to tho se n o te d ab o ve a do ubl e stain fo r e ndo sp o r e s

,

h as b e e n d e vis e d and is fr e qu e ntly u se d .

The re a re c e rtain o the rstains whi ch may o r may no t indicat e

that th e pr o ce ss o f staining is a che m ical o ne but the y d o ,

indicate at any rate that th e r e is s o m e diff e re ntiati o n in

structur e in th e vari o us parts o f th e c e ll S o m e b a cte ria wh e n .

s tain e d with a w e ak s o luti o n o f m e thyl e n e blu e giv e th e b i

 R P
M I C O S C O I CA L E X At fi i
I
N A T ON
“

OF B AC T ER I A 10 1

p o lar stain Oth e r bacte ria wh e n stain e d with me thyle ne blu e

.
,

o r similar s tai ns pr e se nt gran ul e s o f vari o us siz e s withi n th e

,

c e l l pro to plasm usually in a p o la r o r e quat o rial po itio n

, s .

Th e se p e culiar staining r e actio n s may in di cat e a variatio n in

th e ch e mical c o mp o siti o n o f th e pr o t o plasm o r th e variati o ns
,

may b e m e r e ly physical o ne s I n G ram s stain th e principl e

.
’

invo lve d is undo ubte dly o f a che mical nature .

 P A RT IV . TA X O N O M Y

C HAP T ER X

CLAS S I FI CATI ON OF B ACTERIA

In tro ducti o n Th e r e is co n side rabl e co nfusio n at th e
.

pr e s e nt time and th e re has b e e n fo r y e ars in th e n o me nclatur e

, ,

and classificatio n o f bacte ria This co nfusio n has larg e ly

.

arise n o n acco unt o f th e fact that th e maj o ity o f th e w o rk

r

o n th e subj e ct o f bact e ri o l o gy h a s b e e n d o n e by path o l o gi s ts

and o the rs inte re st e d in th e r e sults o f th e Vital activity o f

bacte ria rath e r than th e ir e xact p o sitio n amo ng th e plants .

Ve ry little syst e matic wo r k has b e e n do n e o n th e bact e ria

by b o tanists .

Th e first o bs e rv e rs o f bact e ria w e r e p o sitive that th e y w e r e

ani mals and w e r e inclin e d to b e li e ve that th e y b e l o nge d to th e

infus o ria Th e y have since b e e n place d in th e plant kingdo m
.
,

alth o ugh th e y p o s se ss many things in co mm o n with ce rtain

animals .

I t may b e stat e d at th e o uts e t that th e r e are many cl assifi ca

tio ns o f th e bacte ria F o rme rly e ve ry autho r o f a t e xtb o o k

.

o n th e subj e ct o f bacte rio l o gy made a ne w cla ssificatio n .

Th e w o rk e r s in th e va ri o us fi e ld s al so mad e cla ssifi cati o ns ,

as ,
fo r e xampl e th e b o tanists path o l o gists agriculturists
, , , ,

10 2
 L
C A S S IF I CA I O T N OF B ACT ERIA 10
3

ch e mists bre we rs e tc Until r e ce ntly no att e mpt at uni

, ,
.

fo rmity has b e e n made Ob vi o usly any e ffo rt at th e co r

.
,

r e lati o n o f th e vari o us pro p o s e d classificatio n s in a b o o k o f

this sco p e wo uld b e imp o ssibl e Th e r e fo re th e a imwill b e
.

to p r e s e nt fo r d ue c o nsid e r ati o n o nly o ne classificatio n

which will give th e cl e ar e st ide a o f th e characte ristics o f th e
bacte ria As o ur d e finite kno wle dg e o f th e bacte ria ih
.

cre as e the s e classificati o ns must ne c e ssarily b e mo difi e d .

I t is an e stabli she d fact that th e fo rm shap e a nd mann e r , ,

o f gr o wth o f a d e fini t e bact e rium may b e t e mp o rarily m o difi e d

by th e e nviro nme nt whi ch surro unds th e o rganism Th e ide a .

that ne w and we ll d e fi ne d S p e cie s may b e p ro duce d by m o difi

—

catio ns o f th e e nviro nm e nt was h e ld fo rm e rly by so me bacte ri

o l o gi s ts . Thi s was th e the o ry o f pl e o mo rphism I t has since .

b e e n pro ve d that th e o rga ni sm whi ch has appare ntly assum e d

a ne w fo rm will re tu rn to i ts o riginal shap e wh e n plac e d in th e
r e qui ite e nvir o nm e nt I t was als o he ld fo r so me tim e rath e r
s .

t e nacio usly that bacte ria had th e p o we r o f assuming ne w

physio l o gical functio ns To thi s p o we r th e t e rm pl e o ge ny
.

wa s appli e d I t h a s b e e n f o und in this cas e as in th e

.
,

fo rme r that th e bacte rial ce ll will r e tu r n to n o rmal wh e n a

,

n o rmal e nvir o nme nt is pro vide d (se e Chap I I I ) . .

Th e re is no do ubt but that th e last clas sificati o n o f M igula is

as accurate and syste matic a s has b e e n made Thi s cl a ssifi ca .

ti o n h as b e e n ado pte d by th e l e ading Am e rican bacte ri o l o gi sts .

I t se e m s advantag e o u s h o we v e r to mak e a slight r e arrang e

, ,

m e nt o f th e classificati o n A divi sio n can b e made fo r suffi

.
10 4 TAX O N OMY
i t r e aso ns into l o we r a nd hi gh e r bacte ria o r what are
c en ,

t e rm e d by F ische r Hapl o b a cte ria and Tricho bacte ria M igula

, .

divide s all bacte ria into two o rd e r s : th e Eubacte ria o r tru e ,

bacte ria and th e Thi o bact e ria o r sulphur bact e ria Unde r
, , .

th e Eubact e ria M igula place s f o ur famili e s Vi : C o ccace a

, ,
z .
,

B a cte ri ace a Spiril lac e a

,
a nd
,
C hl am yd o b a cte ri ace a We .

b e li e ve that a simpl e r arrang e m e nt can b e mad e by pl a c mg th e

famili e s Co cca ce a B acte ria ce a and Spi rillac e a und e r th e
, ,

sub o rde r o f th e Hapl o b a cte rina ( F isch e r) o r l o w e r bact e ria ,

and th e Chl am yd o b acte ri ace a unde r th e sub o rde r Tri ch o b ac

t e ri a ( Fisch e r) o r high e r bact e ria Th e vari o us ge n e ra in
.

the se famili e s a s give n by M igul a are no t change d .

The r e a re many sp e cie s o f bact e ria whi ch p o sse ss de finit e ,

whil e o th e rs p o sse ss variabl e Cha r acte ri stics I t is p o ssibl e

,
.

to plac e th e m in d e fi nit e g e ne ra a nd to c o n s titut e di s tinct

sp e cie s I t sho uld b e r e m e mb e r e d h o we ve r that the r e is uh

.
, ,

d e r c e rtain c o ndi ti o n s a s t r iki ng simil a r ity in f o rm a nd physi

o l o gical activity am o ng di ff e r e nt sp e ci e s o f bact e ria Fo r .

e xampl e c e rtain bacilli a re so sh o rt that th e y app e ar at tim e s

,

to b e c o cci . I t will b e e a sil y s e e n th e r e fo r e that th e r e is

, ,

always di fficul ty in th e clas sificati o n o f so m e o rganisms Th e .

b e st cla ssificatio n can b e made o nly wh e n all th e vari o us

sp e ci e s o f bact e ria are subj e ct e d as n e arly as p o ssibl e
,
to th e ,

s am e c o nditi o ns o f te mp e ratur e light ga se o us e nvir o nme nt

, , ,

f o o d e tc
, .

Cl a s s ifi ca ti o n o f M i gul a M o difi e d ) — Th e bact e ria are phy

.

co ch ro m e fr e e schi z o myc e t o u s pla nt s w hi ch di vid e

-
in o ne ,
 L F
C AS S I I CA I O T N OF B ACTERI A I O 5

two , thr e e plan e s R e pro ductio n take s place by ve ge tative

or .

mul tipli catio n (fi ssio n) R e sting stage s in th e f o rm o f e n

.

d o sp o re s are pr o duc e d by many sp e ci e s M o tility is n o te d in

.

so me ge n e ra and thi s i d ue to flage lla I n B e gg a to a and

,
s .
i

Spiro ch a ta th e o rgans o f l o c o mo ti o n a re no t de finite ly

kn o wn .

I . ORDER : E UBAC TE R I A (Tru e B acte ria) .

Th e c e lls are de vo id o f any n ucle us (Z e ntral ko rpe r) and

fre e f r o m sulphur and bacte ri o purpur n c o l o rl e ss o r faintly i
,

co l o re d .

I . S ubo rde r : Hapl o bacte ri nce ( Lo we r B acte ri a) .

I . F amily : C o cca ce a (Z o pf) M ig .

Th e ce lls are gl o bular wh e n in a fre e state but in th e vari o us ,

s tag e s o f divisi o n app e ar s o m e what e lliptical A fe w sp e ci e s in .

this famil y are mo til e C e ll divi si o n tak e s plac e in all di re c

.

tio ns o f space F re que ntly th e ce lls re main attach e d to ge the r

.
,

a nd und e r th e s e c o ndi ti o ns u s ually sh o w s o m e flatt e ni ng o f th e

c e ll at th e p o int o f junction with th e ce ll n e xt to it .

Ge nus : S t r e pto co ccus B ill ro th .

Th e ce ll s are gl o bular and d o no t p o ss e ss any o rgans o f

lo co mo tio n C e ll div isio n take s place in o nl y o ne plane
.
.

Usually th e c e ll s r e main unit e d t o g e th e r aft e r divi si o n p ro ,

d ucing chai ns o r di pl o co ccus fo rms N o e nd o sp o re s hav e

.

b e e n no te d .
10 6 TAX ON OMY
Ge nus : M icro co ccus (Halli e r) C o hn .

Th e ce lls a re gl o bular and d o no t p o ss e ss any o rgans o f l o co

m o tio n C e ll divisio n take s place in two plan e s at right angl e s
. .

I f th e c e lls r e mai n attach e d t o g e th e r aft e r c e ll divi si o n ,

m e rism o p e d ia plat e s a re f o rm e d Th e plat e s give th e ap

.

p e a ra nc e o f a r e gular flat ma s s o f c e lls N o e nd o sp o r.e s hav e

b e e n no t e d in this ge nus .

G e nus S arcina Go o d sir .

Th e ce lls are gl o bular and d o no t p o sse ss any o rgans Of l o co

m o ti o n C e ll divi si o n take s place I n thre e plan e s all p e rp e n
.
,

di cul a r and at right angl e s to e ach o th e r I ts c e l ls r e main .

attach e d a ft e r divisi o n cub e l ik e packe ts are fo rme d Th e

,
-
.

co mp o siti o n o f th e m e di a so me tim e s pr e ve nts this typical

cub e f o rmatio n .

G e nus : P lan o co ccus M igula .

Th e c e ll s are gl o bular C e ll divisio n tak e s place in two

.

plan e s at right angl e s S imilar to ge nu s M ic ro co ccus Th e ce lls .

o f this g e nu s a re m o til e p o sse ssi ng o ne o r two lo ng flage lla ‘

.
,

N o e nd o sp o r e s a re pr o duc e d in this g e nus .

Ge nus : P lan o s arci na M igula .

Th e c e lls a re gl o bular C e ll divisio n tak e s plac e in thre e

.

plan e s as in S arcina C e lls a e m o til e havi ng o nl y o ne fl age l

. r ,

lum o n e ach C e ll s usually r e main uni t e d in two s and in te t

.

rads and s e ldo m fo rm pack e ts as S arcina .

 L
C AS S I F I CATI O N OF B AC TERI A 10 7

II . F amily : B acte riace a M igula .

Th e ce lls are cyli ndrical in shap e The y vary in l e ngth .

fro m sho rt almo st sph e rical b o die s to ve ry l o ng rq

, C e ll .

divisio n take s place s in o ne di re ctio n in a plan e p e rp e ndicular

to th e l o ng axis o f th e c e ll S o m e o f th e m e mb e rs o f th i s
.

famil y r e main attache d to ge th e r fo rming thre ads while , ,

o th e r s s e pa r at e fr o m e ach o th e r s o o n aft e r fissi o n .

G e nus : B acte rium Ehr e nb e rg .

Th e ce lls cylind rical Of l o nge r o r sho rte r l e ngth

a re ,
.

Th re ads a re fr e que ntly f o rm e d Th e ce lls d o no t p o sse ss any

.

o rgans o f l o c o m o ti o n E nd o sp o r e s a re p r o duc e d in s o m e
.

fe w sp e ci e s but in th e maj o r ity no s uch f o rmati o n o ccurs

,
.

I t is p o s sibl e that e nd o sp o r e fo rmati o n o ccurs o nly und e r

ce r tain e nviro nme ntal co nditi o ns .

G e nus : B acillus C o hn .

Th e ce lls a re cylindrical o f l o nge r o r sho rte r l e ngth Th e

, .

r o d s a re s o me tim e s o val in S hap e C e ll s a e m o til e and p o s

. r

s e s flage lla which a re di st ribute d o ve r th e e ntir e surface

s .

Endo sp o r e fo r mati o n o ccur s with mark e d r e gul a rity Th e .

bact e ria in thi s g e nu s are m o til e o nly du ri ng c e r tain p e ri o ds

o f th e ir lif e Thi p e ri o d varie s gre atly in l e ngth and o ccur s
. s

o nly i n th e v e g e tativ e stag e .

Ge nu s : P s e udo mo nas M igul a .

Th e c e lls are Cylindrical o f l o ng e r o r sho rte r l e ngth Th e

,
.

ce lls are mo til e and p o sse ss p o lar flag e lla Th e s e flag e lla .
10 8 TAX O N OMY
may vary fro m o ne to tw e lve in numb e r Th e fo rmatio n o f .

e ndo sp o r e s in thi s sp e ci e s i s cla im e d by s o m e I f it o ccur s it .

is e xtre me ly ra r e Occasio nally ce rtain sp e ci e s in this g e nus

.

fo rm th e mse lve s into thre ads o r chains .

III . F amily : Spirillace a M igula .

Th e ce ll s are w o und in th e f o rm o f a spiral o r r e pre s e nting

th e p o rti o n o f a turn o f a spiral I n th e latt e r cas e if th e c e lls
.
,

r e main attach e d to g e th e r in th e fo rm o f a thr e ad a full spiral ,

o f s e v e ral turns is pr o duc e d C e ll di visi o n tak e s plac e in o nly

.

o ne di r e cti o n o f spac e and thi s is transv e rs e to th e l o ng axis

,

o f th e c e ll .

Ge nus : Spiro s o ma M igul a .

Th e ce lls are rigid and b e nt in th e fo rm o f spiral s Th e .

m e mb e rs o f this ge nus are a s a g e n e ral r ul e qui te larg e Th e .

ce lls may b e f r e e o r unit e d to ge th e r into small g e latino us

mas se s S o me o f th e ce lls individually a re surro unde d by
.

a g e latin o u e nve l o p e while o th e rs a re f re e

s .

G e nus : M icr o spira S ch rOte r .

Th e rigid sho rt and b e nt similar to a Co mma

c e lls a re , , .

Wh e n th e c e lls are unite d to ge th e r S shap e d thr e ads are ,

-

fo rm e d Th e ce ll s a re m o tile p o sse s sing u sually o ne fl age l

.
,

lum and rar e ly two o th re e flage lla Th e se flage lla a e ab o ut

r . r

th e sam e l e ngth as th e c e ll N o e ndo spo r e s are f o r m e d

. .

S o m e writ e rs mak e no distincti o n b e tw e e n M icr o spira and

 L
C AS S I F I CA I O T N OF B AC T ERIA 10 9

Spirillum . Th ename Vibrio has also b e e n applie d by so m e

write rs to thi s g e nus .

Ge nu s : Spirillum Ehr e nb e rg .

Th e ce ll s a e rigid usually l o ng and f o rming l o ng scr e wlik e

r , ,

thr e ad s o r in s o m e ca se s o nl y p o rtio ns o f a spiral turn

, , ,
.

C e lls are m o til e and p o sse ss a tuft o f flag e lla at th e p o l e Th e .

flag e lla may o ccur at b o th e nds o f th e spiral and th e y vary ,

gr e atly in numb e r Endo sp o r e fo rmatio n has b e e n o bs e rve d

.

in so m e sp e ci e s .

G e nus : Spiro cha ta Ehr e nb e rg .

Th e ce lls are fl e xibl e spirals ve ry thin and l o ng No ,

.

flage lla are pre s e nt Th e se bacte ria m o ve by r o tati o n simi

.

lar to a scr e w and also by late ral mo tio n similar to a snak e

,
.

Th e l o co m o tiv e o rgans if pr e s e nt a re no t kn o wn N o e ndo .

sp o re s are pr o duce d .

II . S ubo rd e r : Tricho bacte ri na ( Hi ghe r B acte ri a) .

F am il y : C hl a m yd o b acte ri a ce a M igula .

Th e ce lls are cylind rical are unit e d in thre ads and sur
, ,

r o unde d by a sh e ath R e pr o ductio n tak e s place by me ans

.

o f m o til e a nd n o nm o til e g o nidia Th e se go ni dia arise di re ctly

.

fro m th e ve ge tative ce ll s and witho ut any r e sting stage p ro

duce ne w thre ads o f c e ll s .

Ge nus : Chl am yd o th rix M igula .

Th e ce ll s are cylindrical n o nmo til e and arrange d in um

, ,

branch e d thre ads and surr o unde d by a sh e ath o f varying

I IO TAX ON OMY
thi ckn e ss in di ff e r e nt sp e ci e s b e ing th e sam e diam e te r at
,

ap e x and base R e pro ductio n tak e s plac e by m e ans o f g o nidia

.
,

whi ch are r o und and ari se dire ctly fro m th e Ve ge tative c e ll .

Thi s g e nus is call e d Le pto thrix by Kli tzing and S te p to thrix

by Co hn .

G e nus C r e no thrix C o hn .

Th e ce ll s a re unite d to ge the r into fil ame nts which are un

branch e d Th e filam e nts gradually e nlarge to wa r d th e fre e
.

e nd,
thus making a di s tinctio n b e tw e e n th e ap e x and base .

Th e sh e ath whi ch c o v e rs th e filam e nts is thi ck and o ft e n

b e co m e s infiltrat e d with th e hydro xid e o f iro n aft e r b e ing

cast o ff in wat e r in which th e r e is a large amo unt o f iro n .

R e pr o ducti o n tak e s plac e by th e fo rmati o n o f r o u nd g o nidia

which are fo rm e d in th e b e ginning by divisi o n p e rp e ndicular

to th e l o ng axi s o f th e c e ll and lat e r by divi si o n in thre e d ire c

ti o ns o f space Onl y o ne o r p o ssibly two sp e ci e s can b e

.

plac e d in this ge nus .

G e nus : P h ragmi di o thrix E ngl e r .

Th e c e ll in th e b e gi nning f o rm unbranch e d thre ads C e ll

s .

divisi o n take s place in thr e e dire cti o ns o f spac e thu s fo rming,

withi n th e sh e th a mass o f c e lls L ate r th e se c e lls may

a .

bu r st th ro ugh multiply and fo rm bran ch e s aft e r acquiring

, ,

sh e ath s Th e sh e ath in this g e nus is qui te thin and can

.

scarc e ly b e se e n .
 L
C A S S IF I CA I O T N OF B ACT ERI A I I I

G e nus : S ph a ro til us K u tz ing, 183 3 ( Clad o th rix C o hn) .

Th e ce ll s a e cylind rical and th e th re ads a e surro unde d by

r r

she ath s Di cho to mo us b r an chi ng is p re se nt and the r e is no

.
,

diff e re ntiatio n in si e b e tw e e n th e ap e x and ba se o f th e thr e ad

z .

R e pr o du cti o n tak e s pla ce by m e a ns o f g o nidia which swa r m

to ge th e r withi n th e ce ll Th e s e go nidia bur st o ut o f th e

.

ce lls attach the m se lve s to so m e Obj e ct and g ro w into ne w

, ,

th re ads Th e go nidia a e e ndo w e d with flag e lla which are

. r

attache d to ward th e e nd and b e lo w th e p o l e .

II . OR DER THI OBACTE R I A ( S ulphur B acte ria) .

Th e ce p o sse ss a ny nucl e us a nd co ntain sulphur

ll s do no t .

Th e c e ll s a e co l o r l e ss o r pigm e nt e d r o s e vi o l e t o r re d by
r , ,

bacte ri o purpu rin Th e c e lls a re n e ve r pigme nte d gre e n

. .

I . F amily : B e ggi a to ace a Tr e visan .

F ilame nto us ba cte ria which d o no t co ntain bacte rio purpurin .

Th c e lls c o ntain sulphu r g r an ul e s

e R e pr o ducti o n tak e s plac e .

in o ne dir e cti o n o f S pace .

Ge nus : Thio thrix Wino gradsky .

Th e ll s a e no nmo til e a nd th e thr e ads are attach e d to

ce r

so me o bj e ct Th e thre ads a e su rro unde d by a de licate

. r

sh e ath a nd th e c e ll s c o ntain s ulphur g r anul e s Go ni di a a e . r

pro duce d at th e e nd o f th th r e ads Th e se go nidia a re e .

mo til e and finally attach the mse lve s to so me o bj e ct and , ,

 I 2 TAX O N OMY
acco rding to so m e autho rs b e nd at right angle s in th e
, m l d dl e

and gro w into ne w thr e ads .

G e nus : B e ggiato a T r e visan .

Th e thre ads are no t sur ro und e d by a she ath and are fo rm e d

o f flat c e lls Th e c e lls a r n t attach e d
. e Thi s ge nu s mo ve s
o .

by m e ans o f an undulati ng m e mb r ane simila r to Oscil laria .

As th e o rganism m o v e s it r o tat e s o n i ts l o ng axis and swi ngs

,

its fr e e e nds Go nidia are unkno wn and r e pro ductio n take s

.
,

place by a divi sio n and se parati o n o f th e th e ads r .

II . F amily : R h o d o b acte ria ce a ( Wi no gra d skys

’
cl assifi ca

tio n ,
Artificial) .

Th e c e lls co nta in bact e ri o purpurin and o n thi s acco unt may

b e re d r o s e o r Vi o l e t
, , S ulphur granul e s may als o b e ih
.

clude d withi n th e ce l ls .

I S ubfamily : Th io ca p sa ce a .

Th e c e lls are unit e d int o c o l o ni e s . Ce l l divisi o n tak e s place

in thr e e dir e cti o ns o f space .

G e nus : Win o gradsky

Thi o cystis .

Th e c o l o ni e s are small c o mpact and e nv e l o p e d e ith e r

, ,

singly o r in gr o ups by a ge latino us cy st Th e co lo ni e s are .

also capabl e o f bre aking up and th e ce lls m o v mg ab o ut .

G e nus : Wino gradsky

Thio cap sa .

Th e c e ll s are gl o bular in shap e and spr e ad o ut o n a sub

stratum in flat co l o ni e s The se co lo ni e s are surr o unde d by a

.
 L
C A S S IF I CA I O T N OF B ACT ERI A I I 3

c o mm o n ge latino us se c r e ti o n simil ar to a capsul e . Th e ce lls

a re n o nm o til e .

G e nus : Thi o sa rcina Win o g r adsky .

Th eco l o ni e s fo rm packe ts sim ilar to th e ge nus S arcina o f

th e Eubact e ria T h e c e lls a re n o nm o til e
. .

II S ubfamily L am p ro cystace a .

ce lls are fo rme d into fami lie s

Th e . C e ll di visio n take s place
first in thr e e th e n in two dire ctio ns
,
of space .

Ge nu s : La m p ro cystis S ch ro te r .

Th e ce ll s in th e b e ginning are so lid th e n ho ll o w b e co ming , ,

p e rfo rat e d lik e a ne t Th e y se parate into small gro ups and

.

b e co me m o tile .

I I I S ubfamily Thi Op e di a ce a .

ce ll s a re unit e d into co lo ni e s
Th e . Ce ll divisi o n tak e s plac e
in two dire ctio ns o f space .

Ge nus : Thio p e di a Wino gradsky .

Th e famili e s are fo r m e d similar to tub e s and a e co mp o s e d r

o f c e ll s arrang e d in f o urs a nd capabl e o f m o tility .

I V S ubfa m ily Am oe b o b a cte ri ace a .

Th e ce ll are unite d int o co l o ni e s

s . C e ll di visio n tak e s p lac e
in o ne di r e cti o n o f spac e .
1 14 TAX ON OMY
G e nu s : Am oe b a cte r Win o gradsky .

Th e ce lls a re unite d into co lo ni e s and afte r di visio n in o ne ,

dire cti o n o f space r e main attach e d t o ge th e r by thre ads o f

pro to plasm Th e co l o nie s p o sse s s amoe b o id mo til ity Th e
. .

ce lls change fo rm by co ntracti o n and th e spre a di ng o ut o f

th e pr o t o plasm .

G e nu s : Thi o th e ce Win o gradsky .

Th e co l o ni e s a re incl o se d by a thi ck ge latin o us cyst Th e , .

c e ll s are capabl e o f mo ving and are ve ry l o o se ly surro unde d

by a co mmo n ge latin .

G e nus : Thio di ctyo n Win o gradsky .

Th e co l o ni e s a re s o lid no nmo til e and co nsist

, ,
of small ce lls
whi ch are pr e s se d to ge th e r .

V S ubfamily C h ro m o tia ce a .

Th e ce ll s a re fre e and capabl e at all time s of mo tility .

Ge nu s : C hr o matium P e rty .

Th e ce ll s a re mo de rat e ly thick e lliptical , ,

or cylindric e l l ip -

tical in shap e .

G e nus : R h abd o ch ro m a tium Win o gradsky .

Th e ce ll s are fre e ro d S hap e d o r spindl e f o rm Th e y

,
-

,
.

p o sse s s flage lla o n th e p o l e s and are m o til e at all tim e s .

G e nus Thi o spiril l um .

Th e ce lls are fre e co ntinually mo til e and spirally twiste d

, ,
.
 CHAP T ER X I

TILE R ELATI ON S HI P OF B AC TER I A

In tro d ucti o n Th e b o undary b e twe e n th e animal and

.

ve ge table kingdo ms if such th e r e b e is no t de fi nite Th e r e

, , .

h as b e e n much c o nfusi o n i n re ga d to th e accurat e di ff e r e ntia

r

tio n o f th e s e kingdo m s since th e tim e o f th e firs t r e co rds o f mic

ro sco p i c l ife
. Th e indicati o ns a e that it will b e s o m e tim e b e f o r e
r

s ci e nti sts will d e fi nit e ly se ttl e thi s pro bl e m and b e abl e to state
in ce rtain i ns tanc e s that a particular l o w f o rm o f lif e is a pla nt

o r an animal . We r e f e r o bvi o u sly to tho s e ani mal and plant

, ,

fo rms whi ch a re o f th e simpl e st structure and which cl o s e ly

r e se mbl e e ach o th e r physi o lo gically and m o rph o lo gically .

I n all pr o bab ility acc o rding to r e c e nt inv e s tigati o ns this

, ,

pro bl e m will b e gre atly e lucidat e d by th e che mis ts wh o o n th e ,

p e rf e ctio n o f th e ir t e chni q ue will fi nd that th e main diff e r

,

e nce s b e twe e n p l a nts and a nimals a re tho s e o f che mical

structur e .

Th e t e rms ani mal and pla nt w e r e d e vi se d y e ars ago

b e fo r e th e S ingl e ce ll e d fo rm s o f life which p o ss e ss similariti e s

-

to b o th gro ups w e r e k no wn I t is in a m e a sur e s e ns e l e ss to

.

try to apply th e se t e rms to o rganisms fo r which the y w e r e

n e ve r int e nde d Th e t e rm s as have b e e n s tate d by vari o us
.
,

write rs w e re inte nde d to apply to such co ntrast s a s o ak tr e e s

,

1 15
I I 6 TAX O N OM Y

and e l e phants and w e re n e ve r to b e applie d to minut e plants

,

and animals Th e r e are ce rtain things whi ch are co mmo n

.

to plants and ani mal s d o wn to th e v e ry simpl e f o rm s Th e re .

is undo ubt e dly a g r adual transitio n fr o m o ne fo rm to th e

o th e r and c o ns e qu e ntly no dividing lin e can b e mad e b e
,

twe e n th e two Th e two gro ups gradually appro ach e ach

.

o th e r i th e ir structure and physi o l o gical pro ce ss e s and finally

h ,

co al e sce .

C o nc e ive th e n a gr o up o f mi cr o o rganisms at th e p o i nt
wh e re th e plant and ani mal kingdo m s unite ; a gr o up which
i nclud e s animal s and plants O f th e simpl e st anat o mical
st ructu re ; a g ro up in which diff e r e ntiati o n into o ne o r th e
o th e r o f th e o rgani c kingd o ms is difficult a nd ba se d up o n

what littl e is k no wn ab o ut th e ir ch e mi stry anato my and , ,

phy sio l o gy Thi s g ro up w o uld in clud e ce rtain o f th e p ro

.

to o a am o ng th e a nimals and c e rtain o f th e alg a and fungi

z

amo ng th e plants We can mak e no d e fi nite b o undary fo r

.

thi s gro up Ha e ck e l has applie d th e nam e p r o tista to this

'

gro up o f simpl e fo rms o f lif e .

B acte r ia De fi ne d — T O this arbit r ary gr o up o f simpl e

.

ce ll e d fo rms th e class o f micro o rganism s which are d e sig ,

na te d as bact e ria b e l o ng Th e bact e ria a re th e small e st o f

,
.

all kno wn living o rganic b o di e s Th e Siz e o f co urse vari e s .

, ,

with th e sp e ci e s but is fai rly co n stant within th e limit o f th e

,

sam e sp e ci e s .

I t has b e e n d e cid e d by car e ful sci e ntific inv e stigati o ns that

th e bact e r ia taking th e wh o l e class o f o rga nisms int o co n

,
 TH E RELATI ON S HI P OF B ACTERI A I I 7

sid e ratio n S h o w m o r e p o ints o f re s e mblance to th e plants

,

than to th e animals The y have b e e n f o und to re s e mbl e

.

c e rtain o f th e l o w e r pl ants v e ry cl o s e ly Th e bacte ria a re

'

no wd e fi ne d to b e m i c o s co pi c uni ce l l ul a no nnucl e a te d p l a nts

'
r ,
r,
,

d e vo id of chl o o p hyl l and e p ro du ci ng by di e ctfi ssi o n I n so m e

r r r .

o rd e s the f
r m a ti o n of sp e ci al
or e p o ducti ve ce l l s kno w
r r n as

a o ccu rS o m e auth o rs hav e stat e d that r e pr o ducti o n

.

als o tak e s plac e by th e f o rmati o n o f e nd o sp o r e s This pro p o .

siti o u h as b e e n di scusse d in ano the r chapte r .

B act e ria a e th e m o st wid e ly distribut e d and m o st num e r

r

o us Of all l iving o rga nisms Th e surfac e o f th e e arth a nd th e

so il to an av e r age o f fo ur fe e t co ntain bacte ria Th e w

.

ate rs .

o n th e surfac e o f th e e arth c o ntain i nnum e rabl e bact e ria .

Th e air with th e p o s sibl e e xc e pti o n o f that o v e r larg e b o di e s

,

o f wat e r and in hi gh altitud e s c o ntain varying numb e rs o f

,

bacte ria and many sp e ci e s N o fami ly o f plants co mpare s

.

with th e bact e ria in imp o rtance .

P o ints o f R e s e m b l a n ce to O th e r F o rm s o f Life Th e .
-

fo ll o wing facts will se rv e to give an ide a o f th e ge ne ral p o ints

o f r e se mblanc e o f th e bact e ria to th e l o w plant s a nd animals .

I t will b e s e e n that th e y r e se mbl e th e kn o wn plants m o r e

clo se ly than th e animal s and co ns e que ntly are place d in th e

, , ,

lo we st o rde r o f this ki nd go m .

.I I n s o m e S p e ci e s o f bact e ria a thin limiting m e mbran e , ,

which is r e lat e d to ce llul o se o r h e mice llul o s e ch e mi cally is , ,

pre se nt and in thi s particular th e y re se mbl e plant ce lls

,
.

Ho w e ve r r e c e nt inve stigatio ns hav e sh o wn that th e c e ll wall

,
I 18 TAX ON OMY
in a gr e at many o f th e l o we r bacte ria r e se mble s as much if ,

no t m o r e th e chitin o f c e rtain animal c e ll wa l ls than it d o e s

,

th e c e llul o s e o f plants .

2
. Th e f o rm o r shap e o f th e bact e rial c e ll is v e ry similar to

so me o f th e l o w plants such as th e blue gre e n alga Th e

,
-
.

bacte ria in s o me case s fo rm filame nts as plants o f a l o w ,

o rd e r d o .

3 Th e r e is a s imilarity b e tw e e n c e r tain o f th e l o w plant

.

fo rm s such as th e f ungi and th e bact e ria in th e ir physio

, ,

l o gical pr o ce ss e s as fo r e xample th e a ssimilatio n o f nut ritive

, , ,

pro ducts and r e pro ductio n Th e y als o r e s e mbl e ce rtain o f

.

th e pr o t o zo a in th e latt e r p r o c e s s .

4 Th e f o o d supply o f th e bact e ria is c o mp o s e d O f c o mpl e x

.

o rganic mat e rial built up by high e r pla nts a nd animals ,

e xc e pt in th e cas e o f s o m e s o il bact e r ia which a re abl e to

,

ab so rb th e fre e nitro ge n o f th e air and use it fo r nutritive

purp o s e s o r de rive nit ro ge n fro m nitrite s o r nit r ate s This
,
.

o rganic mat e rial must b e i n s o luti o n in o r d e r that it may b e

abso rb e d thro ugh th e ce ll wall I no rganic salts a e fre

. r

qu e ntly pr e s e nt and se e m n e ce ssary fo r pro to plasmic nutri

tio n and to as i st in k e e ping up th e o smo tic pr e ssure o f th e
s

ce ll as is th e case in hi ghe r plants M o st plant s use ve ry

,
.

simpl e sub stance s fo r fo o d .

5
. C e rtain r e c o g niz e d sp e ci e s o f bact e ria hav e th e p o w e r

o f d e riving th e ir nitr o g e n fr o m th e disint e grati o n o f amm o nia

( N H )
a c o mp o u n ds . I n thi s r e sp e ct th e y r e s e mbl e s o m e

plants and diffe r fr o m all ani mals .

 THE RELATI O N S HI P OF B ACT ERI A I I 9

6 . So me bacte ria p o ss e ss th e p o we r o f br e aking

sp e cI e S o f

up carb o n di o xide ( C0 ) into carb o n ( C ) and o xyge n (0) 2

witho ut th e aid o f chlo ro phyll and sunl ight whi ch are use d ,
.

by highe r plants I n s o m e case s th e carb o n and o xyge n are

.

de rive d fro m carb o nate s The y diff e r f ro m all plants and .

animal s in this r e sp e ct .

7 C
. e r tain bact e r ia o f th e s o il tak e up fr e e nitr o g e n o f th e

air and in thi s th e y re se mbl e so me fe w sp e cie s o f fungi and

,

diff e r fro m all animal s .

8 A larg e numb e r o f sp e ci e s o f bact e ria p o sse s s o rgan s o f

.

lo co mo tio n calle d flage lla o r cilia and in thi s r e sp e ct th e y ,

re se mbl e ce rtain animal fo rm s amo ng th e pro to zo a .

9 B
. act e r i al c e ll s p o sse ss n o d e finit e nucl e u s as m o st animal
and plant ce lls d o Th e ce ll pro to plasm is undiffe r e ntiat e d
.

and th e chro matic mate rial distribut e d thr o ugh o ut all th e

parts o f th e ce ll .

1 0 B act e rial c e l ls like all high e r plants d o no t pr o duc e

.
, ,

any ni tro ge n co ntaining e xcre tio ns

-
.

R e l ati o nsh ip to P l ants Ve ge tati o n o n th e e a r th may b e

.

di vide d int o f o ur divi si o ns whi ch a re quit e di s tinct ,

.

I . T hall p h y ta
o A lg ae ,
fun gi ,
e tc .

2 . B ry p hy ta
o M o sse s .

3 . Pt rid p h y t
e o a r
F e ns .

4 . S p m at p h y t
er o a Se e d p lant s .

Th e first thre e di visio ns co mp o se th e C rypto gamia ,

or

no n fl o we ring plants
-
.

Th e micro o rganisms kn o wn as bact e ria b e l o ng to th e

 I 2 0 TAX ON OMY
Thallo phyta Th e Th al l Oph yta co mp o se a m ul titude o f
.

plants whi ch are ve ry primitive and simpl e I n fo rm This .

gro up o f plants are d e vo id o f fi b rO vascular bundl e s ro o ts-

, ,

st e ms and l e ave s Th e ce ll b o dy pr o duce d is call e d a thallus

,
. .

Th e individual c e lls carry o n th e ir m e tab o lic pr o c e ss e s i n

d e p e nde ntly .

Th e Thall o phyta a re subdivid e d acc o rding to s o m e b o tan

,

ists int o th e fo ll o wing gr o up s :

,

I . My x my to ce es li
S m e mo ld s .

2 . S hi z p h y ta
c o B a t ria bl u gr
c e ,
e -
ee n alg a ,
e tc
.

3 . Diat m o ea Diat m o s .

4 . Euph y ce a T r u alge a .

5 . E um y ce t e s Tr u fungi
e .

gre at many auth o ritie s divide th e Thallo phyta into o nly

A
two gr o ups th e fu ngi and th e alg a .

R e l ati o ns hi p to E u m y ce t e s — T h e E umyc e t e s o r fungi

.
, ,

are cl o s e ly r e lat e d to th e bact e r ia in s o m e particular s B ac .

te ria have b e e n call e d S chizo myce te s o r fi ssio n fungi P hysi

,
.

o l o gical l y th e bacte ria e xc e pt c e rtai n o f th e so il fo rms

, ,

agre e with th e fungi in a gr e at many d e tails B o th a re m e ta .

tro phi c and paratro phi c ( Chap X I V) and th e re fo re re q uir e

.
,

o r ganic mat e rial w hi ch h a s b e e n built up by hi gh e r plants

and animals Th e bact e ria an d th e fungi d o no t r e se mbl e

.

e ach o th e r m o rph o l o gically Th e fungi includ e th e m o ld s

.
,

mushro o ms milde ws e tc Th e s e plants lik e th e bact e ria

, ,
.
, ,

a re d e v o id o f chl o r o phyll a nd r e quir e o r gani cfo o d in s o l u

tio n ,

and are no t abl e to build up Carb o hydrate s as high e r plants

 THE RELATI ON S HI P OF B ACTERI A 12 1

do . The y are co mmo nl y fo und to ge th e r with sapro ge nic

bacte ria o n d e caying animal a nd v e ge tabl e mat e rial Th e .

fungi ce lls are granula r no nnucl e at e d and th e pro to plasm

, ,

wate ry and co l o rle s s The y are usual l y fo und co ll cte d to

l

~
. e

ge the r into filame nts call e d hypha Inte rwo ve n hyph a .

a re c o ll e ctively call e d a myc e lium ; Th e fun gi no matt e r ,

what sp e ci e s can b e s e pa r at e d into two di sti nct parts th e

, ,

myce lium o r ve g e tativ part and th e fruiting h e ad o r S p o r e

e ,

b e aring r e pro ductive part N o Such s e paratio n o ccur s in

,
.

th e l o w e r bact e ria I n th e hi gh e r bact e rial f o rm s wh e n re

.
,

pro ductio n take s place all th e ce ll s in th e filam e nt whi ch

,

a re d e s tin e d to r e p r o duc e f o rm go nidi a and th e ve g e tative ,

stat e fo r the se ce lls ce ase s to e xi st Thi s is als o tru e in a

.

m e asur e in re pro ductio n amo ng th e M yxo myc e t e s o r slime ,

mo lds .

Th e fungi a re r e lat e d to th e alga thr o ugh a co mm o n

ance stry The re is a gro up amo ng th e alga o r cl o s e ly

.
,

r e late d to th e m whi ch is call e d th e Phyco m yce ta o r algo

, ,

fungi S o me b o tani sts ho ld th e Vi e w o n se e mingly a go o d

.
,

basis that th e fungi a e m o di fi e d alga which have l o st th e ir

,
r

chl o ro phyll and acquir e d sapro phytic habits .

Th e y e asts o r S acchar o my ce t e s a re c e r tainly v e ry cl o s e ly

re late d to th bacte ria Th e y are unice llula r plants which d o

e .

no t o r di narily pr o duc e filam e nt s and a e d e vo id o f Chl o r o phyll

r .

While having many physio lo gical pr o ce sse s in co mmo n th e y ,

diffe r fro m th e bacte ria in th e m e th o d o f r e pro ductio n th e ,

bacte ria r e pro ducing by fissio n and th e y e asts by budding .

I 2 2 TAX ON OM Y
So me y e asts and mo lds p o sse ss th e p o we r o f pro duc
of th e

ing di se as e co nditio ns in th e human b o dy .

R e l a ti o ns hi p to C yano ph yce a Th e bact e ria ta king .

th e wh o l e gr o up i nt o c o nsid e r ati o n a re witho ut do ubt

m o re cl o se ly r e late d to th e Cya no ph yce a o r blue gr e e n alga ,
-

than to any o the r plants Th e blu e .

g r e e n alga are singl e ce ll e d plants which -

usually f o rm the mse lve s into th r e ads o r

fil am e nts and r e pr o duc e i n th e main by,

fi ssio n similar to bacte ria (F ig ,

.

Th e filam e nt s branch a nd th e r e is a

S p e cificity O f fu ncti o n a m o ng th e vari

o us c e ll s Th e y a re ve ry cl se ly r e lat e d
. o

to th e Chl am yd o b acte riace a a nd th e

Thi o bacte ria S o m e f o rm s al so r e se mbl e .

th e l o w e r bact e ria in shap e Th e y are .

,
FIG 37 Anab a en
as b e f o r e s tat e d filam e nt o u s a nd many
. .

i bili
ra r a C ya s . no , ,

Ph y t Af t r sp e ci e s p o s se ss sh e aths Th e blue gr e e n
f
’ : e -

S hm dt d W i
.

c i an e s .

alga co mp o se th e r e maining p o rtio n o f

th e c o nn e cting link b e tw e e n th e bact e ria and th e chl o r o

phyll b e a ring plants I n addi tio n to th e ir si m ilarity in

-
.

fo r m th e y s e cr e te a g e latino us substance f ro m the ir ce ll

walls similar to so me sp e ci e s o f bact e ria S o me fo rms
. .

(Oscilla ri a ) a re m o til e Th e y di ffe r fro m th e bact e ria

.

and Sho w an advanc e o ve r th e m in th e p o sse ssio n o f a

primitive nucl e u s and chl o r o phyll in th e o ute r lay e r s o f
the ir pr o to plasm Th e bluish gr e e n co l o ri ng matte r is
.
 TH E RELATI O N S HI P OF B AC TERI A 12 3

carri e d in sp e cial ho ll o w chr o mat o ph o re s Thi chl o r o phyll

. s

do e s no t se e m to b e as co mpl e x a s that o f high e r plants .

C e r tain ba ct e ria fo r e xampl e

,
B act e rium vi r id e
, Van
T e igm an co ntain a slight amo unt o f g re e n co l o ring matt e r
,

similar to th e chlo ro phyll o f th e blue g re e n alga Th e y -

.

are o n th e who l e m o r e hi ghly diffe r e ntiat e d than th e

bact e ria B e side s re pro ducing by fi ssio n th e Cya no ph yce a
.

at ce rtain p e ri o d s may r e pro duce by fo rming g o nidia similar

to th e high e r bact e r ia .

I t sh o uld b e r e m e mb e r e d that th e r e is no g e n e tic r e lati o n

shi p b e twe e n th e bact e ria and th e CyanOph yce a Th e y a re .

,

h o w e ve r r e late d th ro ugh a co mmo n anc e s to r

,
.

R e l a ti o ns hi p to E uph yce a . Th e E up h yce a o r tru e alg a , ,

vary in siz e and structu r e fro m simpl e mic ro sco pic unice l ,

l ul ar fo rm s to highl y d e v e l o p e d and c mpl e x b o di e s

o Th e .

Eup h yce a a re no t r e lat e d e xc e pt by a co mm o n s t o ck to th e

C ya nOph yce a o r blu e g r e e n alg a

,
-
C e rtain o f th e uni ce l
.

l ul ar f o rms r e se mbl e th e bact e ria in th e ir m o rph o l o gy but ,

d iff e r fro m the m in that th e y r e pro duce by o Osp o re fo r ma z

tio n in th e p o ss e ssio n o f chl o ro phyll a nd chro mo g e nic pla stids

, ,

a nd al so in th e chara ct e r o f th e c e ll wall Th e similarity o f

.

th e bact e r ia a nd th e Eup h yce a i s o nly sup e rficial Th e .

two gr o up s ca nno t b e sh o w n to b e r lat e d di r e ctly

e .

R e l ti nshi p to M y m yce te s o
a o xo S l im e M o l d s
r Th e s e .

l o w fo rms o f lif e are mo s t f r e qu e ntly fo und up o n d e cayi ng l o g s ,

l e ave s e tc At ce r tain p e rio d s they sugge s t a r e latio nship

,
.

with so me o f th e singl e ce lle d animals and th e fungi S o m e

-
.
12 4 TAX ON OMY
bio l o gists class th e m as anim als S tructur e s call e d sp o rangia
.
,

c o ntaining minut e micro sco pic S p o re s whi ch are pro vide d

with a c e ll wall a nucl e us pro to pla sm a nd capabl e o f ge ne rat
, , ,

ing a ne w b o dy are p re se nt Th e s e b o die s are o nly slightly

,
.

re sistant and a re sp e cific r e pr o ductiv e c e lls The y d o no t .

s e rve to carry th e o rganism o ve r unfavo rabl e p e rio ds as th e

e nd o sp o r e s o f bact e ria d o R e pr o ductio n am o ng th e bac
.

t e ria is a much S impl e r pro ce s s than it is in th e slime mo lds .

Th e M yx o myc e t e s d o no t r e se mbl e th e bact e ria cl o s e ly .

R e l a ti o nshi p to M yxo b a cte ri a ce a Th e myxo bact e ria

a re r o dlik e o rganisms whi ch a re m o til e a nd multiply by fi s

sio n Th e y s e cre te a firm ge latino us base and o ft e n fo rm

.

aggre gat e s o f s e ve ral c e lls Th e fo rmati n o f th e se aggr e ga

. o

tio ns o f ce ll s u sually pr e c e de s a s tag e in which th e r o ds b e

co me e ncy st e d and go into th e r e sting stag e o r b e co m e e n
cy st e d and f o rm S p o r e s Th e ce l l s und e rg o e l o ngatio n and
. .

co nstrictio n in th e ce nt e r be fo re divisio n and n e ve r r e main

attach e d to ge th e r in chains Th e o rga nism mo ve s in a glid
.

ing fashio n no t unlike B e ggiato a Lo co mo to r o rgan s a re .

unkn o wn Th e ma sse s o f th e ce ll s a e r e ddi sh in app e ar

. r

ance M yx o bacte ria are co mm o n in m o i st plac e s as o n

.
,

d e caying wo o d o r o n fungi Th e ir Optimum te mp e r ature is

.

ab o ut 3 C This g ro up o f m i cro O ganism s co nne cts th e

0
°
. r

high e r o r Thi o bact e ria and th e Cya nOph yce a o r blue gr e e n -

alga and th e M yxo myce te s o slim e m o ld s This gro up r .

( M yx o bacte ria) is so m e time s made to co nstitut e a thi rd o rd e r

unde r th e ba ct e ria .
 THE RELATI ON S HI P OF B ACTERI A 12
5

t i zo a acte ria al so p o ss e ss so m e
—B
.

R e l a i o nsh p to P ro to .

p o ints Of r e se mblance to ce rtain singl e c e ll e d animal s b e l o ng -

ing to th e p ro to zo a The r e is a sp e ci e s P o l yto m auv e l l a

.
, ,

b e l o nging to th e o r de r o f F l age ll ata whi ch is o vo id Or b acil ,

lus like in shap e and which p o sse s se s p o lar

-

flage lla Th e numb e r o f the se flage lla is

.

quite de fini te fo r th e sp e cie s The r e are .

o th e r sp e ci e s whi ch p o s s e ss a tuft o f flag e lla

at th e p o l e s similar to th e lo pho tri cho us

and amphit rich o u s bacte ria S till o th e r .

fo rm s p o ss e s s a pul sating vacuo l e which

sugg e sts a r e se mblance to a s t r uctur e o f
that natur e in B e ggiato a Th e ce ll wall in .

ce rtain sp e ci e s o f bact e ria is ve ry similar

to th e c e ll wall o f c e rtai n S p o r o z o a which

b e lo ng to th e pr o to z o a (Fig F .
3 3 _ Dim p h IG . , or a

Th e pro c e s s o f e ncystm e nt which tak e s

r diata S h wi g a . o n
,
l t nuc e us, co n ra c

place wh e n so m e o f th e se f o rms o f p ro til va l t e cuo e . e c .

(P r t z ) Aft r o o oo n e
to o a co m e I n c o ntact w1th u nfavo rabl e
.

S h m idt d W i
z
c an e s .

su rr o undi ngs cl o se ly r e se mbl e s th e fo r ma

,

tio n o f e ndo sp o r e s by so m e sp e ci e s o f bacte ria I n distin ct .

co ntra st to th e bacte ria th e pro to zo a and particula rly th e

, ,

fl age l l a ta p o sse ss a w e ll d e fi ne d nucl e u s

,
-
S o m e o f th e p ro .

to o a are q uit e simila r to th e fu ngi a nd parti cula r ly th e

z

y e ast s a s fo r e xampl e Tricho mo nas inte stinalis and Tri

, , ,

ch o m o nas va gi nali s .

Th e M yx o myc e t e s o r slim e m o ld s b e ar many p o ints o f

, ,
12 6 TAX ON OM Y
r e se mblance to th e pro to zo a and als o an
,

p o ints o f r e se mblance to th e myxo bacte ria .

Th e bact e ria S h o w th e m o st ma rk e d sim ila r iti e s

C yano phyta o r blu e gr e e n alga a nd th e F lag e llata am o ng

,
-

th e pr o t o z o a. Th e y sho uld b e co n sid e r e d as a s e pa r ate

gro up p o sse ssing many p o i nts o f r e se mblance to th e vari o us
,

l o w fo rm s o f plant and animal life .

PA RT v . G E N E RA L P H Y S I O LO G Y

CHAP TER X I I

L
R E ATI O N To E NVIR ONM ENT
I ntro d uc i o n t physi o l o gy is m e ant th e scie nce o f
. By

functio n I n P a r t I V w l e a rne d ab o ut what bacte ria are

. e ,

a nd in P art V we a e to l e a r n what bact e ria d o

r I n co n .

si d e ring th e m rph o l o gy o f bact e r ia o ne o f th e st r iki ng facts

o ,

b ro ught o ut was th e simplicity o f fo rm B act e ria are r e duc e d .

to alm o st th e limit o f simplicity so fa r as f o rm and st r uctur e

a re c o nc e r ne d Ge ne rally sp e aki ng s t r uctu r e and fu ncti o n are

.
,

r e lat e d Th e o rgani sm s whi ch have th e mo s t co mpl e x func

.

tio ns a e th o s e which have th e m o st co mplicate d fo r ms B ut

r .

in th e ba ct e ria th e r e is a s tri ki ng e xc e pti o n to thi s g e ne ral rul e .

Th bact e ria a e v e ry simpl e so fa r a s th e ir fo rm i s co nc e rne d

e r ,

but th e ir functio ns are v e ry co mpl e x and it is o ne o f th e ,

w o nde r s o f bio lo gy h o w such a co mplicate d physio l o gy has

de ve lo p e d o n so simpl e a mo rph o l o gy .

F uncti n s o I n th e s ubj e ct o f phy si o l o gy we hav e to co n

.

sid e r no t o nly th e fu ncti o ns o f bact e ria that i s what th e y d o , , ,

b u t we hav e al so to c o nsid e r th e e ff e ct whi ch th e e nvir o nm e nt

h as up o n th e o rg anism and th e m o dificati o n s w hi ch th e se

e xt e rnal c o n di ti o ns hav e o n th e fun cti o ns .

12 7
12 8 GENERAL P HY S I OL O GY
r
Exte nal C o n di ti o ns Th e e xt e rnal c o nditi o ns may b e
.

with o ut e ffe ct o n th e bact e ria o r th e y may stimulat e activity

, ,

that is th e y may have a to nic e ffe ct o r o n th e o th e r hand the y

, , ,

m ay hav e a r e pr e s sive e ff e ct a nd it is th e pu rp o s e in this co n

,

ne ctio n to c o n sid e r th e m o s t imp o rta nt o f th e e xt e rnal co n

d i tio ns and d e t e rmin e wh e th e r th e i r acti o n is s timulativ e o r

r e pr e ssive and find o ut if p o ssibl e th e co nditio ns which a re b e s t

,

adapt e d fo r th e gro wth o f th e bact e ria o r co nditi o ns whi ch will

l e ad to th e ir de s tructi o n m o s t quickly .

Fo o d Al l fo rms Of lif e mu s t have fo o d and th e bact e ria

.
,

a re no e xc e pti o ns Lik e o th e r fo rm s o f life als o th e y may b e

.
, ,

abl e to ge t al o ng with a ve ry small am o unt that is o n a , ,

maint e nance ratio n o r the y may d o a gre at de al b e tt e r o n a

,

diffe re nt o r large r amo unt that is o n a gro wing r atio n AS

, ,
.

to th e actual am o u nt which is n e e d e d by th e bact e ria it is to

b e n o te d that th e bact e r ia a re di vid e d int o two clas se s One .

class a re accusto m e d to live upo n th e body o f a nimals o r plants ,

and are kno wn as p a asi te s Th e se r e quir e a co n side rabl e amo un t

r .

o f fo o d .Th e o th e r class liv e o n d e ad o r ga nic matt e r and a re ,

kn o wn as th e sap o p hyte s S o m e o f th e s e may sub si s t o n a

r .

surprisingly small amo unt o f o rganic fo o d M e ade B o lto n .

s o me y e ars ago call e d atte ntio n to th e fact that ce rtain wat e r

bacte ria w e r e abl e to live and gro w with gr e at rapidity in
d o ubl e di s tille d wate r wh e r e th e o nly s o urc e o f o rganic matte r
-

wa s th e di r t o n th e sid e s o f th e v e s se l s whi ch h e wa s unabl e to

,

r e m o ve by th e m o st ca r e ful m e th o ds o f cl e ani ng Thi s is o ne .

e xtr e m e M a ny o f th e bact e ria h o w e v e r r e q uir e a c o nsid e r

.
, ,
 RE LAT I ON TO E NVI RONM EN T 12 9

able amo unt o f fo o d sub stanc e On th e o th e r hand th e e .

,
x

c e ss o f fo o d 5 no t witho ut its e ff e ct I t is impo ssibl e fo r

1 .

many o f th e bacte ria to live in f o o ds that a e to o co n ce ntrate d r

‘
.

R e fe r e nc e h a s alr e ady b e e n mad e to th e fact that wh e n th e

bacte rial c e ll is surro unde d by a substa nce that h as gr e at e r

o sm o tic pr e s s u r e tha n th e pr o t o pla m o f th e c e ll th e c e ll is
s

plasmo lyze d B acte ria a re O f c o u r s e u nabl e to g ro w u nde r

.
, ,

the se co nditio ns I f th e n th e ir fo o d sub stance is to o co n

.
, ,

ce ntra te d th e y a e u nabl e to mak e'

,
r u se o f it and thi s fact is ,

take n advantage o f in th e p r e se rvatio n o f a numb e r o f fo o d

sub stanc e s C o nde nse d milk fo r instanc e k e e p s b e cau se
.
, ,

the bact e ria in it a re u nabl e to mak e use o f th e f o o d which is

pr e s e nt the r e in a to o co nce ntrat e d fo rm Th e same thing .

is tru e o f sirups At tim e s ch e mical sub s tanc e s a e add e d

. r

which incr e ase th e o smo tic pr e ssu r e as is true in th e case ,

o f bri ne .

Ch e m i cal C o m po s iti o n o f F o o d F o o d must c o ntain ce r

.

tain che m ical e l e me nt s Tho se whi ch are n e ce ssary are carb o n

.
,

o xyg e n hydro g e n nit r o g e n ph o sph o r u s sulphu r i r o n and cal

, , , , , ,

ci um . N o s ub s ta nc e is valuabl e a s a f o o d which d o e s no t fur

ni sh th e ne c e ss ary ch e mical e l e m e nt I t is t r u e that no t all

s .

substance s furnishi ng th e n e ce ssary e l e me nts a re n e c e ssarily

go o d fo o d sub s tanc e s I t is said that o ne ma n s fo o d is a n
.
’

o th e r ma n s p o i so n S O am o ng bact e ria ; o ne substa nc e may

’
.

se r v e a s a m e dium fo th e d e v e l o pm e nt fo r o ne ki nd o f bac
r

t e r ia and b e ab s lute ly p r o hibitiv e t th e g ro wth o f ano the r

o o .

Wate r .B act e ria r e quir e wat e r fo r gr o wth Th e y may in .

 3 0 GENERA L P HY S I O L O GY
fact b e co nsid e re d as aquatic plants Th e r e aso n fo r thi s is .

p e r fe ctly e vide nt The y se cure th e ir f o o d th r o ugh ab so p

. r

tio n and it is ab so lut e ly n e ce ssary that th e y b e sur ro unde d by

,

m o isture I t is tru e that bacte ria may gro w upo n so lid m e

.

dia but th e y can o nly d o so wh e n th e y a e s u rr unde d by at

,
r o ,

le ast a film o f mo i stu re I f the y b e co me co mpl e te ly drie d

,
.
,

th e y a re u nabl e to fe e d Th e y a re ho we ve r abl e to maintain

.
, ,

th e ir e xi ste nce fo r so m e tim e in a de siccat e d co nditio n but a re ,

no t abl e to gr o w wh e n in thi s s tat e S o far as th e ability o f a

.

m icro O ga nis m to with sta nd d e siccati o n is c o nc e r n e d d iff e r e nt

r ,

kinds o f bact e ria vary gre atly Th e sp o re s a re able to with

.

s ta nd an almo s t unlimit e d am o unt o f d e siccati o n I t is fo r .

thi s r e a so n larg e ly that th e y a e so valuabl e to th e bact e ria

, ,
r .

Th e v e g e tativ e c e ll is no t with o ut thi s p o w e r h o w e v e r , but ,

th e diff e r e nt ki nd s vary gr e atly in th e ir ability to with s tand

co mpl e te d rying F o r e xampl e such a n o rganism as th e

.
,

M sp c o mma o f Asiatic ch o l e ra can with s tand d ryi ng fo r

.

o nly a fe w h o urs o r a fe w days at m o st

,
Th e sam e is tru e .

also o f th e M pn e um o nia and B act influe nza ; but o the r

. .

v e ge tative c e lls such a s th e B act diphthe ria may withstand

,
.
,

th e e ff e ct o f d e siccati o n fo r many day s in s o m e cas e s a s ,

l o ng as 9 4 day s R e f e r e nc e wa s mad e in a pr e c e ding para

1 .

graph to th e e ff e ct o f th e d e n sity o f th e m e dium o n th e bact e ria ,

and it will b e r e call e d that wh e n th e culture m e dium is l e s s

d e nse than th e pro t o pla sm o f th e c e ll th e c e ll is in a s tate o f ,

turge sce nc e that wh e n th e culture m e dium is m o r e de n se than

th e pr o t o plasm th e c e ll is pla sm o lyz e d
,
I t is tru e al so that .
 RELATI ON TO ENV I R O NM ENT I 3 I

th e de n sity o f cultur e m e di um may hav e i ts to nic e ffe ct o n th e

b cte ria C e r tain ba cte ria that a accusto me d to a co n
a . re

sid e a b l e c o nc e nt r ati o n will b e a ff e ct e d by a di mi nut io n in th e

r

de g re e o f co nce ntratio nand so m e f o rm s will mo ve to wards o r

,

away fro m a diff e r e nt co nce nt r atio n f ro m that to which th e y

a re accu s t o m e d .

O xyge n . I t is a g e n e ral l aw Of phy si o l o gy t hat all lif e

mus t br e athe I n 86 P aste ur disco ve r e d a m icro Organism

. 1 1

which gre w b e s t in th e abs e nce o f the fr e e o xyg e n o f th e air ,

and S i nc e that tim e we have co m e to divide bact e ria into two

classe s so far as th e ir r e latio n to th e fr e e o xyge n o f th e air
,

is co nce rne d One class kn o wn as th e a e o be s live s in th e

°

.
,
r ,

fr e e o xyg e n o f th e air and a re co mpa r abl e in th e i r o xyge n re ,

q u i re m e nts to ,
o rdi nary f o ms o f lif e T h e o th e r class kn o wn
r .
,

as th e anae o be s gro ws b e st in th e ab se nce o f fr e e o xyg e n and

r , ,

s o me o f the m gro w o nly wh e n fr e e o xyg e n is r e mo ve d I t is .

no t to b e und e r st o o d h o w e ve r that th e s e ana e r o bic micr o

, ,

o rganisms a re a n e xc e pti o n to th e g e ne ral rul e that all life

mus t bre athe Th e s e f o rms d o bre ath e Th e diff e re nce is th e

. .

s o urce o f the ir o xyge n Th e ana e ro b e s tak e th e ir o xyge n

.

fro m o rganic co mp o unds rath e r than fro m th e fr e e o xyge n o f

th e air . S o m e o f th e bact e ria can gro w u nd e r b o th a éro b ic
and ana e ro bic co nditio ns ; so m e gro w b e st fo r instanc e unde r , ,

a e ro bic co nditio ns but have th e faculty o f gro wi ng unde r

‘

anae ro bic co n di ti o ns S o m e bacte ria gro w b e s t u nde r aua e

.
'

ro bic co nditio ns but have th e faculty o f gr o wing und e r ae ro bic

,

co nditi o ns So that in r e ality we have fo ur clas se s o f bacte ri a

.
13 2 GENE RA L P HY S I O L O GY
h e re . Oneclass kn o wn as th e strict o r o bl iga te ae ro be s can
, ,

gr o w o nly und e r strict a e r o bic co ndi tio ns ; th e o bl igate

a na éro be s can gr o w o nly und e r s trict a na éro b i c c o ndi ti o ns th e

facul tative ana é o b s gr o w b e s t unde r a e r o bic c o nditio n s but

r e ,

have th e faculty o f g ro wing u nde r anae r o bic co nditi o ns ; and

th e fa cul ta ti ve a e o be s g r o w b e st und e r ana e r o bic c o ndi ti o ns but
’ ’

r ,

have th e faculty o f g ro wing unde r a e ro bic co nditi o ns Auae .

r o b e s a e wid e ly di s tribut e d in natur e Th e y are f o und fo r

r .
,

e xampl e i n th e s o il c o nstantly a nd i n gr e at numb e rs

,
F urth e r
.

m o re the y o ccur i th e upp e r laye rs o f th e s o il unde r co ndi

,
h

tio ns wh e re o xyg e n is c o nstantly pr e s e nt ; and it is a matt e r

o f sp e culati o n as to h o w th e c o nditi o ns fav o rabl e fo r th e ir

d e ve lo pme nt a re s e cur e d Up o n o ur artificial cultur e m e dia

.

th e y are o bligate a na éro b e s I t is th o ught by so me that

.

p e rhaps in th e ir natural habitat th e y a re no t so particular in

th e ir o xyg e n re quir e m e nts o r what is q uite as like ly th e y
, ,

may e nte r into partn e rship with ce rtain o th e r fo rms o f life

which furnish th e m anae ro bic co nditio ns B ut whate ve r th e
‘

e xplanati o n it is a w e ll kn o wn fact that th e s e f o rms a re wid e ly

,
-

di s tribut e d and am o ng th e m so m e o f th e di s e a se pr o duci ng -

ba ct e ria as fo r e xampl e B t e tani (l o ckjaw) and B act w e lchii

, , ,
. .

( gas e o us e mphy se ma) .

Ana e o bi c Cul tu e M e tho ds

r r I n o rd e r to cul tivat e th e s e
.

g e rm s o n artificial m e dia it is ne c e ssary to e mpl o y sp e cial

m e th o ds A gr e at many diff e r e nt m e th o d s have b e e n de scrib e d
.

but th e y can all b e gro up e d unde r th e f o llo wing h e ads :

1 . R e plac e m e nt o f air I n thi s cas e air is r e pla ce d by s o m e
.
 R ELATI O N To E NV I R ONME NT 13 3

ine rt gas as hydr o g e n o r carb o n dio xide F o r thi s purp o se

,
.

sp e cial apparatus is n e c e ssary and a g r e at vari e ty o f fo rms

,

o f apparatus has b e e n d e vis e d 2 Abs o rpti o n Of o xyg e n

. . .

Thi s can b e d o n e by m e ans o f c e rtain ch e mical S l ib stance s ,

Such as an alkalin e s o lutio n o f pyr o gallic acid He re to o th e .

, ,

fo rms o f apparatus are num e ro us A v e ry simpl e fo rm o f.

apparatus is mad e by pushi ng th e co tto n plug o f a t e st tub e

cultur e into th e tub e and th e n filling up with an alkalin e so lu
ti o n o f pyro gallic acid and clo sing th e t e st tub e WI th a rubb e r
sto pp e r 3 Exhaustio n o f air This is do n e by m e ans o f an
. . .

air pump Any o f th e ab o ve m e th o ds are e qually go o d and

.
,

in fact may fr e qu e ntly b e c o mbin e d to advantag e

, 4 Ex . .

cl usio n o f o xyg e n Thi s is acco mpl ish e d by p o uring o n to p o f

.

th e cultur e e ith e r m o r e cul tur e m e di a o r ste r il e Oil 5 G r o wth . .

in th e pre se nce o f air Thi s is do n e e ith e r by gro wing with

.

so me o th e r o rgani sm that is ae ro bic o r so m e r e ducing sub

stance such as litmus o r fo rmic acid
,
.

Wh e n P aste ur disco ve r e d the se o rganisms h e tho ught ,

that the y w e re th e o rganisms which carrie d o n fe rme n tatio n ,

and h e de fi ne d f e rm e ntati o n as lif e witho ut air This o n fur .

th e r s tudy has b e e n fo und to b e wi th o ut basis o f fact but it is ,

true that a co nsid erabl e numb e r Of f e rm e ntati o ns are p ro

d uc d by a na e r o b ic bact e ria
e Am o ng th e se may b e m e nti o ne d
.

th e butyric acid f e rm e ntati o n th e f e rm e ntati o n Of c e llul o s e

, ,

th e r e tti ng o f flax and putr e facti o n

,
R e f e r e nc e has alr e ady
.

b e e n made to th e fact that the se anae r o bic bacte ria may p ro

duce di s e ase .
13 4 GENERAL PHY S I OL O GY
Te m pe r atu r e .T e mp e ratur e has a pro fo und infl u e nc e
—

up o n th e l o w e r as w e ll as o n th e high e r f o rms o f life Eve ry .

o r ganism b e c o m e s habituat e d o r attu ne d to c e rtain t e mp e r

atur e s I f th e functi o ns o f life b e co m e adjuste d to thi s par

.

ticul a r t e mp e ratur e any c o nsid e rabl e variati o n a ff e cts th e s e

,

functio n s An incr e ase o f t e mp e ratur e withi n ce rtain li mits

.

incre as e s th e activity o f th e ce ll Th e p o w e r o f abso rptio n

.

is incr e as e d and th e e xcre tio ns are acce l e rate d so that th e wh o l e ,

life pro ce ss b e co m e s m o r e active Variatio ns fro m this te m

.

e ra tur e e ith e r ab o v e o r b e l o w d e cr e a se th e activity B e

p , ,
.

yo nd a c e rtain limit functio nal activity c e as e s and gro wth st o p s ,

and fi nally ce rtain mark e d variatio n s r e nde r th e e xist e nce o f

th e o rganism imp o s sibl e T h e t e mp e r atu r e limits withi n w hi ch
.

th e lif e o f th e bact e ria is p o ssibl e a re wid e in a g e n e ral way ,

e xt e ndi ng fr o m th e t e mp e r atur e o f b o iling wat e r d o wn to

practically ab so lute z e ro T h e t e mp e ratu re limits within

.

whi ch functio nal activiti e s o ccur a e v e ry much narro we r and r

fo r m o s t f r ms o f lif e a re b e tw e e n z e r o and a t e mp e rature

o

slightly ab o ve that o f th e animal b o dy I n co nside ring th e .

e ff e ct o f t e mp e ratur e o n bact e ria th e r e a re s e v e ral cardinal

,

p o ints whi ch must b e tak e n into co n side ratio n Th e re is th e .

m axi mum te mp e ra ture which is th e high e st t e mp e ratur e at

,

which th e bact e ria are abl e to carry o n the ir life pr o ce sse s .

Th e p ti mum te m p e ra tu e is that t e mp e ratur e at which th e

o r

bact e ria gro w b e st Th e mi ni mum te mp e atu e is th e l o w e st

. r r

te mp e ratur e at which th e bact e ria a re capabl e o f gro wing and ,

th e the rmal d e ath p o i nt is that t e mp e ratur e at whi ch bact e ria

 RELATI O N To E N VI R ONMEN T 13 5

a re d e str o ye d Th e th e rmal d e ath p o int will vary de p e nding

.
, ,

amo ng o th e r thing s up o n th tim e th e bacte ria a e sub

,
e r

j e ct e d to thi s t e mp e r atu r e T h e th e rmal d e ath p o i nt is usu

.

ally d e fine d a s th e d e gr e e o i he at r e quir e d to kill th e b a cte ria '

wh e n it is appli e d fo r te n m inut e s in a m o ist co nditio n Th e .

e ff e ct o f a ny gl v e n t e mp e ratur e up o n a bact e rium d e p e nds

up o n its c o nditio n that is wh e th e r it is in a v e ge tative o r

, ,

a late nt stage Wh e n th e bact e rium is in an active ly gro wing

.

c o nditi o n it is much m o re susce ptibl e to e xt e rnal influe nc e s

than wh e n it is in a late nt co nditio n and atte ntio n has alre ady ,

b e e n call e d to th e fact that in th e sp o re stat e th e bact e ria are

unusually r e sistant Ano the r facto r o f a go o d de al o f imp o r tance
.

i s th e kind o f h e at that is e mpl o y e d that is wh e th e r it is m o i s t ,

o r dry . M o ist h e at is much m o r e e ff e ctive than dry h e at at

th e sam e t e mp e ratur e .

Acc o rding as bact e ria a re affe ct e d by h e at th e y a re divid e d

i nt o thre e classe s Th o s e whi ch have a l o w o ptimum te m
.

p e
r a tru e a re k no wn a s t h e p s ych p h il i c b acte i a Th
roe s e r .

gro w at a te mp e r ature o nly a fe w de gre e s ab o ve ze ro Th e .

bact e ria which have a m e dium o ptimum te mp e rature are

kn o wn as th e m e so phil i c bacte i a and are m o re abundant than r

any o th e r class This g ro up may we ll b e divide d into two

.

subgro ups : o ne in whi ch th e o ptimum t e mp e ratur e is ab o ut

that o f a summe r s day ( 5 to 3 C ) and include s pa r ticul arly
’
2
°
0
°
.

th o se bacte ria which live o n de ad o rganic matte r th e sapro ,

p yh t e s ; and th e di s e as e pr o du cing-
bact e r ia o r para sit e s which , ,

have as the ir o ptim um te mp e rature that o f th e b o dy o f th e ir

13 6 GENERAL P HY S I O L O GY
h o st (3 7 to 40 Th e third clas s a re th o s e which hav e a
° °

hi gh o ptimum te mp e rature fr e qu e ntly ranging fr o m 5 0 to

,
°

7
°
0 C and .a re kn o wn as th e the rm o p hi l i c bacte ri a .

Wh e n th e t e mp e rature go e s ab o ve th e maximum o r falls b e

l o w th e minimum th e bact e ria go int o a stat e which r e s e mbl e s
,

that o f d e ath and is sp o ke n o f as rigo r e ith e r h e at rig o r o r co ld

,

rigo r Unl e ss th e state o f rigo r co ntinue s to o l o ng th e bact e ria

.

are capabl e o f r e c o v e ring wh e n br o ught int o n o rmal te m

p e ra ture s Th e r e is
. no d e finit e chang e w hi ch tak e s plac e as
th e r e sul t o f e xp o sur e to e xtr e m e s o f t e mp e ratur e but th e ,

change is cumul ative and may finally b e d e structive Th e .

zo n e o f h e at rigo r is much narro w e r than that o f Co ld rigo r ,

and h e nce th e the rmal d e ath p o i nt m o r e clo se ly fo llo ws th e

maximum than do e s th e co ld d e ath p o int fo llo w th e minimum .

I t is a p o pular n o ti o n that c o ld is fr e qu e ntly and rapidly d e

structive to bacte rial life but thi s id e a is no t b o rn e o ut by e

,
x

p e rim e nt fo r it is f o und that wh e n bact e ria a re s ubj e ct e d to

,

co ld e ith e r that o f o rdi nary ice o r much l o w e r te mp e ratur e s

, ,

th e y ca n p e rsist fo r v e ry l o ng p e ri o ds o f time A numb e r O .f

y e ars ago P rudd e n fro ze B p ro di gio sus and fo und that th e y

.

w e re living at th e e nd o f thirty s e v e n days in much r e duc e d

-

numb e rs but o nly o n th e fi fty fi rst day had th e y e ntire ly d is

,
-

app e are d S edgwick like wise a numb e r o f y e ars ago f o und

.
, , ,

that th e B typh o sus wh e n fro z e n in ice p e rsi st e d in co nside r

.

abl e numb e rs e ve n fo r a p e rio d o f 03 day s F r o m this it is 1 .

cl e ar that o rdi nary co ld canno t b e de p e nde d upo n to kill

bacte ria e ve n tho se o f th e mo st de licat e so rt Wh e n bacte ria
,
.
 RELATI O N TO ENV I RON M ENT I
37

a re subj e cte d to e xtre m e s o f co ld such as may no w r e adily

,

b e o btai ne d by th e u se o f liquid air it has b e e n f o u nd by ,

R ave n e l M cF ayd e n and o th e rs that th e y may liv e fo r l o ng

, ,

p e rio ds o f tim e and inde e d may b e littl e change d I t is

, , .

to b e und e rst o o d in all O f th e s e cas e s that s o m e i ndividuals

r e adily succumb but that c e rtain o th e rs p e r sist and this is ,

f o und to b e th e case wh e n e ve r bacte ria are br o ught into un

favo rabl e surro undings Th e r e is gr e at diff e r e nce in th e
.

p o we r s o f re si stance t o wards any finfav o rabl e co nditio ns by

th e di ff e r e nt individuals in th e sam e cultur e .

Co mpar e d with o th e r fo rms o f life bact e ria a re m o re r e sist

ant as will b e s e e n in th e fo llo wing tabl e
,

Co mparati ve P owe r o f Vario us Orga nis ms to re s ist Cha nge s o f Te mpe rature

COLD D E REE
G S C .

B a t ria v g tativ
c e ,
e e e

p r s o e 350

4 to

Th e te mp e ratur e o f th e bacte rial ce ll vari e s with th e te m

p e r a ture o f th e s urr o und i ng s in th e sam e way that th e t e mp e r

atur e o f th e co ld bl o o d e d animal vari e s Th e bact e ria are th e re

-
.

f o r e sp o k e n o f a s po i ko l o the rmi c in co ntradi stinctio n to tho s e

'

o rgani s ms which hav e a c o nsta nt t e mp e ratur e as fo r instanc e , , ,

13 8 GENERAL PHY S I OL O GY
th e warm bl o o de d animals whi ch are kn o wn as ho mo the rmic
-

, .

F urth e rm o r e it is f o und that so m e Of th e bact e ria a re abl e to

,

gro w in a wide te mp e ratur e range That is th e ir minimum

.
,

t e mp e rature is wide ly s e parat e d fro m th e ir maximum te m

p e ra tu re
,
a nd th e ir O ptimum i s usually a wid e z o n e Th e se .

o rga nisms a re sp o k e n o f a s e u ythe mi c r M o s t o f th e sapr o

r .

p yh te s b e l o ng to this class and a re thus di stinguish e d fr o m th e

cla ss o f bacte ria which are d e scrib e d as ste n the mi c whi ch haveo r ,

a narro w t e mp e rature range S uch bacte ria are th e patho

.

g e nic o r dise ase p ro d ucing o n e s M any o f th e se o n acco unt o f

~
.
,

th e ir parasitic m o de o f life h ave b e co m e adapte d to a partic

ular t e mp e rature and o nly Slight variati o ns fro m this inte rfe r e
,

with th e ir life pro c e ss e s in a ve ry striking way S uch a g e rm is .

th e B act tub e rcul o si s which has b e c o m e adapt e d to a parasitic

.
,

m o de o f life and is d e p e nde nt up o n a v e ry narro w te mp e ra

ture rang e Th e p sychro phi lic bact e ria w e re f o und by F o ste r
.

gro wing in m e lting I ce F isch e r fo und th e m in se a wat e r

. .

M any o th e r s hav e f o und that th e y e xist in s o il and m ilk and

vari o us o th e r mate rial s Th e y gr o w quite rapidly e ve n at a
.

t e mp e ratur e o nly slightly ab o ve ze r o The s e p sych ro L .

philic bacte ria are similar to th e re d sn o w plants fo und in

th e n o rth and th e di at o ms that swarm th e Arctic S e as so far
,

as th e ir r e sistance to c o ld is co nce rn e d .

The rm al De a th P o i nt Th e d e t e rminati o n o f th e th e rmal

.

de ath p o int is a matte r o f so much practical imp o rtance that

it is w o rth whil e to c o nsid e r it at s o me l e ngth h e r e I t is .

i mp o rtant b e caus e it is d e sirabl e to apply as l o w a t e mp e ratur e

 RELATI ON TO ENV I RONMEN T 1
39

as p o ssibl e to vari o us fo o d substance s so as no t to i njur e th e m ,

a nd at th e sam e tim e th e t e mp e r atur e must b e s uffici e ntly

high to de st r o y all ge rm s capabl e o f p ro ducing di se as e Th e .

t e mp e ratur e n e c e ssary to kil l bact e ria in a g e n e ral weiy is no t

far fr o m 60 C wh e n this is appli e d fo te n minut e s in a mo i st
°
. r

c o nditi o n Wh e n sp o re s are pre s e nt a t e mp e r ature o f 9 to

.
,
0
°

C is n e c e s sary and so m e tim e s thi s mu s t b e appli e d by

°
100 .
,
.

th e int e rmitt e nt m e th o d in o rd e r to pr o duc e c o mpl e t e st e r ility .

Th e th e rmal d e ath p o int o f di ff e r e nt o rganisms vary d e p e ndi ng ,

up o n inh e r e nt charact e ristics as s e e n in th e f o ll o wing tabl e

,

M icrcsp ira co mm a
Str p t
e u
o co cc s n py g o e es

B a ill t yp h u
c us os s

B a t ri um di p hth ri
c e e a

Mi n ( p i unk n
cro co cc s s eces o wn)

Th e the r mal de ath p o int vari e s with the r e actio n o f th e

me dium up o n whi ch th e ge rm is g ro wn and p e rhaps al so with
that o f th e m e dium in which th e t e st is made Th e sam e va r i .

ati o n amo ng individual s that was n o t e d ab o ve in co n sid e ring

th e e ff e ct o f c o ld up o n bact e ria is n o t e d h e r e I f a quantita .

tive de t e rminatio n is made o f th e numb e r o f bact e ria kill e d at

vario us t e mp e ratur e s just b e l o w th e th e r mal d e ath p o int it ,

will b e f o u nd that within a fe w de g r e e s th e g r e at maj o rity

o f th e i ndi vidual s i n a singl e cultur e a re kill e d and thi s we ,

may S p e ak o f a s th e no rmal the rm al de ath p o i nt C e rtain in .

di v id ual s p oss e ss a high p o w e r O f r e si s tanc e and r e quir e a

high e r d e gre e o f h e at to kill the m Th e n e ce ssary h e at re .

140 G ENERAL P HYS I OL OGY
quire d to kil l th e se is sp o ke n o f as th e a bso l ute the mal d e ath r

p o i nt The r e have b e e n two m e th o d s wide ly u se d fo r th e

.

de te rmi natio n o f th e th e rmal de ath p o int : o ne de vi se d by

S t e rnb e rg wh o us e s a gla ss bulb and th e o th e r d e s c r ib e d by th e
, ,

C o mmitt e e o n S tandard P r o c e du r e s app o int e d by th e Am e ri

can P ublic He alth Ass o ciatio n I n b o th o f th e s e m e tho ds.

o nly th e ab so lut e th e rmal d e ath p o int i s d e t e rmi ne d a nd e ach ,

m e th o d has ce rtain advantag e s and c e rtain di sadvantage s .

A capilla r y b e nt tub e m e tho d whi ch wa d e vi e d in th e lab o r a

,
s s

to ry o f o ne o f us o ff e rs c e rtai n advantag e s o ve r b o th o f th e
,

o th e r m e th o d s De ath f r o m h e at i s u nd o ubt e dl y d ue to th e
.

co agulatio n o f th e pro t o pla sm and th e di ff e re nt p ro to pla sms

,

have di ff e re nt p o ints o f co agulati o n One diff e r e nc e whi ch may .

acco unt fo r th e se variati o ns is th e diff e r e nt amo unt f m o is o

tur e pre s e nt Thi s wo ul d te nd to e xplain e p e cially th e

.
,
s ,

high th e rmal de ath p o int o f sp o r e s co mpare d with th e v e ge

ta tiv e c e ll .

Egg alb um i n 5 0 pe r t wat r

ce n e co agulat at 5 6 C es
°
.

Egg alb m i n nt wat r agulat at 7 4 8

— ° °
u 25 p e r ce e co es 0 C .

Egg alb m i n n t wat r ag lat at 8 —9 ° °

u 1 8 p e r ce e co u es 0 0 C .

Egg alb um i n n t wat r ag lat at 4 5 C °

6p e r ce e co u es 1 .

Egg alb m i n d agulat at 6 ° °

u r y co es I 0
-
170 C .

Th e Acti o n h e mi cal s o n B acte ri

of C R e f e r e nc e has a
— .

al r e ady b e e n made to th ch e mical r e actio n o f th e m e dia

e ,

and th e i nflu e nc e whi ch it h as up o n th e de ve lo pm e nt o f th e

bact e ria I t was n o t e d that th e bact e ria p r e f e rr e d a n e utral
.

o r S lightly alkalin e m e dium rath e r tha n an acid o ne alth o ugh ,

th e y will to l e r at e o ne that is slightly acid .

 RELATI O N To E NVI RONM EN T 14 1

C h e mical s influe nce bacte ria in much th e sam e way that

o th e r e xt e rnal ag e nt s d o in that unde r ce rtain co n di tio ns
ch e m ical substance s are to nic i e th e y have a stimulating
,
. .

e ff e ct . O th e r ch e mical s o r di ff e r e nt co nc e ntrati o ns o f th e
,

same che mical may pro duce a r e pr e ssive actio n l e adi ng

, ,

ultimat e ly if th e actio n is p e rsiste d in to th e de ath o f th e

, ,

bacte ria .

One o f th e m o st st iking e ff e cts o f ch e micals o n bact e r ia is

r

th e p o w e r th e y hav e to induc e a nd co nt o l l o c o m o ti o n r T he r .

fact that bacte ria se e m to b e att r act e d to ce r tain substance s is

a matt e r o f l o ng s tandi ng o b se rvatio n ; fo r in stance in s tag ,

nant wate r o r in put rid s o lutio n s it is o ft e n se e n that th e bac

te ria and o th e r Cil i at e d o rganisms co ll e ct in mas se s aro und
, ,

c e rtain pa r ti cl e s o f th e d e caying o r putr e fying sub stance s .

With hi gh e r animal s thi s wo uld b e calle d in stinct This .

pro p e r ty is wide spr e ad thro ugho ut b o th th e plant and

animal ki ngdo m and is ve ry fr e qu e ntly n o te d amo ng th e
,

bact e ria Thi s ph e no m e n o n is kn o wn as che mo taxis o r

.

che mo tr o pism .

Th e ph e nom e no n may b e n o t e d u nd e r di ffe r e nt c o nditi o ns

fo i ns ta nc e if th e ch e mi cal s ub sta nc e h a s a nut r itiv e value

r , ,

it is so m e tim e s sp o k e n o f as t p ho t o p i m ( attractio n to wards

ro r s

fo o d) if th e ch e mical sub st nce is a g s it is so m e time s calle d

a a ,

a é t pi s m
ro ro Th e simpl e s t ma nif e stati o n o f thi s ph e no me no n
.

i s th e m o v e m e nt o f bact e ria t wa r d s f r e e o xyg e n

o I t h as al .

r e ady b e e n n t e d that ba ct e ria in th e hanging dro p p re pa r a

o -

tio n f r e qu e ntly co ll e ct a t th e o ute r e dge f th e d ro p Thi s is o .

14 2 GE NERAL P HY S I O L O GY
d ue to th e influe nce o f th e o xyge n o f th e air
ch e m o ta xi c .

That it is th e o xyge n that attracts can b e sho wn in ano th e r

way ; wh e r e an algal ce ll is in th e sam e so lutio n with bact e ria ,

and is active ly e vo lving o xyge n a s it manufacture s its fo o d ,

th e bact e ria in th e ne ighb o rh o o d are attract e d to it Eu .

a B C

FI G 3 9
. p tr m f th hr m phyll f ba t ri —p rp ri h wi ng b
.
— a, s ec u o e c o o o c e o u u n, s o a

so rpti ba d at on d n s An (i vi ibl ) ab rpti

A ba d a an A 1
1 . n s e so on n

h b d t rm in d by m n f th b l m t r at A
as een e e ne b th ba t ia ea s o e o o e e .
, e c er

a re a ggr g at d hi fly in th r gi
se e n f th
e ab pti n ba d Th
e c e e e o ns o e so r o n s . e

a m lati f ba t ia i th r g i f ab rb d rg y m d t th
ccu u on o c er n e se e o ns o so e e ne se e s ue o e

fa t that th m v ing ba t ria a

c et pa fr m a r gi f high rgy t
o c e c nno ss o e on o e ne o o ne

o f l w with t a vi l t ti m l
o whi h i m p l th m ba k ag ain ( Fr m
ou o en s u us c e s e c . o

Eng lm ann 83 ) e ,
’
a
.

ge h na nn has made use o f this pr o p e rty to S ho w th e infl ue nce

o f th e di ff e r e nt c o l o r s o f th e sp e ctrum o n a filam e nt o f algal

c e lls Th e bact e ria c o ll e ct in th e zo ne s o f gre at e s t ab so rp

.

tio n o in o th e r wo r ds in that pa r t o f th e filame nt wh e r e

,
r, ,

o xyg e n i s m o s t rapidly e v o lvi ng ( F ig .

N um e r o us i no rganic salts a re capabl e o f pro ducing a ch e m

o ta xi c i nflu e nc e o n bact e ria S o m e ch e mical sub s ta nc e s p o . r ;

duce an att r active e ffe ct S uch substanc e s are sp o k e n f as . o

 RELATI O N TO ENV I R O NM EN T 14
3
’

b e ing p o i ti l y che m taxi c; o th e r ch e mical sub s tance s r e p e l

s ve o

th e ba ct ria and a e sp o ke n o f a n ga ti v l y he m o t ti c
e r s e e c ac .

P f e i ff e r f o u nd that th e s alt s o f th e alkali e s e x e rt a v e ry p o si

tive e ff e ct ; th t th e alk li ne e a r th s e xe rt a le ss a cl ive in

a a

fl u nce
e . I no r ga nic a cid s a re g e ne r ally r e p e ll nt e xc e pt e ,

ph o sph o ric acid and th e ph o sphat e s o f th mu scl e O

,
e . r

a n i c co mp u nds a e f roe q u e ntly att rra ctiv e T hi s might

g .

r e adily b e e xp e cte d b e cau e it h as b e e n so f r e q ue ntly Ob se rve d

,
s

that d co mp o sing o rg ni c matt e r is att r activ e to bacte ria

e a ,

a nd thi s might in all lik e lih o o d b e att r ibut e d to o rga ni c sub

, ,

st nce s M any O f th e o rganic a cids are att r active S uch

a . .

nit ro ge n co mp o und s a s u re a sp a gin p e pto ne a nd m e at e a, r , ,

x

t r a ct a e v e ry att r active Gly ce rine is ne ut r al o r witho ut

r .

e ff e ct whil e al co h o l pr o duc e s r e p e ll e nt acti o n

,
.

Th e di ff e r e nt co nc e nt r ati o ns o f so luti o ns pr o du ce di ff e r e nt

e ff e cts ; p o ta ssium chl o rid e attra cts in dilute so lutio ns but

r e p e l s in s t ro ng r so lutio ns I t is imp o s sibl e to p r e di ct what
e .

e ff e ct a ny ch e mical s ubsta nce may hav e o n th e b ct e r ia o n th e a

basi s o f its ch e mical c o nstitutio n Th e actio n is appa r .

e ntly no t d e t e r mi ne d by th e e l e m e nt s whi ch e nt e r i nt o th e

c mp o u nd but by th e e nti r e co mp o u nd ; fo
o ,
instance mil k r ,

s uga r p r o duc e s a p o sitiv e ch e m o ta i c e ff e ct in o ne p e r c e nt x

s luti o ns whil e g r ap e sugar r e quir e s a t n p e

o ,
c e nt s o lutio n e r .

I t i s c o nce ivabl that thi s ph e n m e no n m y b e o f use to th e

e o a

bact e ria fo instanc e in h e lping th m to se l e ct th e ir fo o d ; but

,
r ,
e

it pp e a r s that fo o d sub tance s a e no t alway s p o sitive ly

a s r

ch e m t o i c a nd th e bact e ria have no p o w e r appar e ntly o f

ax , , ,
144 GENERAL P HY S I OL O GY
di ff e re ntiating b e tw e e n a substance whi ch is a g o o d fo o d and
a p o iso n AS an e xampl e F i sch e r has cite d th e fact that a
.
,

p e pto ne s o lutio n to which a littl e co rr o sive s ublimate h as b e e n

adde d e xe rts a p o sitive ch e m o ta ic influ e nc e and th e bacte ria
x ,

rush into th e s o lutio n o nly to b e de stro y e d by th e pr e s e nce o f

th e sublimat e I t has b e e n th o ught that thi s attracti o n in
.

dica te d o n th e part o f th e bact e ria a p o w e r o f se l e cti o n and

, , ,

that the y w e r e attract e d and purp o se ly m o ve d to wa r ds th e se

substance s F r o m th e wo r k o f o th e rs h o we ve r it s e e ms
.
, ,

lik e ly that thi s is no t th e ca se but that th e bacte ria swim

,

ab o ut in an aiml e ss fa shi o n and wh e n the y co m e in co ntact

,

with th e s e ch e m o taxic substance s a re und e r th e ir influe nce .

Th e y find co nditi o ns th e re m o re favo rabl e and h e nce re ,

main in th e z o n e ; but c e rtain bacte ria in th e so lutio n may

fail to ge t into this zo n e and th e re fo r e do no t co m e und e r its
,

influe nce .

Th e ph e n o m e n o n o f ch e m o tro pism is o f g e n e ral int e r e st

th ro ugho ut bio lo gy and e nabl e s us to e xplain m o ve m e nt s o f

,

many kinds o f life that wo uld o th e rwise b e ve ry diffi cul t to

unde rstand As e xampl e s o f thi s we may cit e th e attractive
.

influe n ce whi ch th e f e mal e c e lls have fo r th e mal e c e lls amo ng

plants and amo ng animals Ove rto n n o t e d that th e bacte ria
.

co lle ct aro und co njugating S pir o gyra wh e r e th e tub e s ari se

,
.

Th e p o ll e n tub e m o v e s d o wn th e stam e n o f fl o w e rs in all pr o b

,

ability unde r th e influe nc e o f thi s phe n o m e n o n Th e spe r .

m ato o a o f plant and animal c e ll s are attractiv e to th e e gg c e lls

z

b e caus e o f th e e xcr e tio n by th e s e e gg c e lls o f c e rtain sub

, ,
 RELATI ON TO E NV I RONM EN T 14 5

stance s which have a ch e m ta ic infl ue nce o n th e sp e rmato zo a ;

o x

and this wo uld e xplain th e phe n o m e no n o f f e rtilizatio n whi ch

w o uld b e difli cul t o n any o the r basis .

Th e whit e bl o o d co rpu s cl e s a e susc e ptibl e to che m o ta ic

r x

influ e nce s and c e rtain m e tamo rpho se s are d ue to thi s Th e

,
.

change fro m th e tadp o l e stage to th e f r o g stag e is bro ught

ab o ut b e cau se o f th e ch e m o taxi c influe nce whi ch ce rtain tissu e s ,

unde rgo ing d e ge ne ratio n have o n th e whi t e bl o o d co rpu scl e s

,
.

T h e sam e e xp l a na ti h is us e d fo r ch ange s whi ch tak e plac e

o
'

in o l d age th e de ge ne r atio n o f ce rtain ti ssue s th e l o ss o f

, ,

pigm e nt in hair e tc Th e ph e no m e no n o f i nflammatio n is

,
.

e xplain e d o n thi s ba sis Th e invadi ng m icro Organism e

. x

e rts a ch e m o ta xi c influ e nc e o n th e whit e bl o o d c e lls which ,

cause th e m to l e ave th e bl o o d ve ss e ls and to co ll e ct aro und

th e invad e r s and thi s ph e no m e no n is r e ally th e basi s o f M e t e h
,

ni ko ff s th e o ry o f i m munity which is so wid e ly acc e pt e d at

’
,

th e pr e s e nt tim e .

Th e ph e no m e n o n o f ch e m o taxis can b e stu di e d und e r th e

mi cr o s co p e by a m e th o d wh ich was d e vis e d by P f e iff e r and ,

whi ch is carrie d o ut as fo ll o ws
S e l e ct e d capillary glass tub e s are plac e d in watch gla ss e s

co ntaining th e so lutio n to b e t e ste d ; the y are th e n place d

unde r a vacuum pump and th e air is d r awn o ut so as to co m
p l e te l y
,
fi l l th e tub e s with th e s o luti o n T h e tub e s a re th e n
.

tho ro ughly rinse d and th e o p e n e nd o f th e fill e d tub is

,
e

plung e d into a hanging d ro p o f th e bacte ria to b e studi e d .

Th e s o luti o n gradually diff us e s o ut a nd th e ba ct e ria which

, ,
14 6 GENERAL P HY S I O L O GY
at fir st w e r e e ve nly di stribut e d thro ugho ut th e hanging dro p ,

make th e ir way into th e tub e and co ll e ct in it .

C h e mical s ub sta nc e s m ay hav e a n e ntir e ly di ffe r e nt acti o n

fro m that di scu ss e d ab o ve ; nam e ly a ge rmi cidal actio n Thi s

, .

d e structive actio n is fr e qu e ntly sp o ke n o f as an antis e pti c

o r di si nf e cti ng acti o n . B y a n a nti s e ptic is m e ant a sub

stance which i nte rfe r e s with o r p r e ve nt th e gr o wth o f bacte ria

s

whi l e di sinfe ctant are tho se substan ce s which de stro y tho se

s

bact e ria capabl e o f p ro ducing infe ctio n Al l disinfe cta nts ve ry

.
,

natu r ally p o ss e ss antise ptic pr o p e rti e s but it do e s no t ne ce s

, ,

saril y f o ll o w that all anti se ptic s p o sse s s di sinf e cting qualiti e s .

A di si nf e ctant may co mpl e t e ly d e s t r o y bact ria a nd th e p ro

e ,

c e ss may b e co mpl e te in th e s e nse o f ste rilizatio n o r it may ,

b e i nc o mpl e t e th e ch e mical b e i ng abl e to d e str o y o nly c e rtain

,

c e ll s A c o nside rabl e numb e r o f th e di sinf e ctants fo r e xampl e

.
, ,
\
a re r e adily fatal to th e v e g e tativ e c e ll s but a e unab l e to kill
,
r
~

th e sp o r e s . Th e di si nfe cti ng acti o n o f a sub stanc e i s influ e nc e d

by many facto rs S o m e o f th e se facto r s a re r e side nt in th e

.

c e ll s the m se lv e s and o th e r s e xist in th e e xte rnal e nvi ro nm e nt .

Th e pr o t o pla s mic c o nt e nt s o f va r i o u s bact e ria di ff e r to a

c o n side r abl e e xt e nt in th e i r r e si sta nce to ch e mical s ubstance s .

Th e v e g e tativ e c e ll s o f s o m e sp e ci e s a e tw e nty tim e s a s r e si s t

r

a nt as a e o th e r s wh e n act e d up o n by th e s am e di sinf e cta nt

r

and unde r th e sam e co nditio ns Th e n to o th e individual s in

.
, ,

th e sam e cul tu r e di ff e r in th e ir p o w e r s o f r e si s ta nc e so m e i ndi

,

v i d ual s b e i ng much m o r e r e sista nt tha n o th e r s .

C e rtain o f th e in o rganic ch e micals a re v e ry p o w e rful dis

 RELAT I O N TO ENV I RO NMEN T 47

infe ctants M any o f th e ch e mical e l e m e nts such a s chl o rin e

.
, ,

br o mine i o din e and o xyge n a e go o d di sinfe ctants Th e

, , ,
r .

h e avy me tal s such a s go ld silve r a nd m e r cury have quit e

’

, , , ,

pr o no unc e d disinf e cting value Oth e r s su ch a s ni ck e l zinc .

, , ,

and COpp e r have o nly a slight val ue ; whi l e o th e r s as l e ad and

, ,

tin have no ne Th e acid s and alk li e s d o no t have a marke d

,
. a

ge rmicidal value e xc e pt wh e n th e y are us e d in st ro ng so lutio ns ,

a nd th e n th e y hav e a d e s tru ctiv e acti o n o n m e tal s and fab r i cs .

M e cu i c chl id e o r co r ro sive s ub limat e i s o ne o f th e m o st

r r or , ,

e ff e ctiv e o f th e in o r ga ni c salt s Thi s h as an antis e ptic a ctio n .

in a o ne millio nth s lutio n and a di sinfe ctant acti o n in wate r

-
o ,

in a o ne to fi e hu ndr e d th o usa ndth s o l u

v tio n This ch e mical
-
.

uni t e s with pro t ein matt e r a nd its use a s a p r acti cal di sinf e ct
,

a nt i s limit e d O u that a cc o u nt C e rtain o f th e s il ve s a l ts . r

a re u se d pa r ticula rly in m e di cine a s anti se ptics Li m e is

, ,
.

o f g r e at valu e as a di si nf e cta nt but wh e n it i s air s lack e d ,

p o sse sse s no value A f r e shl y pr e par e d lim e wate r s o lutio n o r

.
,

a white wa sh make s an e xce ll e nt disinf e ctant and is e sp e cially

, ,

valuabl e in disinfe cting a s tabl e o r ba rn b e cause it h as ce rtain ,

p ro p e rti e s in additio n to its ge r micidal o n e s that are m o st

b e ne ficial such as ab so rptio n o f o do r s r e fle ctio n o f light e tc
, , ,
.

Chl id e of l i m is als o a n e xc e ll e nt di si nfe cta nt

or e I t h as re .

ce ntl y b e e n S h o w n to b e v e ry e ff e ctiv e as a di si nfe cta nt o f

se wag e e ffl u e nt s As a ga se o u s di i nf e ctant s ul p hu h as had

. s r

a ve ry pro mine nt plac and it is p p ula rly b e lie ve d that th ee ,

o

bu rning f a littl e sulphu r in a r o o m p ro duce s a go o d d isin

o

fe ctant . S ul phur h as a g o o d d e al Of value but no t n e arly as ,

much as is o rdinarily b e lie ve d I n o rde r to disinfe ct a r o o m
.

having a th o u sand cubic f e e t o f air space ( 10 by 10 by 10

it is n e ce ssary that f ro m th re e to fi v e p o u nds o f sulphur b e
burn e d in it F urth e rmo r e since bu r ning o r o xidi zing o f
.
,

sulphur f o rms sulphur dio xide and sulphu r dio xid e is no t a ,

g o o d di sinfe ctant it i s no t e no ugh m e re ly to bu rn sulphur

,
.

Th e re must b e m o i s tu r e p r e se nt M o i stur e can b e Obtain d . e

by m e an s o f ste am Th e activ e di sinfe cting age nt f o rm e d by

.

th e uni o n o f th e sulphu r di o xid e and th e wat e r i s sulphur o us

acid S ulphu ro us acid is a go o d di sinf e ctant but it is also an

.

ide al bl e aching ag e nt and whe n it is us e d all o f th e d rap e rie s

,

in a r o o m are de st ro y e d Unl e ss thi s o ccur th e di sinf cting

. s, e

pro c e s s has no t b e e n pr o p e rly carri e d o ut a nd if it is do ne o f , ,

co ur se th e drap e ri e s wall pap e r o r what e ve r it may b e a re

, , ,

r e nd e re d us e l e s s A go o d de al o f mi s und e rs tanding the n

.
, ,

h as pr e vail e d in th e u se o f sulphur a nd it is no t p o pular at ,

th e pr e se nt tim e fo r family use Th e re a re co nditi o ns h o w

.
,

e v e r u nd e r whi ch it w o r ks w e ll and fo r c e rtai n pu rp o s e s it has

, ,

no c o mp e tit o r . Thi s is tru e wh e r e it is de si r abl e to kill high e r

f o rms Of life su ch as ins e ct s a nd r o de nts and it h as ve ry
, ,

gre at valu e fo ce rtain line s o f wo rk su ch a s in th e di sinfe ctio n

r ,

o f th e h o lds o f i nf e ct e d S hip s wh e r e it i s d e sir e d to ki ll rats ,

mice and i nse cts

,
.

F o rm al de hyd e is a ch e mical sub s tanc e whi ch has c o m e into

use as a di si nf e ctant within r e c e nt y e ars a nd has b e c o m e

d e s e rve dly p pular I t is a gas fo r m e d by th e i nco mpl e te

o .

o xidati o n o f m e thyl al co h o l Alth o ugh a gas it is put up o n

.
,
 RELATI O N TO ENVI R ONM ENT 14 9

th e marke t as a so lutio n I n its o rdi nary fo rm th e wate ry

.

s o lutio n co ntai ns 4 p e r c e nt o f th e gas F o r purp o se s o f dis

0 .

infe ctio n fo rmalde hyd e may b e u se d e ith e r as a gas o r a so lutio n .

I ts u se s a s a gas will b e c o nsid e r e d fir s t .

Th e gas may b e pr o du ce d in th e spac e to b e di sinfe ct e d

dir e ctly f ro m th e m e thyl al co ho l and fo r thi s purp o se a numb e r

,

o f sp e cial lamps hav e b e e n ma nufactu r e d Th e ch e ap e r o ne s

.

a re un s ati s fact o ry in that th e y a e lik e ly to giv e an un e v e n

r

p ro duct ; th e m o r e e xp e nsive o ne s m ay b e quit e sati sfacto ry ,

but b o th have th e di sadvantage s Of l e aving a fi re burning in a

cl o se d r o o m which is liabl e to l e ad to accide nts so that th e
, ,

gas is alm o s t unif o rmly pr o duc e d at th e pr e se nt tim e by

r e ge ne r ati ng it f ro m th e so luti n I t might naturally b e
o .

e xp e ct e d that thi s w o uld b e a simpl e matt e r and that all ,

that w o uld b e n e ce ssary to d o w o uld b e to e vap o rate th e

so lutio n but wh e n fo rmalde hyde so lutio n e vap o rat e s unde r
,

o r di nary c o ndi ti o ns it is lik e ly to p o lym e riz e ; that is two o r ,

mo re m o l ecul e s unite to fo rm a ve ry diffe re nt substanc e .

S o m e o f th e p o lym e r s in this cas e a re s o lid so that if an

‘

, , ,

o rdi nary pan o f f o r mald e hyd e i s e vap o rat e d a c o nsid e rabl e

amo unt o f a white s lid is f r me d kno wn as pa r afo rmalde hyde

o o ,
.

I n thi s ca se th e f o r mald e hyd e in s t e ad o f b e i ng giv e n Off a s a

,

gas go e s into thi s p o lyme riz e d fo rm and is l o st as a di sinfe ctant

,
.

Th e dang e r o f p o lym e rizati o n is o v e rc o m e in s e v e ral way s ;

o ne o f th e m e ans that h as b e e n m o st w i d e ly us e d is th e rapid

h e ating o f th e so lutio n ab o ve th e b o iling p o int in an auto clave

, , ,

o r in sp e cial r e t o rts
. I t has als o b e e n f o und that if glyce rine
 5 0 GENERAL P HY S I O L O GY
is add e d in a small amo unt name ly 10 p e r c e nt that p o ly , ,

me i a tio n is pr e ve nt e d
r z .

The re a e a numb e r o f pi e c e s o f appa r atus o n th e mark e t

r ,

s o ld at a ve ry r e a so nabl p ri c whi ch d o e x ce ll e nt se rvic e as

e e ,

r e ge n e r ato rs Am o ng th e l o we r p ri ce d o ne s th o se havi ng a
.
-

Co pp e r r e t o r t h e at e d by a P r im u l mp a e p r bably th e m o s t
,
s a ,
r o

satisfacto ry Th e y a e so a rr ange d that th y ca n b e r un o ut

. r e

side o f th e spac e to b e di si nf e ct e d Th e ga s p ro duc e d by .

th e m is l e d into th e apartm e nt thro ugh a k e yh o l e o o th e r r

s mall O p e ni ng I n thi s way th e o p e r at o r h as th e apparatus

.

und e r h i c o nt ro l du ri ng th e e nti r e o p e ratio n Th e ga s can

s .

also b e r e g e ne rat e d f r o m th e parafo rmald e hyd e o r th e so lid ,

fo rm I n this cas e th e whit e mat e rial whi ch is put up u nd e r

.
,

di ff e re nt nam e s is h e at e d o v e r a flame and bro k e n up into

,

th e gas . T h e gas may b e r e g e n e rat e d al so fr o m th e s o luti o n

whe n th e latt e r is spray e d in ve ry fi ne dro p s o n a la rg e surfac e .

I f th e dr o p s a re s mall a nd suffi ci e ntly s e pa r at e d f r o m o ne

an o th e r so that th e y d o no t run to ge th e r and h e nc e

,

e vap o r at e rapidl y littl e p o lym e rizati o n tak e s pla ce

,
Th e se .

principle s a re at th e basi s o f what is k no wn a s th e

C hicag o B o ard o f He alth m e th o d o f f o r mald e hyd e disi nf e cti o n .

This m e th o d is u se ful at time s but th difli cul ty co m e s und e r ,

e ,

o rdi nary c o nditi o n s in g e tti ng e no ugh f o r m ald e hyd e spray e d

,

o n to th e sh e e ts which a re u se d to pro duc e s ati s fa ct o r y di si m

fe ctio n b e f o r e it is ne c e ssa r y fo r th e o p e rat o r to l e av e th e r o o m .

I n many way s th e m o s t s tisfact o ry m e th o d o f r e g e ne r ating th e

a

gas is to bring it in co ntact with so m e ch e mical substance with

 RELATI O N TO ENV I R O NM EN T S I

whi ch e ith e r th e wate r in th e s o lutio n o r a part o f th e fo rm al d e

hyde r e cts p ro ducing h e at and th e e vo lutio n o f gas whi ch
a , ,

r e a ctio n lib e r at e s th e fo rmalde hyde ga s Th e sub stance mo s t.

f re qu e ntly u se d is p o ta ssium p e rmanganat e Whe n p o tassium

’

p e rmanganat e a nd fo rmalde hyd e a e b ro ught to g e th e r in th e

r

right p rOp o tio ns th e r e is a vi o l e nt r e actio n a nd th e fo rm al d e

r ,

hyde gas is rapidly give n o ff to ge the r with a co n side rable

,

amo unt o f mo i stur e A la rg e r am o unt o f fo r malde hyde is

.

n e e d e d to disi nfe ct a give n spac e b y thi s m e th o d than tho se

t

c o nside re d ab o ve b e caus e o f th e fact that pa r t o f it is wa st e d

in th e ch e mical c o mbi nati o n so that thi s m e th o d is s o m e what
,

m o r e e xp e nsive than o th e rs B ut d ue to th e fact that it can

.
,

b e us e d with o ut any sp e cial apparatus a nd that p r actically ,

th e mat e rials a re univ e rsally availabl e and that its e ff ci e ncy i

is ve ry high it is at th e pr e s e nt time th e m e th o d par e xce ll e nce

,
.

F o rmalde hyde gas i s e xc e e dingly e ffi ci e nt in it g e rmicidal s

actio n o n bacte ria I t has a m o r e o r l e ss s e l e ctive actio n and

.

do e s no t affe ct o r at l e as t r e adi ly many o th e r kind s Of ce lls

, ,
,

s uch as th o s e that c o mp o se th e tis s u e s o f high e r pla nts and

animals I t d o e s no t kill plants o r o rdina ry animals su ch

.
,

as ins e cts rats and mi ce in th e sam e co nc nt r atio n r e qui re d

, , ,
e

fo r th e d e structi o n o f bact e ria This se le ctive actio n is dis

.

advantage o us as w e ll as de sirabl e On acco unt o f this fact

.

sulphur is still u se d wh e r e it is n e ce ssary to kill o the r fo rms

o f lif e than bact e ria .

F o rmald e hyd e gas h as practically th e sam e sp e cific gravity

as air I t th e re fo re r e adily di ff us e s thro ugho ut th e air space

. .
15 2 GENERAL P HY S I O L O GY
It has ho w e ve r no mark e d p o w e r o f p e n e trati o n and in o rd e r
, , ,

to b e e ff e ctiv e all articl e s to b e di s m fe cte d mu s t hav e th e ir

,

s urfa ce s w e ll e xp o s e d a nd in its p r actical applicati o n it i s a

,

p o int o f co nsid e r abl e imp o r tance to se e that all mate rial s to

b e di si nf e ct e d are arra ng e d so that th e ga s can r e adily r e a ch
th e m Th e r e is so m e diff e re nce o f o pinio n a s to th e ne c e ssity
.

o f m o i s tur e I t is usually s tat e d that a c o nsid e rabl e am o u nt

.

o f m o i s tur e i s d esirabl e and t hi s i s pr o bably tru e ; but if th e

,

amo unt is suff ci e nt so that it co nde n se s o n an Obj e ct it can

i ,

b e v e ry r e adily s e e n that it might int e rf e r e with th e ab so rpti o n

o f th e gas .

T h e am o u nt o f f o r mald e hyd e to b e u se d in di sinfe cti o n i s a

matt e r o f imp o rtanc e Th e re is th e sam e dange r that no t

.

e n o ugh may b e us e d as th e r e i s in th e cas e o f sulphur In a .

g e n e ral way o ne pint o r p o und o f f o rmald e hyde S h o uld b e

us e d fo r e ve r y th o usa nd cubic f e e t o f space
.

Th e di sadvantag e s wh e n c o mpa r e d with sulphur a re fir s t

, ,

th e c o st w hi ch is s o m e what m o r e e sp e cially wh e r e u se d with

, ,

p o tassium p e r manganat e ; se co nd th e fact that it d o e s no t ,

a ffe ct animal lif e and th e r e fo r e is no t availabl e fo ce rtain

,
r

kinds o f di sinf e ctio n ; and thi rd th e stro ng Od o r which fre

,

qu e ntly p e rsi sts fo so m e p e rio d o f tim e B ut th e adv an

r .

tage s far o ffs e t th e se Th e Odo r whi ch p e r si st s aft e r fo rmal

.

d e h yd e h a s b e e n us e d may b e n e ut r aliz e d to a c o n sid e r abl e

e xt e nt by th e us e o f amm o nia Th e amm o nia is spri nkl e d

.

ab o ut th e r o o m fi v e parts b e ing us ed to fo ur parts o f th e fo r

,

m al d e h yd e .
 RE LATI O N TO E NV I R ONMEN T 15
3

Fo rmalde hyde may u se d as a liquid di sinfe ctant and is

be ,

str o ngly r e co mme nde d by many fo r th e purp o se o f disinf e cting

sputum urin e a nd fe ce s
, ,
.

I n th e pra ctical u se o f f o rmald e hyd e fo r disinfe cting h r o o m

o r an apartm e nt it is w e ll to r e m e mb e r that fir s t o f all th e

,

ro o m sh o uld b e tightly cl o se d all cracks ab o ut th e windo ws ,

and do o rs sho uld b e pa st e d up with ne w spap e r strips o r a ,

mate rial e sp e cially pr e par e d fo r th e pu rp o s e r e gi ste rs fi re , ,

place s e tc tightly cl o s e d l e aving o nly an e xit to b e made

,
.
, ,

tight o n th e o utside Al l articl e s in th e r o o m ne e ding di sin

.

fe ctio n sh o uld b e so arrang e d that th e ir surfac e s a e e xp o s e d r .

Thi s m e ans that Cl o thi ng sh o uld b e sp re ad o ut o n a lin e o r

o v e r chairs that th e b e ddi ng sh ould b e tak e n o ff a nd lik e wi se

,

S pr e ad ab o ut that th e pill o w s a nd ti cks S h o uld b e r e m o v e d

,

fr o m th e b e d and arrange d so that th e gas can r e adily r e ach

all parts o f th e m and if it is tho ught n e c e ssary an Op e ni ng
, ,

made so that th e gas can r e adily ge t int o th e inte rio r B u re au .

drawe rs trunks e tc must b e Op e ne d up and articl e s th e r e in

, ,
.
,

spre ad o ut Wh e re p o tassium p e r manganat e o r ano the r

.
,

ch e m ical is to b e use d fo r r e ge n e rating th e gas it is n e c e ssa ry

, ,

to hav e a la r g e c o ntai ne r Wh e r e a co nside rabl e am o unt o f

.

wo rk is to b e d ne it is de sirabl e to have a sp e cial pi e ce o f ap

o ,

paratus made su ch a s that sugg e ste d by th e I llino i s S tate

,

B o ard o f He alth Thi s co nsi st s o f a tin pail with l o w st r aight

.

side s surm o unt d by a flaring to p th e who l e to b e co ve r e d o n

e ,

th e o ut sid e with a sb e t o s pap e r a nd pr o vid e d with a d o ubl e

s

tin b o tto m Whe r e th e amo unt o f wo rk do e s no t warrant

.
1
54 GENERAL P HY S I O L O GY
g e tting thi s apparatus a go o d size d m e tal pail can b e use d
,
-

but it is b e st to place this in a m e tal wa shtub o r o n a pi e ce o f

a sb e sto s o f so me siz e The r e is by thi s me tho d co nside r abl e
.

dange r o f spu r ting d ue to th e vi o l e nt ch e mical actio n whi ch

,

go e s o n I t is th e re f o r e u nd e i r abl e to use m o re th a n thre e

. s

p o und s o f p e rmang nat e in o ne r e ce pta cl e ; if m o r e is n e e d e d

a ,

it is b e st to have ano th e r co ntaine r Th e ro o m o r apartm e nts

.

sho uld r e main cl o se d at l e ast e ight h o urs ; it will the n b e p o s

sibl e fo r s o me o ne t rush in and o p e n up a wi ndo w a nd th e n
o ,

as s o o n as p o sibl e th e spac e sh o ul d b e th o ro ughl y air e d a nd

s , ,

if th o ught n e c ssary th e O do r can b e n e ut r aliz e d by th e us e o f

e ,

ammo nia .

Al l di sinf e cti o n o ught to b e t e s t e d by m e an s o f culture s .

Th e t e st i s a v e ry simpl e o ne A standard o rgani sm su ch as

.
,

M py o g e ne s var au r e us is spr e ad o n ste ril e squar e s o f c o tt o n

. .
,

cl o th all o w e d to dry and p ut in a st e ril e t e st tub e Thi s

, ,
.

st e ril e t e s t tub e is plac e d in th e r o o m and th e co tto n plug

r e mo ve d Afte r di sinfe cti o n th e t e st o utfit is take n to th e
.

lab o rato ry and th e co tto n clo th i th e n dro pp e d into cultur e

s

m e dia I f th e m e dia r e main st e ril e th pro c e ss o f di sinfe c

.
,
e

tio n was satisfacto ry o th e rwise it is to b e r e p e ate d

,
.
 CHAP T ER XI I I
E FFE C T OF EXTE R NAL C ONDI TI O NS ON B AC TER I A
( C o nti nue d )

Li gh t.
—T h e r e latio n o f pro to pla sm to l ight varie s with
diff e re nt typ e s o f o rgani sms Light a s a fo r m o f radiant e ne r gy
.

is utiliz e d by ce r tain livi ng fo r m s in buil di ng up th e ir fo o d

mate r ial I n th e gr e e n plants fo r in s ta nce th e chl o ro phyll
.
, ,

e nabl e s th e o r ga ni s m to ab so r b a nd u se th e su n s e ne rgy in
’

co nstructing its fo o d mat e rial and th e g r e ate s t amo unt o f

,

assimilatio n o ccur s in th e ab so rptio n band s o f th e chl o r o phyll .

F o rm s that d e r ive th e i r e ne r gy f r o m o rganic s o urc e d o no t s

n e e d light C o l o rl e ss plants and animals o btai n th e i r e ne rgy

.

by br e aking do wn hi ghly co mpl e x f o o d sub stanc s and a e thus e r

inde p e nde nt o f th e e ne r gy o f light S inc e th e ba ct e ria a e

, . r

co l o rl e ss o rgani sms we wo ul d natu r ally e xp e ct that light wa s

no t e ss e ntial to th e m a nd thi s i s t r u e in th e mai n
,
T h e ma .

j o ri ty o f bact e ria g r o w b e s t in th e da r k A small

. gr o up o f

bacte ria that will b e r e f e rre d to lat e r o btain the i r e ne r gy mu ch

in th e same way as th e g r e e n plants d o a nd a re th e r e fo r e
de p e nd e nt up o n sunlight fo the i r e ne rgy Th e gr e at maj o rity
r .

o f ba ct e r ia h w e v e r p r e f e r da r k ne ss rath e r tha n light a nd fo r

,
o , ,

the s e fo r m s l ight is no t o nly u nne ce ssa ry but it may act as a

I SS
15 6 GENERAL P HY S I O L O GY
p o i so n Th e ge rmicidal p o we r o f light was fi r st cl e arly
di r e ct .

de fine d and p ro ve d e xpe rim e ntally by Do wns and B lunt in

187 8 .Thi s g e rmicidal actio n o f di re ct sunlight s e e ms to b e
particularly w e ll ma rk e d in th e ca se o f th o s e bact e ria which
pr o duce dise ase and is th e r e fo r e a matte r o f a gr e at de al o f

F I G 40
. . pp ara
— A f a g lati
e lt r f B a t ri
nce o e ne cu u e o c e um a thra i xp d t
n c s, e o se o

lig ht v r ly th ar a E d th i bat d f
th e o e o n e e , an en ncu e or 48 h r I n th ar a
ou s . e e

En l ni hav d v l p d (Fr m Ward

o co o es e e e o e . o ,

practical imp o rtance N o t o nly a re bacte ria r e adily kill e d

.

in di r e ct su nlight wh e n th e y a re in a v e g e tativ e s tag e but ,

M a r shall Wa r d has sh o wn that s o m e Of th e sp o r e s a re m o r e

quickly kill e d than a e th e ve ge tative ce lls r .

I n o r d e r to d e m o n s t r at e th e acti o n o f di r e ct su nlight o n

bacte ria it is o nly ne ce ssary to se e d a n agar plate h e avily

with th e o rgani sm to b e t e ste d Wh e n th e agar is tho ro ughly .
 EFFE CT OF E XTERNAL CO NDI TI ON S ON BACTERI A 1
57

so li di fi e d th e unde r side o f th e P e tri di sh is pa r tially co ve r e d

,

with so me o paque sub stance a s black pap e r Thi s may b e

,
.

put o n in th e f o rm o f characte rs as fo i nstanc e th e name

, ,
r ,

o f th e m i cro Orga ni sm u se d o r th e y e ar Th e P e tri di sh i s th e n

.

e xp o s e d to th e a cti o n o f di r e ct s unl ight fo r so m e tim e and i s

the n i ncubat e d I f th e e xp o sur e is sufl i cie nt th o se parts that

.
‘
,

have b e e n di re ctly e xp o se d will b e cl e ar whil e a gro wth will ,

app e ar o n th e p r o te cte d parts This acti o n is w e ll

.

illu s t r at e d in th e acco mpanyi ng fi gu re I n thi s e xp e r im e nt it

.

i s ne c e s ary fo r th e sunlight to pass thr o ugh th e gla ss a nd in

s ,

d o ing so a gre at d al o f th e e n e r gy is l o st Th e w o r k o f
e .

W e inz i rl was o f th e gre ate s t impo r tance in thi s co nne ctio n fo r ,

h e sh o w e d that it wa s p o s ibl e to e xp s e th e bact e ria witho ut

s o

th e i nt e rp o se d glas s di r e ctly to th e ray s o f th e sun and that ,

wh e n thi s was d o ne th e time r e q uire d to kill th e bact e ria was

ve ry much l e ss than had b e e n fo r m e rly supp o se d Th e acti o n .

o f th e sunlight i n kill i ng th e bact e ria is r e ma r kabl e and fre ,

que ntly an e xp o sur e o f o nly a fe w minut e s fi v e o r te n is , ,

sufficie nt to kill all o f th e g e rms AS indi cat e d ab o ve this

.
,

ge rmicidal actio n o f th e sunl ight o n bact ria is a matt e r o f e

g r e at hygi ni c imp o rtanc e and th e l ife o f th e di e ase pr o ducing

e ,
s -

bact ria o ut side o f th e b o dy is r e nde r e d ve ry diffi cult whe re it

e

i s p o s ibl e fo r th e di r e ct su nlight to ge t at th e m
s Undo ubt e dly .

th e d e s tructi n f g e r m s tak e s plac e in gr e t numb e rs i n na

o o a

tu r e pa r ticula rly in t r anspare nt sub stanc e s as wat e r B e

, ,
.

cause o i th e g e rmicidal a ctio n o f sunlight bact e ri in natu r al ,

a,

wate r s a l e s s abundant in th e upp e r lay e r s than at lo we r

,
re
 5 8 GENERAL P HY S I O L O GY
l e ve ls m o re in th e mo rning than at night ; and B uchne r de te r
,

mine d that th e light co uld p e n e trate wat e r to th e de pth o f a

m e te r o mo r e and that wh e n bacte ria w e r e e xp o se d at va rio us
r ,

d e pths the y w e r e aff e cte d by th e sunlight in inve r se p ro p o rtio n

to th e d e pth b e l o w th e s u r fac e Th e e ff e ct o f th e su nlight
.

va ri e s with its inte n ity and h e nce va ri e s with th e se aso n o f

s ,

th e y e ar th e tim e o f day a nd th e co nditi o n o f th e sky

, ,
In .

summe r th e acti o n is se ve ral time s as pro no unce d as in winte r .

A g r e at d e al o f di s cu ssi o n h a s o ccurr e d r e gardi ng th e m e th o d

o f th e acti o n O f s u nlight o n th e bact e ria At fir s t it was .

th o ught that p e rhap s it wa s d ue to h e at but it ca n b e r e adi ly ,

S h o wn by int e rp o si ng a s c r e e n
,
that th e actio n take s plac e
,

e ntir e ly i nd e p e nd e nt o f th e h e at I t h as b e e n f o und that it

.

vari e s with th e kind o f light ; that is if whit e light is b ro ke n ,

up into its co mp o n e nt parts c e r tain parts o f th e sp e ctrum are

,

much m o re e ff e ctiv e than o th e rs I f a scr e e n is place d b e tw e e n

.

th e sun and th e bact e ria which shuts o ut all but th e l e ss

,

r e fracte d rays such as may b e do ne by th e use o f a s cre e n

,

o f p o tas sium bichr o mat e it will b e f o und that th e r e is littl e o r

,

no acti o n ; that i s th e se r ays a re no t e ff e ctiv e

,
B ut if a .

scre e n is inte rp o se d b e twe e n th e bact e ria and th e sunlight

whi ch al l o w s th e m o r e highly r e fra cte d rays to pa ss thro ugh ,

such a sc r e e n fo instan ce a s o ne co mpo se d o f an amm o

,
r ,

nia cal s o luti o n o f Co pp e r sulphat e it is fo und that th e

.

maximum e ff e ct is Obtai ne d Or if a sp e ctrum is thr o wn o n a

.
,

he avily s e e de d agar plate it will b e fo und that th e gre ate s t

,

e ff e ct i s in th e blu e Vi o l e t rays that i tdi minish e s v e ry quickly

-

,
 EFFE CT OF E XTERNAL CO NDI TI O N S ON B AC T ERI A 1
59

on b o th side s and is practically n o thi ng in th e ult r a vi o l et o r

,
-

in th e e d ray s r Th e e xact p o i nt wh e r e acti o n b e gi ns and e nds

.

di ff e rs with vario us f o rms but ge n e rally it b e gins in th e blu e ,

e nd o f th e gr e e n and r e ach e s a maximum in th e vi o l e t a nd ,

di mini sh e s fro m th e vi o l e t to th e ultra vi o l e t ( F ig Th e -

.

FI G 4 1
. . Plat
a thrax p r x p d f 5 h r t th lar p tr m in
e o f n s o e s, e o se or ou s o e so s ec u

g t d i bat d f 4 8 h r Th h riz tal li h w th l g th f

Au us , an ncu e or ou s . e o on ne s o s e en o

th p tr m Th v rti al li
e s ec u t F
. li b t rv t
e e c ne s a re no R AUE N H O F E R S
’
ne , u se e o

Sh w th li m it f th pri ipal r gi f th p tr m Th l ar t ar a i
o e s o e nc e o ns o e s ec u . e ce es e s

that wh r f w t p r hav d v l p d in th i bati —wh r

e e e es s o es e e e o e e ncu on e e, co u se

q ntly th ba t ri idal ff t w gr at t ( Fr m Ward

ue ,
e c e c e ec as e es . o ,

same kind o f actio n is o btaine d by e l e ctric light altho ugh , ,

o f co u r se th e e ne rgy is v e ry mu ch l e ss
,
.

E l e ctri ity A g r e at d e al o f i nt e r e s t h a s b e e n manif e st e d

c .

in th e e ff e ct o f e l e ct r i city up o n bact e r ia a nd a g r e at ma ny ,

e xp e rim e nts hav e b e e n p e rf o r m e d to d e t e rmine what thi s e f

1 60 GE NERAL P HY S I OL O GY
fe e t is Aside fr o m th e sci e ntific asp e ct it has b e e n th o ught
.
,

that much practical g o o d might c o m e fro m th e use o f e l e ct ric

ity as a ge rm icidal age nt but S O far th e e xp e rim e nts have
,

b e e n disapp o inting Th e r e is no do ubt but that s tr o ng cur

.

r e nts o f e l e ctricity w o uld aff e ct bacte ria much in th e same way

th at it do e s o th e r fo rms o f lif e but it is ve ry di fli cul t to apply
,
‘

it I f fo r instance a di re ct cu rr e nt is run thr o ugh a cultur e

.
, ,

m e di um th e e l e ctro lytic e ffe ct o n th e m e dium is v e ry mark e d

, ,

and th e r e sulting actio n o n th e bacte ria is d ue to th e ch e mi cal s

f o rm e d and no t to th e cu rr e nt itse lf An i nt e r e ti ng co nt ri
. s

b utio n to thi s subj e ct was mad e by Z e it in 9 and hi s

1 0 1,

c o nclusio ns are as fo ll o ws :
I A c o ntinu o u s curr e nt o f 2 60 to 3 2 0 milliamp e r e s
.

pas se d thr o ugh b o uill o n culture s kills bact e ria o f l o w th e rmal

de ath p o ints in te n minut e s by th e pro ductio n o f h e at 9 8 5 C .
°
.

Th e antis e ptics pro duc e d by e l e ctr o ly sis du ring thi s tim e

a re no t suffi ci e nt to pr e v e nt th e gr o wth o f e v e n no n sp o r e -

b e aring bacte ria Th e e ffe ct is a pu r e ly phy sical o ne

. .

2 A c o ntinu o us curr e nt o f f o rty e ight milliamp e r e s

.
-

pass e d thro ugh b o uill o n cultur e s fo r fro m two to th r e e h o u rs

do e s no t kill e ve n no n r e si stant f o rm s o f bacte ria Th e
-
.

t e mp e ratur e pr o duce d by such a curr e nt do e s no t ri se ab o ve

3 7 C and th e e l e ctr o lytic pr o ducts a re antis e ptic but no t
°
.
,

ge rmicidal .

3 A c o ntinu o u s cur r e nt o f 100 milliamp e r e s pa sse d

.

thr o ugh b o uill o n culture s fo r 7 5 minut e s kill s all no n r e si stant-

fo rms o f bact e ria e v e n if th e t e mp e ratur e is artificially k e pt

,
 E FF EC T OF E X TERNAL COND I T IONS ON BA CTE R IA 1 61

b e l o w 3 7 C Th e e fi e ct i d ue to th e fo rmatio n o f ge rmicidal
°
. s

e l ctr o lyti c pr o duct s in th e cultur e

e Anthrax sp o r e s a e kill e d
. r

in two h o u r s S ubtili s sp o r e s w e r e still al iv e aft e r th e cu r r e nt

.

wa s pass e d fo thr e e h o u r s
r .

4 . A c o nti n u o u s cu r r e nt pas s e d thr o ugh b o uill o n cultur e s

o f bact e ria p r o duc e s a st r o ngly a cid r e acti o n at th e p o sitiv e

p o l e d ue to th e lib e ratio n o f chl o ri n whi ch co mbine s with o y

, ,
x

g e n to f o r m hyp o ch l o r o u s acid T h e s t r o ngly alkal i ne r e acti o n

.

O f th e b o uill o n cultu r e at th e ne g tiv e p o l e i s d ue to th e fo r

a

matio n o f so di um hydro xid a nd th e lib e r ati o n Of hyd ro ge n in

gas bubbl e s With a curr e nt Of
. mill iamp e r e s fo r two
100

ho urs it r e quir e d
,
mil ligrams o f H 80 to ne utraliz e c c 2 4 I . .

Of th e c ultur e fluid at th e ne gativ e p o l e a nd all th e m o s t r e sist ,

ant fo rms o f bact e ria w e r e de stro y e d at th e p o sitive p o l e ,

including anthrax a nd subtilis sp o r e s At th e ne gative p o l e .

a nthrax sp o r e s w e r e kill e d als o but subtili s sp o re s r e mained

,

al ive fo r f o ur h o urs .

5 . T h e c o nti n u o u s curre nt al o n e by m e ans o f Du B is ,

o

R e ym o nd s m e th o d Of no np o larizing e l e ct r o d e s a nd e xclu si o n
’
,

O f ch e mi c e ff e ct s by i o ns in K rug e r s se nse i s ne ith e r ba ct e ri

’
,

cid al no r anti se ptic T h e appa r e nt a nti se ptic ff e ct o n susp e n

. e

sio n o f bacte ria is d ue to e l e ctri c o mo se Th e co nti nu o us s .

e l e ctric curr e nt h s n a b cte ricidal no r antise ptic pro p e r tie s

o a ,

but can de st ro y bacte ria o nly by its phy sical e ff e cts (h e at)
o r ch e mi c e ff e ct s ( th e p r o ducti o n o f bact e r icidal substanc e s

o f e l e ctr o lysi s) .

6 A magn e tic fi e ld e ith e r with in a h e l ix o f wir e o r b e tw e e n

.
,

H
1 62 GENERAL P HYSIOLOGY
th e p o l e s Of a p o w e rful e l e ctro magn e t has no antis e ptic o r
,

bact e ricidal e ffe cts whate ve r .

7 A
. lt e rnating curr e nts o f a thr e e i nch R uhmk -
o r ff c o il
pass e d thro ugh b o uill o n cul tur e s fo r te n h o urs favo r gro wth
and pigm e nt pro ductio n .

8 High fr e qu e ncy high p o t e ntial cur r e nts

.
,
— T e sla cur
r e nts hav e n e ith e r anti se ptic no r bacte ricidal pro p e rti e s
wh e n pa ss e d a r o und a bact e rial s usp e nsi o n withi n a s o l e no id .

Wh e n e xp o se d to th e brush di scharg e s o zo ne is pr o duce d and

,

kills th e bacte ria .

9 B o
. uill o n and hydr o c e l e fluid cultur e s in t e st tub e s o f
no n r e si s tant f o rms o f bact e r ia c o uld no t b e kill d by R o ntg e n
-
e

rays aft e r fo r ty e ight h o ur s e xp o sure at a di stance o f 2 0 mm

-
’
.

fro m th e tub e .

10 S usp e n si o n s o f bact e ria in agar plat e s and e xp o s e d fo r

.

four h o urs to th e ray s acco rdi ng to R i e de r s plan w e r e no t

,
’
,

kill e d .

1 1 Tub e rcular sputum e xp o se d to th e R o ntge n rays fo r

.

six ho urs at a distance o f m m fr o m th e tub e cause d acut e

,
2 0 .
,

miliary tub e rcul o sis o f all th e guine a pigs in o culat e d with it .

”

R on tge n R ay s I n spit e O f ma ny e xp e r im e nts and th e

.

ho p e ful vi e w o f many it h as b e e n v e ry d e finite ly sh o wn that

,

th e R o ntg e n ray s hav e no bact e ricidal pr o p e rti e s B act e ria .

m ay b e e xp o se d to th e se r ays in cultu r e m e dia fo r practically

inde finit e p e rio d s witho ut b e ing a ff e cte d Th e favo r abl e .

clinical r e sults whi ch hav e b e e n o btaine d in th e use o f th e se

rays in infe ctio us dis e as e s must b e e xplain e d up o n s o m e o the r
 E FF E CT OF E X TERNAL CON D IT I ONS ON BA C T ERIA 16
3

basis I t may p o ssibly b e d ue to th e pr o ductio n o f ce rtain

.

ch e mical substanc e s in th e ti ssu e s as o zo ne o r hyp o chl o ro u s

,

acid o r by a stimulatio n Of th e ce lls to re n e we d activity o r

, ,

an incr e ase d phago cyto si s S o m e e xp e rime nt s have Se e me d

.

to i ndi cat e that pr o l o ng e d e xp o sur e o f cultur e s to th e X r ays -

may sligh tly di minish th e vitality and virul e nce o f th e

g e rms .

M o ve m e nt B acte ria appare ntly gro w b e tte r wh e n th e y

.

a re in a stat e o f r e s t Exp e rim e nt s S h o w that S light m o v e

‘
.

m e nts are no t imp o rtant in aff e cting th e gro wth o f bact e ria ,

but that mo r e vio l e nt m o ve me nt such as , fo r instance might

, ,

b e Obtai ne d by attaching th e cultur e to th e pisto n o f a rapidly

mo ving ste am e ngine o r a S haking machi n e d o e s have an in ,

bi bit o ry acti o n A practical phas e o f thi s subj e ct manif e sts

.

itse lf in a di scussio n Of th e qu e stio n o f th e e ff e ct o f th e mo ve

m e nt o f a st re am o n th e co ntain e d bact e ria I t app e ars fro m .

o ur pr e s e nt kn o wl e dg e that such m o v e m e nt c o uld no t b e much

o f a fact o r in th e purificati o n o f a str e am .

P re s s ur e .Ordinary variati o ns in pr e s sure d o no t a ff e ct

e ith e r th e lif e o r th e virul e nc e o f bact e ria and th e r e is s o m e
,

qu e stio n wh e th e r e xtre me ly high pr e ssur e s aff e ct th e c e ll .

R uss e ll in hi s w o rk at N apl e s f o und th e sam e S p e ci e s o f bac

, ,

te ria b o th at th e surfac e and in de e p —se a dre dgings O the rs .

,

o n th e o th e r ha nd s ay that 6 ,
to 7 atmo sph e r e s Of pr e ssure
00 00

h av e an inhi bit o ry e ffe ct o n putre fa ctiv e pro c e ss e s R o ge r .

claims that a pre ssure O f 2 000 atmo sph e r e s inte rf e r e s with th e

virul e nce Of th e B act anthracis . .
1 64 GENERA L P HY S I OL O GY
As so ci ati o n
. B act e ria are undo ubte dly gr e atly affe ct e d
by the ir asso ciat e s Thi s a sso ciatio n may b e with o th e r
.

bacte ria o r with highe r plants and animal s I n p e rhap s th e

.

maj o rity o f case s no influ e nce is e x e rte d by o ne o rgani sm

up o n an o th e r but ve ry fre que ntly o ne sp e ci e s r e tard s th e
,

gro wth O f an o th e r ; that is th e re is a ntago ni sm Thi s is v e ry

,
.

fre que ntly s e e n in cas e s wh e r e a culture m e dium is ino culat e d

with s e ve ral di ffe r e nt Sp e ci e s One sp e ci e s g e ts th e upp e r
.

hand and gro ws fo r a p e ri o d to th e practical e xclusio n Of all

o th e r s ; th e n c o n di tio n s chang e a nd ano th e r f o rm app e ar s
,

and so o n That the r e is a v e ry mark e d antago ni s m by s o me

.

bact e ria fo r o the rs is sh o wn by a numb e r o f e xp e rim e n ts that

have b e e n made whi ch cl e arly de mo nst r at e th a t One ge rm may
c o mpl e t e ly d e st ro y an o the r pro bably thro ugh th e b y pro ducts
,
-

which it pro duc e s Exampl e s o f thi s have b e e n in stanc e d by

.

a numb e r Of inve stigato rs Th e asso ciatio n o f o rganisms at

.

tim e s is advantage o us to o ne o r b o th Th e asso ciatio n o f th e

.

r o o t n o dul e bact e ria with th e l e gumin o us plant is a cla ssical

-

e xampl e o f sym bi o si s and it i s th o ught that many bact e ria

,

may live in symbio tic r e latio nshi p with o th e rs e sp e cially ,

c e rtain anae ro b e s and ae r o b e s Our kn o wl e dg e o f th is is ve ry

.

inco mpl e te and fragme ntary .

 C HAP TE R X I V

M E TA B O LI S M OF BAC T ER IA

I ntro d u cti o n pre vi o us chapte rs we hav e co nside r e d

. In

th e r e lati o n O f bact e ria to th e ir e nv r o nm e nt

i W e hav e no w .

to c o n sid e r th e p r o ce ss e s inh e r e nt in th e bact e rial c e ll that i s , ,

th e fu ncti o ns o f th e c e ll a nd und e r this will c o m e a discussi o n

,

of : I ,th e g e n e ral charact e r o f th e lif e pr o c e ss e s ; 2 s o urc e s ,

and e xp e nditure s Of e ne rgy ; and 3 r e spirati o n Th e pr o d , .

ucts o f m e tab o lism wil l b e l e ft fo r di scu ssi o n in a s e parat e

chapte r .

G e ne al Ch aracte r Of Life P r o ce s s e s
r
— Th e w o rk o f th e .

living c e ll is d e t e r mine d in pa r t by its e nviro nm e nt but ce rtain ,

functi o n s b e l o ng to th e ce l l and withi n o rdinary limits are no t

,

pro fo undly aff cte d e xce pt a s to rat e by e xt e rnal co nditio ns

e , , .

The se g e ne r al functio ns a re : 1 assimilatio n ; 2 re spiratio n ;

, ,

a nd 3 gro wth a nd m o v e m e nt
,
T o th e e g e n e ral functi o ns a re
. s

to b e add e d th e sp e cializ e d a ctiviti e s o f th e c e ll that have b e e n

acquire d sub se que ntly in th e de ve lo pm e ntal hi s to ry o f th e

.

o r ga ni m s S u ch sp e cial fu ncti o n s are pigm e nt pro ducti o n

.
,

f e r m e ntatio n di se ase pro ducti o n e tc Al l Of th e life pro ce sse s

, ,
.

Of th e ce ll a re e mbrac e d in th e t e r m m e ta bo l i s m w hi ch acc o rd , ,

ing to Vi ne s include s all Of th e ch e mical chang e s whi ch go

,

165
1 66 GENERAL PHYSIO L OGY
on in th e living pro to plasm Th e living ce ll is in a state o f un
.

s tabl e e q uilibrium P r o t o pla s m is b e ing c o nstantly built up and

.

just as co n stantly br o k e n do wn The se two pr o ce sse s th e

.
,

bui ldi ng up and th e t e ari ng do w n always gO hand in hand , .

Th e bu ilding up c o nst r uctiv e o r synth e tic pro c e ss is give n

, ,

a sp e cial nam e a na bo l i sm Th e te a ring do wn de structive

,
.
, ,

o r analytic pr o c e ss is sp o k e n o f as ca ta bo l i s m .

Anab o li s m . Anab o l i sm includ e s acco rdi ng to Vin e s , ,

all o f th e ch e mical pro ce sse s go ing o n in th e ce ll wh ich l e ad to

th e f o rmati o n o f c o mpl e x sub s ta nce s fr o m simpl e o n e s Th e .

ce ll is co ns tantly abs o rbing raw mate rial as fo o d as similating ,

it a nd b uil di ng it up into pro to plasm Th e e xact way in which

, .

assim ilatio n is acco mplishe d is unkn o wn Th e r e quire me nt s .

O f bact e r ia s o far as th e ir f o o d is c o nc e rn e d is fir s t that it b e

, ,

s o lubl e altho ugh it is true a s will b e p o i nt e d o ut late r mo re

, ,

S p e cifically that th e bact e ria a re abl e to pr o duc e s ubsta nc e s

,

which dis so lve in so lubl e mate rial a nd make it p o ssible fo r

the m to tak e up sub s tance s that a e no t o lubl e Th e s e c o nd
r s .

r e quir e m e nt is that th e y have th e right kind o f f o o d and th e ,

bacte ria a s a who le ca n make use f a gre at many di ffe r e nt O

ki nds o f f o o d a nd may b e clas sifi e d o n thi s basis B act e ria

,
.
,

with th e e xc e ptio n O f o ne gr o up a re unabl e to make use O f th e

,

carb o n di o xide o f th e air as a s o urce o f carb o n S om e bac .

t e ria are able to liv e o n d e ad o rgani c matte r ; ano th e r gro up

r e quire that th e ir f o o d b e livi ng o rgani c matt e r A cl assifi ca .

tio n O f bacte ria acc rdi ng to th e ir nutritive r quir e me nt has

o e

b e e n pr p o s e d by F i sch e r a nd is as fo ll o ws :
o ,
 ME TAB OLISM OF B A C T ERIA 167

I . ro to tro phi c B acte ria

P .

N i trifyi ng ba ct e ria bact e ria o f ro o t n o dul e s sulphur and

, ,

iro n bacte ria o ccur o nl y in th e Op e n in nature n e ve r parasitic , ,

always mo no tr o phi c .

II . M e tatro p hi c B acte ria .

Z ym o ge nic sapro g e ni c and sapro phytic bacte ria ; o ccur in

, ,

th e o p e n and up o n th e e xt e r n l a nd int e r nal s urfac e s o f th e

a

b o dy s o me time s parasitic (fa cul tativ e p arasite s) m o no tro phi c

, , ,

o r p o lytr o phic .

I II aratro phi c B acte ria

. P .

O ccur o nly in th e tis s u e s and v e ss e ls O f living o rganisms ,

true ( Obligato ry) parasite s .

C a ta b o l i s m B y catab o li s m is m e ant all th e ch e mical

.

pro ce sse s g o ing o n in th e ce ll which l e ad to th e fo rmatio n o f

S impl e sub s tance s fro m co mpl e x o n e s ( Vine ) Th e ce ll s .

pro to plasm is co ns tantly b e ing br o ke n do wn a nd b y—pro ducts

f o rm e d Anab o li s m and catab o li sm a re co nstantly go ing o n
.

in th e s ame ce ll I f o rganic matte r o f th e ce ll is o f a hi gh e r

.

sta te than th e fo o d fr o m which it wa s fo rme d th e anab o lic ,

pr o ce ss has b e e n gre ate r than th e catab o lic p ro ce ss and th e ,

o rganism that builds up o rganic matt e r can b e pr o p e rly sp o k e n

o f as an anab o lic o rgani s m I f a s a r e s ult o f S i m ilar pr o c e s se s

.

in an o rgani sm th e e nd pro ducts are l o we r than tho se o f th e

,

f o o d th e o rga m sm 8 said to b e catab o li c ; S O that whil e th e

,
1

anab o lic and ca tab o lic pro ce sse s are co ncur r e nt in th e same
1 68 G ENERAL P HYSIO L O GY
c e ll r e sult is e ith e r o ne o r th e o th e r and o rgani sms may
th e ,

b cl a
e sse d a s anab o lic o rga ni s ms o r as catab o l ic o rga ni sms .

Th e anab o lic o rgani sms a re th e gr e e n plants a nd th e p r t o o

tro phi c bacte ria Th e catab o lic o rganisms a e th e ani mals . r ,

fungi m e tatro phic a nd paratro phi c bacte ria The se di ffe re nt

,
.

FI G 4 2
. .
— I llustrating M e tabo li sm t a ll AB r p . Th e s u q are r pr e e se n s ce . e te

t f d t ri g th ll B C r pr nt a ab li m CD r pr t
se n s oo en e n e ce . e e se s n o s . e e se n s

catab li m B CD r pr t m tab li m DE r pr t th pr d t
o s . e e se n s e o s . e e se n s e o uc s

Of m tab li m e b r pr t f d f i m pl m p iti
o s . d r pr a t e e se n s oo o S e co os on . e e e se n s

co m pl x b y pr d te A rg a i m pr d i g ynth ti ha g
-
o indi at d by
uc s . no n s o uc n s e cc n e s, c e

th lin i all d a ab li rg ani m An rg a i m pr d i g th

e e ao s c e an n o c o s . o n s o uc n e ana

lyti ha g indi at d by th li
c c n e s, i all d a atab li c e e ne or a o
' '
s c e c o c

org ani m s .

classe s Of o rganisms are Of mutual aid to e ach o th e r in th e

unive rse Th e anab o lic o rgani sms build up th e f o o d mate rial
.

f ro m simpl e so urce s i nto a fo rm th at is availabl e fo r th e cata

h o lic o rgani s ms Th e se in turn te ar do wn th e co mpl e x .
, ,

o rganic matt e r and r e turn it to th e e arth in a S impl e f o rm

r e ady to b e again tak e n up by th e anab o lic o rgani sms .

R e spira ti o n — F o o d is n e ce ssary to s u s tain lif e but f o o d

.

al o n e will no t e nabl e lif e to go o n F o o d co nsum e d p re . re

s e nts a s o urce o r supply o f e ne rgy but this is no t availabl e ,

until it is o xidize d and th e p o t e ntial e ne rgy l o cke d up in th e

fo o d co nsum e d is transfo rm e d into th e kin e tic e n e rgy O f wo rk .
 ME TAB OL I S M OP B ACTERIA f1 69

C o al r e pr e se nts st o r e d up e n e rgy but we must transf o rm th e

-

, .

late nt e ne rgy that is impri so ne d in th e co al into th e active

e ne r gy o f m o ti o n b e f o r e it b e c o m e s availabl e This is do ne by .

co mbustio n o o xidati o n a nd by placi ng th e co al urfd e r co n

,
r ,

d itio ns wh e r e th e afli nity o f th e ca rb o n fo r o xyg e n will b e

satisfi e d and th e kin e ti c e ne rgy will b e r e l e ase d in h e at Th e .

fo o d co nsume d by all o rgani sms functio ns like co al and th e ,

pro ce ss by which it is o xidize d is kn o wn as r e spiratio n I t .

was fo r me rly tho ught that o xy ge n in a fr e e stat e was ne ce s

sary but it is no w kno wn that it is no t Th e o xyg e n may b e
,
.

Obtai ne d f r o m o rga ni c c o mp o u nd s as in th e case o f th e

,

ana e ro b e s but th e amo unt Of e n e rgy Obtai ne d wh e n th e o xy

'

g e n i s d e riv e d i n th i s w a y is much l e s s than wh e n th e fr ee

o xyg e n i s u se d F o r e xampl e wh e n 100 grams Of sugar

.
,
0

a re o xidiz e d to ca rb o n di o xid e and wat e r 3 93 9 cal o ri e s Of h e at

a re Obtain e d but wh e n th e s am e am o unt o f sugar is o xidiz e d

,

to butyri c a cid hyd r o g e n a nd carb o n di o xid e o nly 4 4 cal o ri e s

, , ,
1

a re O btai ne d AS a r e sult O f th e pr o c e s s O f r e spirati o n ca rb o n

.

di o xide is uni ve rsally pro duce d as w e ll a s wat e r I n addi tio n

,
.

to thi s e n e rgy i s di ssipat e d in th e f o r m O f h e at light and

, , ,

m o tio n I n th e cas e o f th e bacte ria th e h e at f o r m e d du ring

.

th e p r o c e s s O f r e spirati o n i s no t appar e nt o r e asily m e asur e d

e x ce pt in c e rtain c o ndi ti o n s wh e r e a high d e gr e e O f h e at is

pro duce d F o r i nstance in ce rtai n fe rme nting masse s th e

t

.
,

t e mp e rature may b e raise d to 6 o r 7 0 C and th e bacte ria 0

° °
.

have so me r e latio n to th e sp o ntan e o us co mbustio n Of hay ,

c o tto n waste e tc En e rgy is s o me time s give n Off in th e fo r m

,
.
I 7 0 GENE R AL P HY S I OLO GY
of light as in ph o sph o re sce nce This characte ristic is co m
,
.

m o n in va r i o us fo rms o f lif e particularly in marin e animal

,

fo rms amo ng th e pho to bacte ria The se a re sapro phyt e s

.

whi ch live in salt wate r and gro w e sp e cially o n p e pto ne s s uch ,

as are co mmo n in de ad fi sh Th e y pro duce light o nl y in th e

.

pr e s e nce o f o xyge n and th e phe no me no n o f pho spho r e sce nc e

,

is intimate ly asso ciate d with r e spirati o n Th e e n e rgy Ob

.

taine d as a r e s ult o f r e spiratio n sh o w s its e lf als o in m o ti o n ,

and a go o d e xampl e O f this is fr e que ntly s e e n in hanging

dro p pr e paratio n whe r e th e a éro b ic bact e ria l o se th e ir p o w e r
,

O f m o ti o n wh e n fr e e acc e s s to th e o xyg e n o f th e air is d e ni e d

the m .
 C HAP TER XV

P R ODU C TS OF M E TA B O L I S M

In tro d ucti o n B act e ria di ff e r fr o m o the r plants in g e n e ral

.

in that th e ir catab o lic p r o p e r ti e s e xc e e d th e ir anab o lic

,
.

The y are p re e mi ne ntly analytical rathe r than sy nthe tic

Or ga ni sms I n th e m ul tipli cati o n o f c e ll s a la rg e amo unt o f
.

o rganic matt e r i s ac cum ul at e d i n th e pro to plasm but this,

is r e lative ly small co mpa r e d with th e di sinte grating pr o ce ss e s

which th e y are able to inaugurate I t is pro p o se d in this .

chapt e r to co nside r so me Of th e pro ducts Of m e tab o lis m prin ,

cip al l y in o r d e r that we may u nd e r s tand so m e thi ng m o r e o f

,

th e natur e o f th e chang e s whi ch go o n in th e bact e r ial c e ll ,

but als o b e caus e th e de t e rminatio n o f s o me o f the s e b y pro d -

ucts i s a matt e r O f gr e at imp o rta nc e in th e di ff e r e ntiati o n

o f vari o u s bact e ria I n c o nsid e ri ng th e p r o ducts o f m e tab

.

o l i sm,
ho we ve r it will b e d ifli cul t to di stingui sh b e twe e n
,

tho se whi ch a e p ro duce d withi n th e ce ll a nd th o se sub stance s

r

whi ch a e fo r me d o utside o f th e ce ll as a r e sul t o f th e b r e ak

r

ing do wn Of th e fo o d sub s tan ce o th e ab st r actio n O f c e rtain

r

e l e m e nt th e r e f r o m
s I n c e r tain ca s e s it i s imp o ssibl e to k no w
.

whe the r a pa r ticula r p ro duct is actually fo r me d withi n th e

ce ll o r whe the r it is a split pro duct Ve ry fre que ntly fr o m .
,

17 1
 7 2 G ENERAL P HY SIO L O GY
th e sam e nutri e nt m e di um a gr e at vari e ty o f pro ducts is
,

pr o duce d by a pa r ticular o rgani sm I t ve ry f r e que ntly hap .

p e ns ho w e ve r that o ne is m o re p ro mine nt than th e o th e r s

, , ,

and ce rtain bacte ria have c o m e to b e kno wn a s bacte ria which

pro duc e c e r tain cha r a ct e ristic b y pr o ducts as fo r i nstance
-

, , ,

th e ac e tic acid th e lactic acid bact e ria ; but in all o f th e s e

,

cas e s o the r p ro duct s are fo rm e d as w e ll Th e se a re o nl y th e .

m o st striking b y—pr o ducts Th e pro ducts p ro duce d by th e

.

bacte ria u sually i nhibit th e ir gro wth if th e y are all o we d to

a ccumulate S o m e tim e s th e b y pro ducts are O f such a na
.
-

tur e that th e y actually de st ro y th ce ll s which pro du ce th e me

aft e r a co n side rabl e amo unt o f th e sub sta nce has b e e n p ro

d uce d . Th e am o u nt o f s ub s tanc e whi ch di ff e r e nt bact e ria a e r

abl e to to l e rate vari e s with diffe r e nt sp e ci e s F o r e xampl e .

,

th e l actic acid bact e r ium h as i ts g r o wth i nhi bit e d wh e n p e r

c e nt Of acid is fo r m e d Th e ace tic acid bact e rium ca n pr o duce

.

10
p e r ce nt o f a ce tic acid b e f o r e i ts gr o wth is p r e v e nt e d and ,

y e ast can pro duc e alc o ho l t th e e xt e nt f 5 p e r c e nt b e fo r e

o O 1

its furthe r d e v e l o pm e nt is i nhibit e d B y r e m o vi ng th e so l .

ubl e pr o ducts it is p o s sibl e fo r th e bact e ria to co nti nue the ir

activity Thi s can b e do n e by ne ut r alizing th e m with che m
.

ical sub stance s o r in so m e case s by gr o wi ng th e o rgani sm in

,

a p e rm abl e sack such a s o ne made O f co ll o dio n Th e b y

e ,
.

p ro ducts h e re are r e m o v e d by dialy sis p e rmitting th e furthe r ,

gro wth Of th e o rgani sm which may co ntinue u ntil all th e fo o d

,

substanc e s are us e d up I n c o n sid e ri ng th e pro ducts o f

.

m e tab o lism att e ntio n S h o uld b e call e d to th e di ff e r e nce

,
 P RO DUC T S OF ME T AB OLISM 1 73

b e tw e e n th e e xtra c e llular a nd th e int r a ce llular pro ducts

‘
- -
.

M any pro ducts a re f o rm e d that are Of su ch a natur e that th e y

ca n pass thr o ugh th e c e ll wall r e a d ily a nd a re e xcr e t e d The se .

a e kno wn as e xtra c e llul a r pr o duct s and a re to b e distin

r -

g u i sh e d sharply fr o m th e S plit pr o ducts whi ch a re r e f e r r e d to

ab o ve M any o f th e pr o ducts o f m e tab o li sm ho we ve r are

.
, ,

O f such a natur e that th e y d o no t r e adily pass thr o ugh th e

c e ll walls and a re lib e r ate d o nl y o n th e de ath o f th e ce ll .

The se a e th e i ntra—ce ll ul ar p ro ducts Al l c e lls in th e ir

r .

m e tab o lism p ro duce c rtain b y pr o ducts such as carb o n

e -

di o xid e w t e r nit ro g e n wa st e pr o du cts and e n e r gy B ut in

,
a , , ,
.

additio n to the se many o rgani sms pro duce in co nside rabl e

,

amo unt de fi nit b y pro ducts by which the y are characte r

e -

i ed
z . M any o r gani sm s fo r i nstan ce pr o duc e substanc e s
, ,

whi ch have th e p w e r O f r e ducing o th e r substance s Th e se

o .

r e da cti o n s ubsta nce s are f o und am o ng th e d e nitifying bac

t e ria wh e r e th e nitrate s are r e duce d to th e nitrite s a nd al so ,

whe r e co l o r e d sub ta nce s a e r e duce d to co l o rl e ss o l e uc o

s r r

p ro ducts An xampl e o f thi s is s e e n in th e gro wth Of many

. e

bacte ria in litmus milk Th e co l o ro f th e litmu s is fr e que ntly

.

l o s t co mpl e te ly d ue to th e fo rmatio n o f s o me r e duci ng sub

,

s ta nce . C ultu r e m e dia c o ntai ni ng m e thyl e ne b l ue a e al so r

r e duc e d by c e r tain bact e ria N e utral e d wh e n adde d to

. r ,

cul tu re m e dia is r e duc e d by c e rtain f o rm s and thi s r e a ctio n

, ,

is ve ry us e ful in th e di ffe r e ntiati n o f va r i o u s bact e ria o It .

was fo rm e rly supp o se d that this r e ductio n wa s d ue to th e split

ting Off o f o xyge n fr o m th e r e ducibl e sub stanc e but it is no w ,
I
74 GENERAL P HY S I OL O GY
kn o wn that thi s e xplanatio n is no t sufli cie nt since so me ,

substance s a re r e duc e d as fo r e xampl e m e thyl e n e blue

, , , ,

whi ch d o no t co ntain o xyge n I n the s e cas e s it may b e that

.

nasce nt hydro ge n i s adde d C e rtain bact e ria p ro du ce .

o xi d ati ve ch nge s
a Th e ace tic acid ba ct e ria o xidiz e e thyl
.

alco ho l to ace tic acid Th e nit rifyi ng bacte ria o xidize

.

ammo nia and th e nitrite s to ni trat e s Th e sulphur bacte ria .

o xidiz e sulphur e tt e d hydr o g e n to sulphur and th e ir o n bac

,

t e ria o xidiz e th e f e rro us to th e fe rric salt s Gase s a re i re .

que ntly pro duce d as th e r e s ul t o f th e me tab o lism o f bact e ria .

Am o ng th e m o r e c o mm o n ga se s a re ca rb o n di o xid e hydr o ,

g e n,
nitr o g e n ,
m e tha ne a n d
,
amm o nia Th e s e. ga s e s a re

fr e que ntly o f a go o d de al o f value in di ff e re ntiating o ne o r

g a ni sm fr o m an o th e r C l o s e ly alli e d bact e ria di ff e r gr e atly
.

in the ir p o w e r to f e rm e nt ce r tain substanc e s One may p ro .

duce a n abundance Of gas in a fo o d m e di um in whi ch a cl o s e

r e lative will pro duce n o ne Th e gase us b y pro duct s o f
. o -

bact e ria have b e e n car e fully studi e d in a numb e r o f ca se s ,

but th e co mpl e te analysi s o f th e gas e s is a matt e r o f co nsid

e rab l e ch e m i cal di fficulty An app r o xim at e m e th o d h o w
.
,

e ve r ,
fo r d e t e r m ining th e am o un t o f ga s pr o duc e d a nd its

f o rmula is ve ry co mm o nly u se d in th e bacte rio lo gical lab o ra

,

t o ry and is kno wn as th e f rm e ntatio n tub e which was intro

e

d uce d fo this purp o s e by Th e o bald S mith

r .

B act e ria ve ry f r e qu e ntly pr o duce acid a s a r e sul t o f th e

s

de co mp o siti o n o f sugar Th e acids m o st co mmo nly p o

. r

d uce d a re lactic ac e tic and butyric

, ,
He r e again t h e acid
.
, ,
 PR O D UC T S OF ME T AB OL I S M 17
5

p ro d uctio n s e rve s a s a m ans o f di ffe r e ntiating cl o se ly all ie d

e

bacte ria Th e me tho d O f de te cting a cid pr ductio n U ually

. o s

u se d is to g ro w th e ge rm s in milk co ntaining l itmus o r in ,

s ugar m e dia to w hi ch lit m us is adde d I n this th e f o rmatio n

.

of cid i s i ndi cat e d by th e cha ng e o f th e blu e litmus to a re d

a

co l o r M any bact e ria pr o duce an al k l i rath e r than an acid

. a .

Thi s is de te cte d in th e same way as th e acid S o me bacte ria .

a e abl e to p r o duc e
r ithe r an acid o r an alkali de p e ndi ng
e ,

up o n th e natu r e o f th e i r f o o d mate rial Wh e n th e B act . .

diphthe ri a fo r e xampl e is gro wn in a cultur e m e di um co n

, ,

tai ning sugar (and all m e di a made fro m m e at co ntain a small

am o unt unl e ss sp e cial me ans fo r r e mo vi ng it have b e e n e m
pl o ye d) it pro duce s fir st an acid r e actio n and th e n whe n th e
, ,

suga r is us e d up it p ro duce s an alkaline re actio n in th e cultur e

m e di um I t has b e e n fo und that whe n thi s ge r m is p ro
.

d ucing acid it d o e s no t f o rm i ts t o xi n r e adily but wh e n th e ,

suga r is r e m o ve d the n th e to xin pro ducti o n go e s o n much

,

m o re r e adily so that it b e co me s a matte r o f a gre at de al o f

,

practical imp o r tance in th e p ro ductio n o f th e to in to b e x ,

us d f r th manufactur e o f antito xin to have th e culture

e o e ,

m e dium fre e f ro m sugar to sta r t with s that th e ge rm may o

b e gi n its to xin p ro ducti o n imm e diate ly A numb e r O f by .

p r o ducts o f a go o d de al o f imp o r tan ce co ntain nitro ge n .

Am o ng th e simpl e r O f th e ni t ge no us su bs tance s a re indo l

ro

a nd S kat o l I nd l is v e ry co mm o nly p r duc e d by bacte ria

. o o ,

a nd si nce it ca n b
,
d e t e ct e d by a co l r r e a cti o n it is a pr o d
e o ,

uct o f a g o o d d e al o f imp o rtanc e in th e di ff e r e nti ati o n o f

17 6 GENERA L PHYS I OLOGY
micr o o rganisms Wh e n ind l in th e p r e se nce o f a nitrite
. o

is tr e ate d with sulphuric a cid th e re is d ve l o p e d a pink c o l o r

e .

I n o r d e r to d e t e rmin e th e n wh e th e r o no t a pa r ticul ar cul

r

tur e h as i ndo l pr e se nt o r no t it is o nl y n e ce ssary to add a so lu

ti o n o f nitrite a nd s o me sulphuric acid I t wa s o riginally .

supp o s e d that indo l was pro duce d by putre factive bact e ria
al o n e but no w it is kn o wn that it is pro duce d by many o th e r
,

bact e ria S o me tim e s it is p ro duce d in v e ry small amo unt s

.
,

and at o th e r time s m o r e abundantly S o m e tim e s also ce r .

tain bacte ria pro duce in addi tio n to th e indo l a nitrite I n .

thi s ca se it is o nly n e ce ssary t add th e sulphuric acid in o r de r

o

to ge t th e charact e ri s tic r e acti o n Thi s r e actio n is kno wn as

.

th e cho l e a re d re acti o n and was at fi r s t supp o s e d to b e char

r ,

a cte ri stic o f th e M sp c o mma o f Asiatic ch o l e ra but it i s no w

.
,

kn o wn that a co nside rabl e n umb e r o f bact e ria giv e thi s

ch o l e ra re d r e actio n a nd in m aki ng a d e te r mi natio n o f th e
,

pr e s e nce o f indo l sulphu ric acid sh o uld b e add d fi r s t a nd th e

,
e

nitrite afte rwards in o r de r to diffe r e ntiat e b e tw e e n th e simpl e

pr o ductio n o f indo l and th e pr o ductio n O f th e ni t r o so indo l -
.

O th e r nitr o g e n b y pr o duct s o f g r e at imp o rta nc e pro duc e d

-

by bact e ria a e th e e nzym e s The se a re substanc e s that

r .

stand midway b e tw e n living and n o nliving matt e r

e .

B a cte ri al Enzym e s and th e ir M e ch a ni s m o f Acti o n .

Up to th e pr e s e nt tim e no e nzym e has b e e n i so lat e d in a pur e

state C o n se que ntly n o thi ng de fi nite is kn o wn in r e gard to
.
,

th e ch e m i s try Of th e se c o mp o u nd s E nzym e s a re r e co g niz e d

.

by th e vario u s ch e mi cal change s which th e y pr o duce Th ey .

 PR O D UC T S OF ME T A B OL I SM 1
77

a re pro bably sp e cific sub s tance s Diff e re nt e nzyme s hav e .

b e e n n o te d which are abl e to pro duc e f e r m e ntatio n s o f

s a cchar o se ( inv e rti ng e nzym e s ) o f d e xtr o se lact o se c e llul o se
, ,
.

( amyl o lytic e n zym e s) pr o ducin

,
g e v e ntually ca r b o n di o xid e

and wate r ; whi ch c o agulate milk simi lar to r e nn e t ( r e nn e t

e nzym e s) ; whi ch d ig s t p r o t e in pr o duci ng ami no acids
e ,
e tc -

,
.
,

(pr o te o lytic e nzym e s) ; and that split fat s int o fatty acids
a nd glyc e rin e ( st e at o lytic e nzym e s ) .

C o ll o idal s o luti o n s o f m e tal s such as platinum S h o w

, ,

many similaritie s in actio n to kno wn e nzym e s The se co l .

lo ids can b e made to pro duc e chang e s in pro t e in s almo st

ide ntical with tho se pro duce d by tru e e nzyme s Th e y a e . r

also aff e cte d by ch e mical change s and by t e mp e ratur e in th e

same way as e nz ym e s .

Th e r e a e many writ e rs wh o ho ld th e vi e w that all so call e d

r -

e nzym e a cti n S d ue to phy si cal cha ng e s p r o duc e d in th e

o I

vario u s o rganic substance s and no t to any de finite ch e mical

,

re acti o ns Th e magne tism in th e magn e tic bar is in many

.

ways similar to an e nzym e Wh e n h e at e d magne tism and

.

e nzym e s di s app e ar wh e n tr e ate d with acids and alkali e s

,

th e y di sapp e ar and n e ith e r can b e i so late d in a pur e state

,
.

Ar th us thi nks that e nzym e s a re no t f o rms o f matt e r but that ,

th e y a e fo rms o f e n e rgy
r .

E nzym e s p o sse s s th e p o w e r o f pr o duci ng ch e mical chang e s

witho ut e nte ring to any g re at e xte nt into th e r e actio n Th e y .

can pr o du ce a n o xidatio n o f a sugar co mp o und into ca rb o n

di o xide and wate r and can br e ak do wn p ro te in co mp o unds
,

N
 7 8 GENERA L P HYSIOLOGY
into ca rb o n dio xide wat e r and amin o acids Th e se change s
, ,
-
.

can al so b e b ro ught ab o ut by s t ro ng r e age nt s ( acids and alka

li e s) o r by sup e rhe ate d ste am und e r pre ssure but no t with
, ,

n e a rly S O g r e at e as e as by e nzym e s B act e rial e nzym e s are .

m o st active wh e n th e y co m e dire ctly fro m living bacte rial

ce ll s Th e filtrat e s o f cultur e s o f bact e ria a nd de ad bacte ria
.

p o ss e ss ve ry littl e e nzym e co mpare d with th e live ce lls .

Th e int r a c e llula r e nzym e s a e di vid e d i nt o two g e n e ral

-
r

class e s : o xidi zing o hydrati ng and r e ducing o r de hydrating

r , ,

e nzym e s Oxidati o n is acc o mpli sh e d by m e an s o f catalyti c

.

age nts e nzyme s within th e bact e rial ce ll One o f th e m o st

, , .

studi e d o f th e o xidizing intra c e llul ar e nzym e s is catala se

-
.

Thi s e nzym e as w e ll a s m o s t o f th e o th e r s h as th e p o w e r o f
, ,

de co mp o sing hydro ge n dio xide and lib e rati ng na sce nt o y x

g e n
. M a ny bact e rial culture s co ntain sub stance s which ca n
d e co mp o se hydro ge n di o xide ( B act tub e rcul o sis B act .
,
.

di phth e riae M py o g e n e s var aur e us)

,
. C atalas e is di s tinct
. .

and se pa r ate fro m th e to xin .

Th e qu e sti o n o f th e o ccurr e nc e O f r e ducing e ny m e s Of z

bacte rial o rigin is s till an u ns e ttl e d pro bl e m I n all pro b .

ability th e y d o o ccur in so m e sp e ci e s .

I t was f o rm e rly supp o s e d by so m e inv e s tigat o rs that all

th e ch e m ical r e acti o ns w hi ch o ccu rr e d wh e n animal and v e ge

tabl e c e ll s w e re pre s e nt we r e d ue to th e ce lls the m se lve s a s

,

a who l e Th e re actio n s have since b e e n pro ve d to b e d ue to

.
(

th e d e fi ni te ch e mical s ubsta nc e s whi ch we no w kn o w as e u

zym e s and whi ch a re s e cr e te d by th e ce ll pro to plasm .

 P RO DUCT S OF ME TAB OLISM 1
79

E nzym e actio n is in no mate rial way di ff e re nt fro m o the r

ch e mical r e actio ns Th e e nzym e r e actio n s a re re ve r sibl e and
.

co nse qu e ntly th e r e is a te nde ncy fo r an e quilibrium to b e

e s tabli sh e d by b o th si d e s o f th e e quati o n Enzym e s act like
.

all catalytic age nts thus incre a sing th e sp e e d o f th e re actio n

, .

I n so m e cas e s it s e e ms that th e sp e e d o f th e r e acti o n i ncr e as e s

th e am o u nt o f e nzym e pr o duc e d . I t is no t kn o wn what th e

me tho ds o f catalysis in e nzyme s are I t has b e e n n o te d al so

.

th t e nzyme s hast e n th e synth e sis Of S o me co mp o unds Em

a
'
.

zym e s m ay p o ssibly fo rm lo o se ch e mi cal co mbinatio ns with

th e sub s ta nc e s o n w hi ch th e y act a nd with wat e r a nd th e r e sult
,

ing c o mp o und b e un stabl e a nd e asily bro k e n do wn and th e

wate r re main co nne cte d with so me O f th e co mp o n e nts o f th e
substance which was act e d up o n Th e re is s o m e e xp e im e n
. r

tal e vide nce to S ho w that e nzym e s may e nte r into lo o se

ch e mical c o mbinatio n with th e substance attack e d I t h as .

al o b e e n sugge st e d that th e e nzyme s may inc r e a se th e i o niza

s

tio n S alt so lutio n s favo r e nzyme actio n e vide ntly o n ac

.

co unt o f actio n o f i o ns .

T h e acti o n o f e nzym e s is sp e cific Th e y d iffe r fro m o rdi ‘

nary ch e m ical r e ag e nts in that th e y attack o nly sp e cific

o rgani c s ub s ta nc e s . F o e xampl e p r o t e o lytic e nz ym e s o nly
r ,

attack p ro t e ins and amylo lytic e nzym e s o nly ca rb o hydrat e

co mp o unds .

I n s m e particul ar s e nzym e s a re r e lat e d to bact e rial t o xin s

o .

Thi s phase o f th e subj e ct h as b e e n discusse d in an o th e r

chapte r .
 GENERAL PHY S I O L O GY
nzym e s fo rm c o llo idal so lutio ns which are o nly slightly
E

dialyzabl e On fil tratio n thro ugh p o rce lain 10 to 5 p e r c e nt

. 2

o f th e e nzym e s a re l o st du r ing e a ch fil trati o n Thi s o f co urse .

, ,
‘

ind ica te s th a t th e y are c o mp o se d o f m o l e cul e s o f larg e siz e .

Enzym e s with fe w e xc e pti o n s are m o r e r e si s tant to o ut sid e

, ,

age nts than th e c e ll s whi ch pro duce th e m Oft e n di lute .

so luti o ns o f ch e mical s will kill bact e r ia as b e f o r e stat e d b u t , ,

will no t d e stro y th e i ntra c e llular e nzym e s F o r malde hyde

-
.

g as ( H C O H ) h o w e v
,
e r in v e ry dilut
,
e s o luti o ns i s v e ry
d e structive to bact e ria and the ir e nzym e s Th e radi um rays .

will al so kill s o me e nzyme s .

Th e t e mp e ratu r e limits O f e nzym e acti o n vary b e tw e e n

35
°
C a
. nd 4 5 C no matt
°
e
. r fr
,
o m what S p e ci e s o f bact e ria th e y
a e d e riv e d
r Enzym e s lik e m o s t t o xin s a re d e stro y e d at
.
, ,

7 0
°
C .wh e n in aqu e o u s s o luti o n s Th e y a r e n o t a.ff e ct e d by
liquid air 19 o r by C wh e n in a d r i e d s tat e
° °
— 0 00 1 .
.

B act e rial e nzym e s in a m o ist c o n di ti o n may b e c o nv e rte d

into inactive co mp o unds ( e n ym o ids) wh e n subj e cte d to z

he at at co mparative ly l o w te mp e ratur e s S unl ight X rays .

,
-

radium rays ma rk e dly inhibit th e actio n o f all e nzyme s e xce pt

,

th e amyl o lytic wh e n th e y a re in aqu e o us s o luti o ns

, .

Enzym e acti o n is r e tard e d by th e accumulati o n o f bact e rial

me tab o lic pro ducts I n so me ca se s tho se m e tab o lic pro ducts

.

a re d e trim e ntal to th e e nzym e s th e m se lv e s This is tru e in .

ace tic acid f e rm e ntati o n wh e n alco h o l and ace ti c acid are pro
d uce d I t can b e s aid h o w e v e r that this i nhibiti o n is pr o b
.
, ,

ably d ue to th e e stablishme nt o f an e quilibrium .

 P RO D UCT S OF ME TAB O L ISM 18 1

Auto l yti c B a cte ri al Enzym e s . A larg e

numb e r o f bact e ria
pro duce pro te o lytic e nzyme s whi ch a re capabl e o f di ge sting
o th e r bact e rial c e ll s a nd al so capabl e o f dig e s ting th e m se lv e s
,

aut ly i ) zym e s o f t hi s cha r act e r a re call e d a u t lytic

( o s s. E n o .

I t h as b e e n n o t e d that in th e gro wth o f so m e sp e ci e s o f bac

te ria o n a r tifi cial m e di a that th e bacte ria inc r e ase in numb e rs

fo r a varying p e ri o d o f tim e th e n gradually d e cr e as e
,
This .

phe no me n o n o ccurs in s o me culture s o f M sp co mma This . .

fact can o nly b e acco unte d fo by asstI m ing in so m e case s that

r

th e pr o t e o lytic e nzym e s o f th e ch o l e r a bact e ria di g e st th e m

s e lve s ( auto lytic) and th e ir n e ighb o rs (is o lytic) .

Gro wth pro c e e ds whe n th e auto lytic e nzym e s are re mo ve d .

I n th o se di s e a s e s d ue to b ct ria whi ch pr o duc e e nd o to xins

a e

it is pro babl e that the s e to xi c pro ducts are lib e rate d fro m
th e bact e r ial ce ll by th e acti o n o f aut o lytic e nzym e s by l e uco ,

cytic e nzym e s a nd by th e p r o t e o lyti c e nzym e s

,
f ti ss ue ce ll s
o

in th e vi ci nity o f th e di se a s e d are a ( typh o id tub e rcul o sis , ,

dyse nte ry ch o l e r a
, ,
I t w o ul d s e e m fr o m what h as b e e n

l e arne d f ro m r e ce nt i nve stigati o n s that th e auto lytic e ny m e s z

are v e ry simila r to nucl e in in s t r uctur e Aut o lysis o f B ac

.

te rium tub e rcul o sis and alli e d f o r ms pr o gr e s se s v e ry sl o wly

I n tim e th e e nd o t o xi ns th e ms e lv e s a e di g e st e d by th e
r

pro te o lytic e nzym e s .

P s e ud o m o nas pyo cya ne a pr o duc e s a pr o t o lytic e nzym e e

which is h e t e ro lytic and furth e rmo re s e l e ctive in its actio n .

Th e e nzym e pr o duc e d is d e cid e dly antago nistic to a B act .

anthracis and B act diphth e ri a I t pre ve nts gro wth ab so

. .
18 2 GENERA L P HYSIOLOGY
l ute l y if it is pro duce d by a vig o ro us cultur e Thi s e nzym e .

can b e e asily iso late d I t is call e d pyo cyanas e

. .

I t is pr o babl e that auto lysis o f bact e ria tak e s place in th e ,

maj o rity o f cas e s wh e n th e bact e ria a re lacking in so m e

,

r e qui sit e fo o d pro duct I t is th e Opini o n Of so me inve s tiga

.

t o r s that n o rmally th e pro te o lytic ( auto lytic) e nzym e s are

, ,

h e ld in ch e ck by th e synthe si s o f th e pro te in fo o d mat e rials

( W e ll s)
. Th e r e is als o t h e p r o bab ility that a ntie n ym e s z

may b e g e n e r ate d .

Anti e nzym e s . E nzym e s l ik e o th e r c o mpl e x m e tab o lic

,

pr o ducts o f bacte ria pr o duce in th e b o dy fl uids o f immunize d

,

animal s antib o di e s Antie n ym e s fo r p r o te o lytic e nzym e s

,
. z

can b e pr o duce d with th e gre at e st e as e Antia m yl o l ytic .

and o th e r e nzym e s a re diffi cul t to pre pare Wh e n co m .

b ine d with th e sp e cific e nzym e s e ith e r i n oi o o o r i n oi t

,
th e ro ,

anti e nzym e r e nd e rs th e f o r m e r inactive Antie n ym e s can . z

b e pr o d uc e d by th e immu nizati o n O f a nimals with e nzym e s

which have b e e n subj e ct e d to h e at at co mparative ly l o w

t e mp e rature s (e n ym o id s) Th e a ntie n ym e s a re r e ally
z . z

d e str o y e d by a h e at o f 7 C 0
°
.

Th e m o st imp o rtant pr o duct s o f m e tab o li sm a re th e p o is o n

o us substanc e s which a re pr o duc e d by bact e r ia The s e sub .

stance s are kn o wn as th e pto ma ns to xins e ndo to xins andi , , ,

th e t o xic bact e rial pr o t e in s Th e se will b e di scusse d in d e tail

.

in Chapte r XXI I .
P A RT VI . B I O L O G Y O F S P E C I AL
I Z E D G RO U P S

C HAP T E R X VI

R O T O T R O P HI C
P B AC TERIA
0

Intro d ucti o n — Th e pro to t ro phi c bact e ria c o mprise tho s e

.

whi ch f ro m a phy sio l o gical standp o int a e th e mo st primi

, ,
r

tive Th e y o btain th e ir fo o d fro m ino rganic so urce s a nd are

.

abl e to build up th e se S impl e sub s tan ce s i nt o th e ir co mple x

pro to plasm I t s e e ms pro babl e that such o rgani sms as the se
.

must have b e e n th e o riginal fo rms o f life o n th e e arth and that ,

th e m e tatr o p h ic a nd paratr o phic hav e d e v e l o p e d fr o m th e m ,

a nd in a se ns e
,
th e se mus t b e co n side r e d as de g e ne rate s
, .

F r o m thi s sta ndp o int th e pr o t o tr o phic bact e ria o ccupy a

ve ry imp o rtant place in th e de ve l o pm e ntal hi sto ry o f life o n th e

e a r th M any Of th e m at th e pr e se nt tim e a s w e ll a s in th e
.
,

pa s t are o f i ne s timabl e se rvice to man Th e se pro to t ro phi c

, .

bacte r a nclude th e nitrifying th e iro n th e s ulphur and th e

i I , , ,

l e gume o r ni tro ge n fi ing bacte ria

,
-
x .

N i trifyi ng B acte ri a Th e nitrifyi ng bacte ria o xidize

.

ammo nia to th e nitrite s and th e nitrat e s The re are s e ve ral .

diff e re nt sp e cie s whi ch b e l o ng to two diff e re nt cla sse s : tho s e

whi ch carry o n th e first part o f th e pro ce ss that is o xidi ze , ,

183
1 84 B I O L O GY OF S PE CI AL I ZE D G R OUP S
ammo nia to nitrite s a re kno wn as th e ni t o so bacte ri a Th e
“

,
r -
.

o th e r class a re th e ni t ifyi ng bacte i a p r o p e r ( F ig

r r .

Th e ch e mi cal cha ng e s br o ught ab o ut by th e ni t r ifyi ng

bact e ria in th e o xidatio n Of amm o nia is r e pre se nt e d by th e

f o ll o wing f o rmula
2 N H3 + O O HN 02 + 2 H2 O .

And th e se co nd stage is r e pr e se nte d by th e

f o rmul a
2 HN02 2 O 2 HN Og
mm
.

FI G .

t ying ba t ria Th e e n e rgy Obtain e d by this o xidati o n p ro

c e .

( ) N itr
1 f o us er

m t ( ) itri
e n s, c
2
e s s e nabl
n e s th e
c
s e bact e ri a t o mak e u se o f
f rm t
th e ca r b o n and nitr o g e n i n an i no rganic f o r m
e e n s X 2 000 .

Aft r B rri e d u an

S t tz ru (Lip e a nd thus to synthe size o rganic matt e r .

m )an
The s e bact e ria no t o nly d o no t r e qui r e o r
.

g a n i c matt e r f o r f o o d but in pu r e cultur e at l e ast th e o r ga nic

, ,

mat e rial inte rfe re s with th e ir d e ve l o pm e nt Th e pro ce ss o f .

nitrificati o n is strictly an o xidizi ng p r o c e ss and in o r de r , ,

that th e pro ce ss may go o n an abundant o xyg e n supply ,

must b e availabl e F urthe rmo r e th e activity o f th e s e o rgan

.
,

ism s is ch e ck e d by th e pr e s e nc e o f co nsid e rabl e amo unts o f

th e acid s so that if th e pr o c e ss c o nti nu e s ba se s mu s t b e sup
, ,

plie d which will unite with th e acid s to f o rm salts Th e .

nitrificati o n Of s o il s is d e p e nd e nt up o n th e acti o n o f th e se
bacte ria and o ur kno wl e dge o f th e life hi sto ry o f th e se ge r ms
,

is a matt e r o f practical imp o rtance wh e n it is applie d to s o il

f e rtility C e rtain phase s o f thi s will b e r e f e rr e d to in ano th e r
.

co nn e cti o n Th e s e bacte ria a re wide ly di stribute d in th e

.
 PR O T O T R OP HIC B ACT ERI A 18 5

s o il and th e imp o rtan ce o f nitrifi catio n in N ature s fo o d

,
’

cycl e canno t b e o ve r e stimat d e .

N i tro ge n fi xi ng B te ri
-
N it r o g n fi xing bact e ria o r
ac a . e -

l e gume bact e ria o ccur in th e n o dul e s whi ch are fo un

,
d o n th e
ro ts o f vario u s l e gumi
o

n o us plants Gro wing .

in th e se no dul e s th s e e

ba cte ria a e able to fixr

th e nitr o g e n o f th e air

into a fo rm which is
availabl e fo r plant use ,

a nd whi l e th e y liv e in

th e pla nt th e y are no t

to b e r e gard e d a s pa r a
F — B a t r id fr m l g m
IG ( ) Fr m u
3 tCS 0 1 e e n a s m e t a
.
44 . . c e o s o e es . I o

M lil t alba ( 5 ) Fr m M di ag
‘

1 ,
v “

e o us . 2 , 3, o e c o

tr o phi c b acteria but ativa ( ) F r m V

s i ia vill Aft r.H 4 o c o so . e ar

ri d B arl w ( L ipm a )
so n a n o . n .

th e y b e l o ng to th e

g ro up unde r discussi o n in this chapt e r ; name ly th e pro to ,

tr o phi c bact e ria The y are ba il li S far a th i r fo rm is co n

. c O s e

c e rn e d and th e re are many di ffe re nt ki nds Wh e the r th e s e

,
.

diffe re nt ki nds are s e pa r ate sp e cie s o r me r e ly va ri e ti e s is a ,

que stio n th at is no t fully se ttl e d The se ge r m s a e no t st rictly

. r

p ro to tr phi c but may b e cultivat e d up o n a r tifi cial m e di a in

o ,

whi ch th e y make use o f o rganic fo o d mat e rial The r e a e . r

di ff e r e nt f rms ass o ciat e d with th e di ff e r e nt plant s

o One .

p e culiarity h o we ve r Of all fo rm s is th e te nde ncy at a

, , ,

ce rtain stage in the ir life histo ry to assum e irre gul ar o r ,

1 86 B I O L O GY OF S PE CI ALI ZE D GROUP S
bizarre S hap e s whi ch are usually kno wn a s bact e ro ids (Fig
, ‘
.

Th e se l e gum e bacte ria make the ir way i nto th e ro o ts

thr o ugh th e ro o t hairs and induce th e fo r matio n o f tub e r
cl e s o r n o dul e s I n th e se th e bact e ria live and rapidl y
,
.

multiply and it is h e r e that th e y take th e e l e m e ntal nit ro

,

g e n ioth e air a nd fix it i n a f o r m availabl e fo r pla nt use .

I t h as b e e n abunda ntly sh o wn that th e p o w e r o f th e s e

l e gumin o us plant s to make use Of th e ni tro ge n Of th e air

de p e nds o n th e activity o f the se micro o rgani sms I f th e se .

l e gum e s a e so wn in st e ril e so il and pro t e cte d f ro m in o cul a

r '

ti o n th e y will c e ase to gro w as so o n as th e nitro ge n Of th e

,

s e e d has di sapp e ar e d ; wh e r e a s o th e r plant s u nde r e xactly

,

th e s am e c o ndi ti o n e xc e pt that th e y hav e b e e n inf e ct e d with

,

pure cultur e s o r so il wa shi ngs co ntinue to gro w Th e se ,

.

l e gum e bact e ria are o f e xt r e me imp o rtance in agricultural

pursuit s and will b e r e f e r r e d to again in a lat e r chapt e r
,
.

Oth e r bact e r ia f o und in th e so il a e abl e to fix ni tro g e n wh e n

, ,
r

gro wing inde p e nd e ntly o f l e gume s o plants r .

S ul phur B acte ri a M a ny bact e ria (put r e factiv e ) are

.

abl e to pr o duce s ulphure tt e d hydr o ge n but th e sulphur bac ,

t e ria pro p e r tak e thi s hyd ro g e n sul phi de that h as b e e n p ro

d uce d by a no th e r g r o up a nd use it a s fo o d o xidi zing int o ,

sulphu ric acid ( Fig The se sulphur bacte ria b e l o ng to

.

two gro ups o ne o f whi ch i s c o l o rl e ss and th e o th e r c o ntains

,

a co l o ring matte r Thi o thrix may b e take n as an e xampl e o f

.

th e filam e nt o u s fo rm s o f th e c o l o rl e ss g r o up T h e Thi o th r ix .

is co mp o se d o f a thre ad o f c e lls which is usually attache d to

 PRO T O T R OPHI C B ACT ERI A 18 7

so me o bj e ct Th i s filame nt o f ce lls is surro und e d by a thin

.

S h e ath . R e pr o ductio n tak e s plac e by m e a ns o f g o nidia .

Th e g o nidi a a e f o rm e d fr o m th e t e r m inal c e lls Of th e filam e nt

r

a nd a e lib e r at e d in th e u s ual way

r C e rtain no nfil a
. m e nto us

fo rms such as B e ggiato a are w e ll

, ,

kno wn B o th o f th e s e f o rm s make
.

us e o f hydr o g e n sul phid e as a f o o d ,

o xidizing it to wat e r and e l e m e ntal

sulphur The se sulphur g r anul e s are

.

sto r e d in th e ce ll itse lf Th e se bac .

t e ria d o no t gro w in highl y nutritio us

cul tu re me di a but a re abl e to o btain
,

th e i r f o o d fro m r e lative ly S impl e sub

sta nce s by m e ans O f th e e n e rgy which
is Obtaine d by th e o xidati o n o f th e

s ulphur e tt e d hydr o g e n The i carb o n

. r

may b e Obtaine d fro m th e air o r F 4 5 — S lph r ba t ria IG . . u u c e .

( ) Th i p i il l m Wi
f ro m small trace s o f f o rm
r u no
ic o r pro p ri g d kii ( ) B ggiat a I

ra s
os

. 2 e o

o ni e acids N itro g e n may b e Oh Aft r C e 01

”

( L pm )
.

m s i . i an .

taine d fr o m th e ai r o f r o m amm o nia

r .

I n th e synth izing o f o r gani c mat e rial th e y w o r k ind e p e n

es

de nt Oi th e rays o f sunlight .

An th e r g r o up o f th e se s ulphur bact e ria a re c o l o r e d purpl e

o

by a co l o ring matte r kn o wn as bact e ri o purpu rin whi ch acts ,

in a way S i m ila r to th e ch l o r o phyll o f gr e e n plants The se .

bacte ria a e la rge micro co cci whi ch have th e co l o ring matte r

r

e v e nl y di s tribut e d t h r o ugh o ut th e p r o t o plasm Th e y have .

 88 B I O L O GY OE S PE CI AL I ZE D GR O UP S
a d o ubl e so urce o f e n e rgy that wh ich is o btaine d by th e use
,

o f sulphu r e tt e d hyd r o g e n in th e g r o up d e scrib ed ab o v e

,
as
,

and also by m e an s o f th e ba ct e ri o purpuri n whi ch e nable s

the m to synth e siz e o rganic matt e r with th e e n e rgy Obtain e d
fro m th e sun s rays ’
.

I r o n B a cte ia rTh e ir o n bact e ria are r e pr e se nt e d by th e

.

o rgani s m C r e no th r ix p o ly sp o ra whi ch was d e sc rib e d by F

, .

C o hn in 187 0 Thi s bacte rium c o n si st s o f a fil ame nt o f c e l ls

.

attach e d to s o m e o bj e ct Th e c e ll s in thi s filam e nt are smalle r

.

at th e bas e and incr e as e in di am e te r high e r up Th e r ange in .

S iz e is fr o m o ne and a half to fi e mic r o ns Th e se bact e ria

v .

o ccur i n tuft s and e ach filam e nt i s s u r r o und e d by a sh e ath .

Th e c e ll s at th e o ut e r e nd o f th e filam e nt b r e ak up i nt o

g o nidia As o rdi n
. a rily se e n th e se bact e ria have th e ir sh e ths
,
a

p e rm e at e d by r e ddi h b ro wn ma sse s o f fe rric Oxid e Th se

s . e

bacte ria tak e f e rri c ca rb o nate a nd o xidiz e it to f e rric hydr o xid e ,

as r e pr e s e nte d in th e fo ll o wing f r mul a : o

2 F e CO3 3 HQO O F e 2:( OH) 6 2 C02 .

AS a r e sult o f thi xidatio n p ro c e s s e ne rgy is Obtain e d whi ch

s o ,

e nabl e s th e s e bact e ria to synth e s i e o r ganic mat e r ial The y

z .

O btain th e ir nitr o g e n fr o m i no rganic s o urc e s and a re no t d e

p e nd e nt up o n o rganic matt e r fo r supp o rt Th e se bact e ria .

gro w r e adil y in wat e rs fr e e f r o m o rganic matt e r pro viding th e

ir o n salt s are pr e se nt Th e y a e wid e ly di st ribut d in natu re
. r e

and are f o und in swamp s m e ado ws and marshe s and a re

, , ,

e sp e cially lik e ly to g r o w i n d e e p w e ll wat e r s whi ch a re ir o n

b e aring S o m e tim e s th e ir gro wth is such as to r e nde r th e

.
 PROT O T R OP HI C B ACTERI A 1 89

wate r unde sirabl e fi r st fo r laundry purp o se s and lat e r fo r

,

drinki ng purp o s e s altho ugh th e o nly Obj e ctio n he r e is an

,

a sth e ti c o ne . Th e ir g r o wth in th e wat e r pip e s o f a city

so me time s o ccurs to su ch an e xt e nt that th e fl o w o f w at e r is
i nt e r f e r e d with Wate r calamiti e s o f this so r t have b e e n e
. x

p e ri e n ce d by a numb e r o f citi e,
s pa r ticul a r ly L ill e and B e rlin .

“
The re is r e aso n to suppo se that th e mass e s Of i ro n o re in
no rthe rn M inne so ta up o n th e M e saba range w e re de p o site d
the re by th e activity o f i ro n bact e ria liv ing in th e wa r m wate rs
o f a n a nci e nt o c e a n We may imagin e such a prim e val se a
.
,

h o t lik e th e g e y se r s O f th e Y e ll o w s t o ne its wat e rs impr e gnat e d

,

with iro n and furni shi ng a spl e ndid fi e ld fo th e p e cul iar ac

r

ti i ti e s O f th e i r o n bact e ria
v . L ivi ng in such a n o c e a n fo th o u
r

sands o i ye a r s as th e y hav e do ne th e re is no thing unre a so nabl e

, ,

in att ributi ng to th e m th e de p o sits o f ir o n o e whi ch du ring

r

th e se latte r day s a e b e i ng min e d fo r co mme rcial purp o se s

r .

(M ac M illa n ) .

An o th e r r e pr e s e ntative o f thi s gr o up is C lad o thrix Od o rife ra ,

which is co mmo nly fo und in th e so il a nd whi ch give s th e p e cul

iar O do r to th e so il such a s that whi ch is particularly n o tice d
aft e r a rain sto rm .
 CHAP T ER X VI I
C HR O M O GENI C B ACTE R I A

In tro d u cti o n S o m e sp e ci e s o f bact e ria wh e n gro wing in

.

culture s e xhibit vario us co l o r s B acte ria o f thi s charact e r are

.

call e d ch o m o ge ni c r .

Th e maj o rity o f bact e ria pr o duc e no c o l o r what e v e r and

app e ar white o r co l o rl e ss wh e n gro wing in culture s Th e y a re .

call e d no n ch o mo ge ni c -
r .

R an ge o f C o l o r T h e chr o m o g e nic bact e ria p r e s e nt i n th e

.
,

vari o us sp e ci e s all th e co l o rs o f th e so lar sp e ctrum Th e

,
.

f o ll o wi ng tabl e gi ve s th e co l o rs o f th e sp e ct r um and an e xampl e

o f a chr o m o g e nic ba ct e rium which will n o rmally pr o duc e p rac

tically th e co rr e sp o nding co l o r in c ultur e s .

Re d B a ill p d igi
c us roFl gg o s us u e

O ra ng e S ar i na a ra ntia a Fl ugg
c u c e

Y ll w
e o Mi r
c o co ccus fl F l ii gg a v us e

Gr n ee B a t ri m viridi K r
c e u s use

Bl ue P d m na yn y
se u o o M ig la
s s c a ne a u

I ndig o P d m na i ndig f ra V g
se u o o s o e o es

Vi l t
o e B a ill vi la
c us J rda n
o ce o us o

The r e are als o sp e cie s o f bact e ria whi ch pr o duce all th e

S had e s b e tw e e n th e vari o u s c o l o rs o f th e sp e ctrum .

Th e c o l o r O f ch r o m o g e nic bact e ria i s d ue to pigm e nt

granule s Th e p o siti o n o f th e pigm e nt gran ul e s in r e latio nto

.

19°
 CH R OM OGENI C B ACTERI A 19 1

th e ce ll is imp o rtant I t h as b e e n f o und that th e re is a gre at

.

variatio n amo ng diff e r e nt sp e cie s in this particular .

i
Lo ca t o n o f Pi gm e nts I n th e cultur e s o f s o m e sp e ci e s o f
. .

bacte ria th e pigme nt granul e s can b e s e e n o utside th e bacte ria ,

b e twe e n th e c e ll s I n cultur e s o f B acillus p ro d igi o sus P se udo

.
,

m o nas indi go fe r a and th e large maj o rity Of th e ch r o mo g e nic

,

bacte ria this co n di tio n I S n o te d Th o s e bacte ria whi ch e x

.

cre te the ir pigme nt are calle d ch mo p a o us ro r .

The r e a re o the r chr o mo ge nic bacte ria whi ch d o no t e xcr e t e

th e pigm e nt gra nul e s but r e tain th e m withi n th e c e ll pr o to

plasm The se bact e ria are alle d chro m at pho o ns Th e

. c o r .

pigme nt granul e s are fo und to b e m o st abundant in th e o ute r

laye r s o f th e c e ll pro to pla sm A large numb e r o f Sp e cie s
.

amo ng th e high e r bacte ria as fo r e xampl e in th e g e nu s Thi o cys

, , ,

tis am o ng th e B e ggia to ace a b e l o ng to thi s class o f chr o m o

,

ge nic o rgani sms P igme nte d p r o t o plasm o f thi s cha r acte r is

.

al so n o t e d in tho se sp e ci e s O f ba cte ria (B act ch l o rinum ) (B acil

.

l uS vi r e ns ,
which a e clo s e ly r e late d to th e Cyano phyta
r ,

( blu e-
gr e e n alg a ) a n d in whi ch c o l o ring matt e r S imilar to
,

chl o ro phyll is fo und .

The r e is still an o th e r cla ss o f chro mo ge nic bact e ria in which ,

by th e aid Of th e mic ro s co p e th e pigm e nt granule s can b e s e e n

,

to b e d e finit e ly c o ntain e d withi n th e c e ll wall C h r o m o g nic . e

bacte ria o f thi s class are call e d pa ach m t pk ns B acillus r ro a o o ro .

vi o lace o us is a typical e xampl e o f a bacte rium b e lo nging to

this class .

P igme nt pr o ducti o n in th e maj o rity o f bact e ria is o f p racti

19 2 B I OL O GY OF S PE C I AL I ZE D G R OUP S
cally no physio l o gical imp o rtance Th e pigme nt granul e s .

a re u s ually e xcr e t e d as rapidly a s th e y a re f o r m e d I n o nly .

th o se cas e s wh e re th e pigm e nt is co mbin e d with th e p ro to

pla sm o f th e c e ll ( ch ro mato ph o ro us) is it Of any imp o rtanc e .

I n this ca se it u nd o ubt e dly se rv e s th e functi o ns O f th e chl o r o

phyll in th e high e r plants I t can th e r e fo r e ab so rb carb o n

.
, ,

di o xid e ( C0 ) and give O ff o xyg e n (O) in th e pr e se n ce o f s un

2

light This fact has b e n d e mo nst r at e d in th e case Of th e

. e

bacte rio purpurin o f th e sulphur bact e ria I n all chr o m o .

par o u s bact e ria th e pigme nt granul e s can o nly b e r e gard e d

,

as e xcre to ry pr o ducts Of th e c e ll N O co nne cti n with th e

. o

nutritive pro ce s se s o f th e c e ll can b e de mo nst r ate d C hro mo .

g e n e si s se e m s to b e an une ss e ntial pro ce ss .

I n r e gard to th e c o nditi o ns und e r whi ch pigm e nt f o rmati o n

take s place it may b e said in th e b e ginning that pigm e nt fo r

matio n rare ly tak e s place in th e ab se nce O f o xyg e n I n cul .

tu re s it is n o t e d that o nly th e s up e rficial lay e rs o f bacte ria

pro duc e any pigm e nt tho se b e n e ath b e ing co l o rl e ss Oxyg e n
,
.

s e e ms to b e a p re r e qui sit e in m o st case s Thi s p o int will b e

.

again r e f e rr e d to P igme nt f o r matio n in m o s t S p e ci e s o f

.

bacte ria tak e s place e qually w e ll in th e light and in th e da rk .

I n a fe w sp e ci e s pigm e nt f o rmati o n is i nhibit e d by da r k ne s s

( ch ro mat o ph o r o u s) B a cillu s myc o id e s var r o se us pro duc e s

. .

pigm e nt o nly in th e da rk High t e mp e ratu re s usually inhibit

.

pigm e nt f o rmatio n ( 3 7 C and ab o ve ) C o l o r e d lights have

°
. .

no e ff e ct o n chr o m o g e ne sis .

S o m e pigm e nt s a re cl o s e ly r e lat e d ch e mically to th e lip o

 R
C H OM O GENI C B AC TERI A 19
3

chr o m e s and o th e r s to th e pto mains Th e r e actio n and .

i
ch e mical co nstitutio n o f th e m e di a o n wh ch bacte ria are gro w
ing mark e dly influe nc s chro mo ge n e si s in so m e case s Acids
e
‘

and alkalie s usually pro duce so m e chang e in th e pigme nt whe n

the y are appli e d to it O ccasio nally th e co l o r may b e d e
.

stro y e d change d to a di ff e r e nt co lo r o r to a co l o r which

, ,

will r e tu rn to th e o riginal wh e n th e r e qui site re actio n is re

sto r e d E F S mi th and o th e rs give th e fo ll o wing e xampl e s
. . .

Th e y e ll o w pigm e nt o f B act e rium ca m p e st e r e mains unc

r

change d and B acillu s p o di gio sus change s fro m re d to carm in e

r

in c e rtai n a cid s o l u ti o ns a nd y e ll o wi sh bro wn in c e r tain alkali e s .

Th e blu e c l o r o f P se udo m o na s syncya ne a is p ro du ce d o nly in

o

acid milk Th e blue g re e n flu o r e sce nt co lo r O fte n se e n in so me

.
-

chr o mo ge nic bacte ria is p r o duce d o nly in alkalin e m e dia .

S o me chr o m o ge nic bact e ria pr o duc e alkali e s ( N H 3,

Thi s de p e nd s o n th e supply o f nitr o g e n and ca rb o n I n .

ge latin P se udo m o nas pyo cya ne a do e s no t f o rm alkali but in ,

ammo nium suc cinate it do e s .

I n thi s la s t m e nti ne d g r o up o f blu e gr e e n fluo r e sc e nt o rgan

-
o -

i sms O f which P se ud mo na s fl uo re sce ns is a typical e xample

,
o ,

it is p r o babl e that th e re are two di s ti nct pigme nts pr o duce d .

J o rda n h a s studi e d th e se pigm e nt s car e fully and s tat e s that

the re is pr e se nt a blue pigme nt call e d pyo cyani n which is

, ,

vi sibl e by ga slight and so lubl e in chl o ro fo r m and a g re e n ,

fluo re sce nt pigm e nt call e d fl uo re scin whi ch is y e llo wi sh

, ,

in gaslight and inso luble in alco ho l and chl o ro fo rm B ac .

t e ria r e adily lo se th e ir p o we r to pro duce pyo cyanin .

o
1 94 B I OL O GY OF S P E CI AL I ZE D GR OUP S
B acte rial pigm e nts are se cr e t e d as co l o rl e ss co mpo unds ,

whi ch in th e maj o rity Of case s are waste p ro ducts o f th e

, ,

bact e rial ce ll Th e se co mpo unds are o f th e natur e o f l e uco

.

base s whi ch assum e co lo r o nly whe n o xidize d by th e air Th e

,
.

y e ll o w and bla ck pigm e nts are distinct o xidati o n pro ducts .

Ch ro m o g e nic bact e ria ca n b e gr o wn und e r c e rtai n co nditi o n s ,

a s fo r e xampl e
,
at hi gh te mpe ratur e s and no pigm e nt will
, ,

b e pro duc e d No n chro m o g e nic fo rm s can be pro duc e d fro m

.
-

o rdi naril y chro m o g e nic f o rm s e tc Th e r e can be no do ubt

, .

that pigm e nt fo rmati o n is an une sse ntial m e tabo l ic pro c e ss to

th e bact e rial c e ll e xc e pt a s m e nti o n e d b e f o r e in th e chr o mat o
, ,

p h o ro us bact e ria V e ry
. fe w chr o m o g e nic bact e ria a re pa th o

g e ni c to a nimal s P se udo m o nas p yo cyane a and th e vario us

.

pyo ge nic mi cro co cci are amo ng th e co mm o n fo rms Chro .

m o ge ni c bact e ria am o ng th e pla nt path o g e ns are no t un

co mmo n .

M igul a h a s arrang e d th e chro m o g e nic bacte ria into thr e e

classe s as fo llo ws Tho se pro d ucing
,
.

( I ) P igm e nts so lubl e in wat e r .

( ) P igm e nts inso lubl e in wate r so lubl e in alc o ho l

2 ,
.

(3 ) P igm e nt s in so lubl e in wat e r and alc o h o l .

P i gm e nts S o l ub l e in W a te r —
Ch ro m o g e ni c bact e ria whi ch
.

pro duce fluo re sc e nt pigme nts whi ch a re so lubl e in th e culture

,

m e dia and diff use thr o ugh it be lo ng to this class P se udo

,
.

m o nas p yo cyane a P se udo m o nas fl uo re sce ns and P se ud o m o

, ,

nas syncyane a a re typi cal o rganisms o f this gro up P se udo .

mo nas p yo cya ne a and P se ud o mo nas fl uo re sce ns re fe rr e d to ,

 R
CH OMO GE NI C BACTERI A 19 5

b
a o ve ,
pro duce pigme n ts ; name ly pyo cyanin and
two ,

fl uo re scin Le d d e rh o se a n alyze d pyo cyanin and f o und it

.

to b e simi l ar in s tructure to so m e o f th e kn o wn pt o mains

( C H
u u N O ) z I t se.e ms to b e d ri v e d f ro m th e a ro matics and

clo se ly re late d to th e anth racine s Pyo cy a nin is no n to xic to

.
-

a nimals .

Whe n tre a te d with acids fl uo re scin be co me s co lo rl e ss

,

and pyo cyanin t urns re d S o lubl e pho s phate s and sulphat e s

.

se e m to be n e ce ssa ry fo r th e p ro ducti o n o f fl uo re scin but no t

pyo cyanin P yo cya nin can be p ro duce d in me d ia d e vo id O f

.

B o th pyo cyanin and fi uo re s cin a re so luble in wate r Pyo .

cyani n is also so lubl e in chlo ro fo rm but inso luble in al co ho l

and fl uo re scin is inso luble in bo th al co ho l and ch lo ro fo rm .

Pi gm e n ts I n s o l ub l e in Wa t e r S o l u b l e in Al co h o l
,
— Th e .

M i cro co ccus p yo ge n e s va rs aure us citre us fl av us and

.
, , ,

B a cillus p ro d igi o us are typi ca l e xampl e s O f chro mo ge nic ba c

s

It is claime d by th o se inv e s tiga to rs wh o ha ve s tudie d th e

p igme nt gra nule s Of this clas s that the y a re fat—like in nature
and simila r to lipo chro m e s The y a re so luble in all fat so l e n ts
. v ,

fo rm de ndritic crysta ls ca n be sapo nifi e d and o n d e co mpo si

, ,

tio n giv e th e a cro l e in re a cti o n ( C I LO) and fat Od o rs Wh e n

S .

co nce n tra t e d sulphuric a cid is add e d th e lipo cya ni n re a ctio n

,

o ccu rs and th e p igm e nt gra nul e s a re ch a ng e d to blu e .

P igm e n ts I ns o l ub l e i n Wa t e r an d Al co h o l —Ve ry fe w .
196 B I OL O GY OE S PE CI AL I ZE D GR O UP S
M ic r o c o ccus va r fl a us a s a typic l e xampl e Th e
ci tre us . v a .

ch e mical stru ct u r e o f th e s e i nso lubl e pigm e nts i s unk no wn .

T h e pigm e nt s O f chr o m o g e nic bact e ria hav e al o b e e n s

cl ssifi e d a s di ffusibl e ( E P S pyo cyane a) and no ndiff u sibl e

a x . .

( E B p ro digio us) Thi s clas sificatio n is o f no practical

x . . s .

value .

S o m e sp e ci e s o f chro m o g e nic bact e ria whi ch attack plants

p ro duc e a b ro wn stain o n th e plant at th e p o int o f i no cul atio n

( B act camp e str e (y e ll o w) o n s t m s and l e ave s o f cucumb e r s
. e

a nd t o mat o e s ) whil e o th e r no n ch r o m o g e nic f o rm s pr o duc e

,
-

th e sam e e ff e ct o n th e plant ( B act e rium so l ance arum o n st e ms

and l e ave s o f cucumb e rs and to mato e s) P lants unde r va ri .

,

o us c o ndi ti o n s sh o w s tai ns w h ich ca nn o t b e a

,
tt ribute d to any
ch r o m o g e nic bact e rium o r o th e r m ic ro o rgani sm Th e se stains .

a re o xidati o n pr o duct s F o r e xampl e wh e n B act e rium vas

.
,

cul a um
r atta ck s sugar cane ve ry o ft e n th e plant S ho ws a
re d s tai n I t h as b e e n n o t e d h o w e v e r that fu ngus gr o wths
.
, ,

a nd i nse ct sti ng s will p r o du ce th e sam e c o l o r This co l ring . o

o f plants has b e e n n o t e d in ma ny sp e ci e s a nd und e r va r i o u s

c o ndi tio ns I t is fre que ntly no t d in gr e e n frui ts and flo w e rs

. e .

Th e s ubj e ct O f th e chr o m o g n c bact e rial pigm e nt s fr o m th e e i

vario us S p e ci e s O f ba cte ria h as no t b e e n e xhaustive ly stu di e d

as ye t M o st o f th e inv e stigati o ns so far hav e b e e n o n th e
.

blue gre e n and fluo re s ce nt pigme nts pyo cyani n and fl uo re scin
-

whi ch we r e re fe rre d to ab o ve .

H i s to i c l Al l us i o ns
r
— Th e re d chr o m o g e nic ba ct e rium
a .

kno wn as B acillus pro d igio sus has a ve ry inte r e sting hi sto ry .

 CH R OM O GENI C B AC T ERI A 19 7

I n th e C hu r ch f S t G e na r in N apl e s twice a y e ar a crytsal

o . o

vial co ntai ning a so lid substance o f a da r k co l o r was e xp o se d in

th e e a r ly d y s und e r a gla ss ca s e to th gaz e o f th e p e o pl e
a e .

This was de clar d by th e p ri sts to b e th e blo o d o f

e e

whi ch co uld b e change d f ro m a s o lid to a li q uid in o ne night .

Wh e n G e ne r al C hampio ne t co mmanding th e F r e nch e nte re d

, ,

N apl e s in 7 9 9 th e m i r a cl e o f th e liqu e fa cti o n O f th e bl o o d wa s

1 ,

no t p e rf o r m e d and th e r e was gr e at dis s e nsi o n am o ng th e

p e o pl e Th e mo nk s d e clar e d that th e lique factio n was th e

.

w o rk O f Go d and that the y co uld ho t bri ng it ab o ut Ho w

,
.

ev r
e ,
G e n e ral C hampio n e t info rm e d the m that th e miracl e
mu st tak e plac e at a d e fi nit e tim e und e r p e nalty o f de ath .

Th e miracl e t o o k plac e as d e si r e d a nd it was f o u nd that th e

,

m nk s p o s se sse d a sub stanc e which th e y ke pt fr o m y e ar to y e ar

o

which wh e n adde d to g e lati no u s mat e rial w o uld bring ab o ut

a liq ue factio n We no w kn w that unkno wn to the m th e
. o , ,

m nk s had a cultu r e f B acillus p ro digio sus whi ch pro duce s

o o ,

a pro t e o lytic e nzym capabl e o f lique fyi ng g e latin Th e same

e .

bact e rium h as b e e n no te d o n bre ad in damp chu rche s o n Oil ,

painti ngs e tc and was inte rpre te d to b e th e bl o o d Of Christ

,
.
,
.
 CHAP T ER XVI I I

PH OTO GE NI C B ACTER I A

Intro d ucti o n . Th e re are ce rtain sp e ci e s o f sapro phytic

bacte ria which p o ss e ss th e p o we r o f thro wing Off light wh e n th e
r e quisit e co nditi o ns are pro vide d B acte ria Of this charact e r
.

a re call e d p ho t ge ni c
o .

Th e p o w e r o f lumin o sity o r p h Osph o re s ce nce is d ue in all

, ,

pro bability to th e co mbu stio n o f vario us f o o d mat e rial s and

to c e ll r e spirati o n just a s th e ph o sph o r e sc e nc e o f in se ct s is a
,

c o mbustio n pro ce ss Lumin o sity its e lf is o f no phy si o l o gical

.

imp o rtance .

Di s tri b uti o n
. Th e maj o r ity O f ph o to g e nic bact e ria a re
-

f o und in salt wate r a nd o n th e fre h a nd d e co mp o sing fle sh o f

s

ma rin e animal s such as fi h Th e ph o sph o r e sce nce Of th e se a

,
s .

is d ue in a larg e part to bact e ria a nd al so to ph o sph o r e sce nt

pro to zo a.

I t has b e e n kn o wn fo r so m e tim e that th e light O ft e n n o t e d

e manati ng fr o m d e c o mp o sing will o ws and o th e r tr e e s and ,

pa r ti cularly fr o m de caying s tump s o f t re e s is d ue to a fungus ,

.

This fungus 18 I n so me case s S imilar to a m o ld and in o th e r ,

cas e s pho to ge nic bacte ria hav b e e n fo und Th e much re

e .

no wne d wil l O th e wi p p h e n m no n i s cau se d by sapr o g e nic

“ ’ ”
-
s o e

19 8
 P HOT O GENI C B ACT ERI A 99

micro o rganism s ( no n pho to ge nic) which ge n e rate infl am m a

-

b l e ga se s such as ph o sph o r e tt e d hydro g e n and m e than e

, .

S o m e v e g e tabl e s such as p o tat o e s b e ans e tc o n d e co m p o

, , ,
.
,

sitio u may give o ff li ght and S ho w th e pr e s e nce o f ph o spho r

e sce nt bact e ria De co mp o si ng m e at s particularly sausag e s
.
,

and fish also may b e co ve r e d with di ffe r e nt Sp e ci e s Of pho s

,

h o re sce nt bact e ria

p .

F o o d R e qui r e m e nts As stat e d ab o ve all th e ph o to g e nic

.
,

bacte ria which o ccur in wate r a re marin e On cultivati o n o f .

th e se bacte ria it is th e r e f o re n e ce ssary fo r th e m e di a to co ntain

two to thr e e p e r c e nt s o dium chl o rid e in addi ti o n to o th e r

salts p e pto n e and so m e carb o n co mp o und such as sugar

, ,

o r glyc e rine .

M agne sium p o tas sium and calcium chl o rid e p o tassium

, ,

s ulphate stimulate th e pro ductio n o f light by bacte ria

, .

Go rham state s that h e has b e e n abl e to gr o w pho spho r e sce nt

bacte ria o n pure ly synthe tic m e di a .

F isch e r s tat e s that th e ph o sph o r e sc e nc e o f th e se a is d ue to

th e g r o wth o f s apr o phytic ph o t o g e nic bact e ria o n d e ad se a

animals and th e subse que nt washing o ff o f th e se f o rms in

‘

la rge numb e rs by th e wave s .

Hans M o l i sch in L e ucht e nd e Pfl an e n r e p o rts th e is o la

z

tio n o f twe nty si sp e ci e s o f ph o to ge nic bacte ria He s tate s

-
x .

that th e fle sh o f cattl e (48 p e r ce nt o f th e sampl e s e xamin e d)

and salt wate r fi sh ( p e r ce nt o f the sampl e s e x min e d) o n
100 a

e xp o su r e to th e air S h o w ph o sph o r e s c e nce nd th e ph o to g e nic

a

bacte ria can b e iso late d F r e sh wa te r fish se ldo m if e ve r S ho w

,

.
2 00 B I O L O GY OF S PE CI A L I ZE D GR OUP S
any ph o sph o r e sce nce Hi s e xp e rim e nts S h o w e d that h o rse
.

fl e sh ( 65 p e r c e nt f sampl e s) a nd cattl fl e sh ( 89 p e ce nt
o e r

o f sampl e s) wh e n pla ce d in th r e e p e c e ntso di um chl ride and r o

all o w e d to st nd in th e air b e cam e ph o sph o r e sce nt a nd th e

a

bact e ria co uld b e i so late d .

O xy ge n R e qui r m nts e
—
O xyg e n is ab so lut e ly ne c e ss ry
e . a

fo r th e pr o ducti o n O f light by bact e ria P h o sph o r e sce nc e i s .

an o xidatio n ph e no m e no n Unde r anae ro bic co nditio ns th e

.

ph o to g e nic bacte ria gro w sl o wly but d o no t p ro duc e any

light until ae ro bic co nditi o n s a re pro vide d Wh e n a highly .

co mbu s tibl e carb o hydrate fo o d is pre se nt ph o spho re sce nc e is ,

gr e atly i ncr e ase d .

F isch e r call s att e nti o n to th e w e ll kn o wn fact that ph o s -

h o re s ce nce is no t n e arly so ma r k e d in s m o o th s e a wat e r a s it

p
i s in r o ugh a nd is m o s t n o tic e abl e o n th e cr e s t o f th e wav e s
,

and in th e wake o f a ship wh e re th e wate r is churne d up .

Wh e n th e o xyg e n o f th e ai r c o m e s in co ntact with th e pho s

p h o re e
sc nt mat e r ial a s it d o e s,
wh e n th e se a wat e r is di s turb e d ,

th e light pr o duc e d is gr e atly augm e nt e d .

T e m p e a tu e R e quire m e nts
r r T h e t e mp e ratur e at whi ch
.

light p r o du ctio n take s pla ce is subj e ct to gr e at va riati n o .

T h e limit s o f ph o sph o r e s c e nce acc o rding to E F S mith a e ,

. .
,
r

fro m 5 C to 2 8 C Light pro ductio n se e m s to b e th e m o s t

—°
.
°
.

int e nse fro m 5 C to 2 0 C Th e ph o sph o r e ce nt bact e ria

°
.
°
. s

o f th e B altic S e a a nd th e N o rth S e a fl o uri sh b e st at 18 C

°
,
.

but ca n gro w w e ll at 0 C r e se mbling in thi s r e sp e ct o the r

°
.
,

inh abitants o f th e N o rth e rn wate rs .

 P H OT O GENI C B AC TERI A 2 OI

t
I so l a i o n . h o spho re ce nt bact e ria ca n b e Obtaine d fo r
—P s

e xp e rim e ntal w o r k by pla ci ng o y s t e r sh e ll s o r th e fl e sh o f

h e rring and h ddo cks in a so lutio n o f two to th r e e p e ce nt

a r

so di um ch l o r id e at 5 C fo r a f w days ‘I h e so lid
°
C to
° ‘

. 10 . e .

mat e rial and th e liquid b e co me bluish gr e n in co l o r and e mit e

a pal e light Th e bact e ria can b e i so lat e d acco rdi ng to th e

.

o r di nary bact e ri o l o gical m e th o ds but must b e gr o w n in m e dia

,

co ntai ni ng a n i nfu si o n O f fi sh e tc a nd two to th r e e p e r c e nt

,
.

o f s o di um chl o r id e .

C ultu r e s in g e lati n c o ntai ni ng th e ab o v e m e nti o n e d ingr e -

di e nts may b e co m e S O lumi no u s as to b e ph o t o graph e d in

th e ir o wn light afte r l o ng e xp o sur e Culture s have b e e n .

r e p o r te d which p r o d uc e e no ugh light fo r a p e r so n to se e th e

face o f a watch in to tal darkn e ss .

Of th e se v e r al vari e ti e s o f ph o t o g e nic bact e r a P se udo i ,

m o nas ph o sph o re sce ns var l iq ue fa cie ns is o ne o f th e m o st co m

.

mo n . Th e Optimum t e mp e ratu r e o f thi s bact e r ium is ab o ut

8 C th e maximum 2 8 C and th e mi ni mum 2

°
°
16 C to . 1
°
.
,
°
C
.
,
—
.

P a th o ge n e s i s E F S mith r e p o r t s o ne di se a se o f animal s
. . .

whi ch is d ue to a di stinctly pho to g e nic bacte rium and that ,

i s th e whit e o r sluggi sh di s e a se o f s and fl e as o n th e S h o r e s o f

F ranc e a nd in th e vicinity o f W o o ds Ho l e M ssachus e tt s ,

a .

N O di s e as e s O f plants hav e b e e n r e p o rt e d as d ue to ph o to g e nic

bacte ria .
 C HAP T ER X IX

Y
Z M O GE NI C B AC TE RIA
n u cti o n D fi ni ti o ns
I tro d . e The r e are c e rtai n sp e cie s
.

o f bact e ria which s e cr e t e withi n th e ir c e ll pr o t o pla s m e nzym e s

o r f e rm e nt lik e
-
sub stance s Th e amo unt o f e nyzm e p ro
.

d uce d by a bact e r ium is m o difi e d by th e quality and th e

am o unt o f th e fo o d whi ch it r e c e ive s Whe n gr o wn up o n a

.

m e di um co ntaining p r o t e in s th e bacte ria pr o duc e pro t e o lytic

,

e nzym e s i n e xc e ss Wh e n g ro wn up o n a sub stance co nta in

.

ing ca rb o hydrate s amylo lytic e nzyme s a re pro duce d N o

, .

e nzym e s a re pr o duc e d with o ut o rganic f o o d o f so me char

acte r .

Th e t e rm zymo g e nic has b e e n appli e d to tho se bacte ria

whi ch pro duce amyl o lytic e nzym e s capabl e o f fe rme nting
carb o hydrat e co mp o unds F e rm e ntati o n as wil l b e p o int e d
.
,

o ut lat e r is a di sti nct e nzym e pr o c e ss

,
As a matt e r o f fact
.
,

all pro c e sse s in which th e e nzym e s o f vari o us kinds are co n

ce rn e d are m o di fi e d f e rme ntatio n s Th e first imp o rtant
'

w o rk o n th e subj e ct o f f e rme ntatio n was do n e by P ast e ur in

185 7 .

Th e zymo g e ni c bacte ria in o ur subse que nt di scussio n s wil l

be co nside re d to b e th o se bact e ria wh o se m e tab o lic pro c e sse s
2 02
 ZYM O GENI C B AC TERI A 2 0
3

are characte riz e d by the hydrating o r o xydi ing actio n o f z

e nzym e s o n carb o hydrat e c o mp o und s .

Al l animal s and plants a re co mp o se d in ad di ti o n to ni tro g ,

e no us c o mp o und s o f a larg e p e r c e ntag e O f carb o n co m

,

p o unds During th e life o f all animal s carb o n in th e fo rm o f

.
,

ca rb o n dio xide ( C0 ) is e xpi r e d co ntinually into th e air

2 .

P a r t o f th e ca rb o n di o xid e th us e xpi r e d by ani mals i s tak e n up

by th e chl o ro phyll b e a ring plants and hi gh e r pigm e nte d alg a

-

and by th e aid o f th e kin e ti c e n e rgy o f th e sunlight is ab

, ,

s o rb e d fixe d and sub se qu e ntly m ade into c o mpl e x o rganic

, ,

co mp o unds I n th e pro ce ss Of plant r e spiratio n a small

.
,

amo unt Of carb o n in th e fo r m o f carb o n di o xid e is again re

tu rne d to th e air Th e re maining p o r tio n o f th e carb o n is
.

no t lib e r at e d until th e plant di e s and is di sint e grat e d Th e re .

is al so a large amo unt o f ca rb o n and nitro ge n li b e rate d whe n

th e animal di e s and is d e c o mp o se d .

Th e carb o n is lib e rat e d f r o m th e a nimal and pla nt b o dy by

th e activity O f ba ct e r ia I t e s cap e s h e r e al so i n th e f o rm o f
.

carb o n di o xide Th e zym o ge nic bacte ria a nd th e sap ro ge nic

.

bacte ria which will b e co n side r e d late r are r e sp o nsibl e fo r

, ,

th e lib e rati o n o f th e s e c o mp o u nds T h e nitr o g e n o u

. s co m
p o unds are act e d up o n chi e fly by th e sap ro ge nic o r putr e
fa ctive bact e ria whil e th e ca rb o hyd r at e co mp o unds and
,

o r ganic acids whi ch a e i n a m e a sur e no n put r e fa ctiv e

,
r a e -

,
r

acte d up o n by th e zymo g e nic o r f e r me ntive bacte ria .

Th e t e rm fe rme ntatio n is use d by vario u s w rite rs in a

“
.

varie ty o f diffe r e nt ways Th e t e rm will b e use d in thi s di s

.
2 04 B I OL O GY OE S PE CI ALI ZE D GROUP S
cussio n m e an th e b r e aking up o f co mpl e x carb o hydrat e
to

co mp o unds by th e a ctio n Of th e e nzym e s o f micro o rgani sm s .

F urth e rm o r e th e t e rms f e r m e nt and e nzym e are

“ ”
,

O ft e n us e d in di ff e r e nt se ns e s F e rm e ntati o n acc o r di ng to
.
,

many write r s is applie d to a dir e ct m e tab o lic f unctio n o f th e

,

viabl e pro to plasm o f c e ll s which se rve s the m a s a s o urce o f

e n e r gy whil e e nzym e a cti o n is no t de p e nde nt up o n livi ng
,

pr o to plasm o f ce lls and do e s no t furni sh a ny e ne rgy to th e

c e lls Th e ab o ve di stinctio ns r ai se many Obj e ctio ns
. .

O r ga niz e d a nd Uno r ga niz e d F e rm e nts — M any writ e rs

.

mak e a divi sio n O f th e ag e nts O f f e rme ntatio n i nto o rgani ze d

fe rm e nts and un o rganize d f e rm e nts o r e nzym e s I t is s tat e d .

that o rganize d f e rm e nt s are livi ng ce ll s su ch as bacte ria , ,

y e a sts m o ld s e tc and a re o nly abl e to p ro duce ch e mical

, ,
.
,

chang e s so l o ng as th e ce ll s a re alive Enzyme s are co n .

sid e re d to b e ch e mi cal s ub s tanc e s th e s e cr e t e d pr o ducts o f

,

ce ll s and capabl e O f p ro duci ng marke d che mi cal r e actio ns

,
”

in o rganic c o mp o und s in th e ab se nc e O f th e c e ll s pr o ducing

th e m (p e psin trypsin ptyali n

, ,
I t is claim e d h o w e v e r
, , ,

that th e o rgani z e d fe r m e nt and th e e nzym e may act in a simi

la r mann e r in many ca se s F i sch e r say s that in b o th ca se s
.

th ch e mical cha ng e s pr o duc e d a re sp e cific a nd a e p e r f o rm e d

e r

with o ut th e l o ss o f e n e rgy and that th e f e rm e nt sub stance s

,

the ms e lve s do no t e nt e r into th e r e actio n F i sch e r al so .

state s that e nzym e s e xe r cise a hydrati ng influe nce whi ch

caus e s th e substance s that are attack e d to tak e up wate r and
fo rm substanc e s o f an e ntir e ly diff e r e nt ch e mical co nstitutio n .
 ZY M O GENI C B AC TERI A 2 05

Th e actio n o f di asta se ( amylo lytic) o n s tarch ( a) and , , in

v e rtase o n ca ne s ug r ( b) may b e giv e n as e xampl e s
a , , .

( )
a C 6I 11005 H2 0 C 6H12 06

( b) C i s on
z H2 0 C 6H12 06 C 6H12 06

( De xtr o s e and L e vul o se ) .

The r e a e no b y pr o ducts such as carb o n di o xide ( CO ) and

r -

o th e r gas e s pr o duc e d with o ut furth e r o xidati o n .

S o m e w r it e r s claim that a nti se ptic s st o p f e r m e ntati o n by

killing th e mi cro o rgani sm s w h il e th e y d o no t s e ri o usly im '

p i r th e e nzym e s but it h a s b e e n n o te d that e nzyme s in dilute

a ,

s o lutio ns a e ju st as susc e ptibl e to th e actio n o f antis e ptics

r .

Th e lat e s t r e se arch o n thi s subj e ct i ndi cat e s that all th e

r acti o ns which ccur du r ing th e f e rme ntatio n o f carb o hy

e o

d a te s are e nzym e p r o c e sse s

r .

As a matt e r o f fact e nzym e s a re s e cr e t e d by all c e ll s cla ss e d

,

by F i sch e r and o th e r s as o rganize d fe r m e nts and it is th e ,

e nzym e s whi ch p r o duc e th e ch e mical cha ng e Th e r e is .

abundant e xp e rime ntal e vide nce to supp o r t thi s state m e nt .

F il tra te s fr o m cul tur e o f bact e ria and bact e r ia kill e d by s uch

s ,

age nts as thymo l co ntain enzym e s o f va rio u s che mical co n

,

stitutio ns . Th e re is no suffici e nt basi s fo r th e di visio n o f

fe r me nts into th e two cla sse s o rganize d and uno rganiz e d ,
.

Th e ch e mi cal pr o ce ss s in b th a e fundam e ntally th e sam e

e o r .

F r m e ntati o n th e r e f r e i s to b e r e ga r d e d as a co mp o sit e
e ,
o ,

ch e mical p r o c e ss o f o xidati o n whi ch i bro ught ab ut by th e

,
s o

hyd r ating actio n o f th e intra c e llular amyl o lytic e nzym e s -

2 06 B I O L O GY OF S PE CI ALI ZE D GROUP S
of bacte ria and s o m e o th e r mi cr o o rganisms o n carb o hydrat e
co mp o unds .

V ari o us substance s are pr o duce d during th e pr o ce ss o f

f e rm e ntatio n G ase s may o r may no t b e pr o duce d ; a cids
.

a nd alc o h o l s a nd e th e rs a nd e st e rs a e f o r me d in s mall quan r

titi e s Th e ga se s which may b e pr o duc e d du ring f e rme ntive

.

p r o ce e a e ca rb o n dio xide o xyge n hydro ge n nitr o g e n

ss s r , , , ,

a nd m tha n oe ma r sh gas Amo ng th e alco ho ls are m e thyl

e r .
,

e thyl butyl p ro pyl glyc e rine and many high e r alco h o ls

, , , ,
.

I t sh o uld b e n o t e d that all f e rm e ntati o n is no t d ue to th e

e nzym e whi ch bact e ria pr o duce A large vari e ty o f y e asts

s .

a nd s o m e sp e ci e s o f m o lds a re c o nc e rn e d in s o m e f e rm e ntiv e

p r o ce s se s Th e maj o rity o f all alco ho lic f e rm e ntatio ns are

.

d ue to y e a s t s ( S accha r o myc e t e s) S o m e bact e ria such as .

,

B a cillu s co li p r o duce small am o unts o f alco ho l

, .

Th e f e rm e ntati o n s d ue di r e ctly to bact e ria will b e bri e fly

co n side re d Th e f e rm e ntive p ro ce ss e s are de signat e d by

.

th m o st imp o r ta nt b y pr o duct pr o duc e d

e -
Thus we S p e ak o f .

lactic acid f rm e ntati o n ace tic acid f e rm e ntati o n and butyric

e , ,

acid f e rm e ntatio n .

I t s e e ms that in th e maj o rity o f bact e rial f e rm e ntati o ns

th e r e is a S p e cific bacte rium o r gro up o f bacte ria which is

p ro mi ne nt in th e pro ce ss .

La cti c Aci d F e rm e nta ti o n — Th e bact e ria c o nc e rn e d in

.

thi s f e rme ntatio n a e v e ry wide ly di stribut e d The y are

r .

fo und chi e fly in milk but may also o ccur in o th e r so lutio ns

, ,

as fo r e xampl e
,
di still e ry mash,
.
 ZYM OGENI C B ACTERI A 2 07

The re a re ab o ut o ne hund re d sp e cie s o f bacte ria whi ch may

cause lactic acid fe r m e ntatio n Th e y can b e gro up e d as th e .

“
lactic acid pro duce r s One o f th e co mmo n sp e ci e s is
.
’

B act a cidi l a cti ci and a v e ry cl o se ly r e lat e d sp e cie s B lactis

.
, ,
°
.

ae ro ge n e s M sp co mma and th e S pirilla b e lo nging to th e

. .

same family B p ro di gio sus a nd many sa r cina fo und in

,
.

b re w e ry mashe s p o s se ss th e p o w e r o f pro ducing lactic acid

,
.

L a ct o se o r milk s uga r ( Cl o ) i s th e o r di nary ca b o h y

gg ll r

d r ate which is acte d up o n by th e lactic acid o rganisms I n .

additio n to lacto se gluco se and cane sugar may b e f e r m e nte d

,
.

S tar ch malto se o c e ll ul o s e a e no t act e d up o n dir e ctly by

, ,
r r

th e e nzym e s o f th e lactic acid bact e ria The y may b e .

change d into th e fo re go ing suga r s h o we ve r by e nzym e , ,

actio n o f o th e r bacte ria and th e n b e sub se que ntly attacke d by

th e lactic acid bact e ri a and chang e d int o lactic acid Th e .

f o ll o wing is appro ximate ly th e ch e mical r e acti o n which take s

place o n lacto se
C mHn n HzO 4 C H3 C H (OH) GOOH (lactic acid ) .

Lactic acid f e rm e ntatio n is an ae ro bic pro ce ss and th e fe r ,

m e ntatio n is imm e di ate ly inhibite d in th e abse nce o f o xyg e n .

Th e maximum a cid f r matio n and ne c e s a r ily th e Optimum

o , ,
s ,

t e mp e r atu r e o f th e lactic acid bacte ria ave rage ab o ut 3 C ,

0
°
.

to 3 5 C in th e maj o rity O f sp e ci e a nd a s l o w as 1 C and

° °
. s, 0 .

hi gh a s 5 2 C in so m e fe w vari e ti e s
°
as . .

L actic acid f e r m ntati o n can p r o gr e ss o nly wh e n th e acid

e

p r duce d is n e ut r aliz e d by so me alkaline sub stance in so lutio n

o ,
2 08 B I O L O GY OF S PE CI AL I ZE D GROUP S
such as cal cium ca rb o nat e e tc L a cti c acid i nhi bit s th e
,
.

gro wth o f m o s t all o f th e Sp e ci e s o f bacte ria co nce r n e d in th e

p ro c e ss wh e n its co nce ntratio n r e ach e s ab o ut p e r c e nt )
e ight t e nths to ( p e r c1e nt ) o ne p e r c e nt .

Ab o ut e ighty p e r c e nt ( 8 p e r c e nt) Of th e f e rm e ntabl e

0

sugar is co nve rte d i nto lactic acid smal l amo unts o f ac e tic , ,

f o rmi c and succinic acids carb o n dio xide hyd ro ge n nitro ge n

, , , , ,

m e than e and alco h o l s u nde r favo r abl co n di tio ns A ce r

, ,
e .

tain amo unt o f lacto se and o th e r sugars alway s r e mains in

s o lutio n unf e rm e nte d .

Ace ti c Aci d F e rm e nta ti o n — Thi s f e rm e ntive o r e nzyme

pro ce ss lik e all o the rs Of this gro up is in part a di stinct
, ,

o xidati o n pr o c e s s Th e bact e rial e nzym e s act o n th e car

.

b o h yd a te s ugars and br e ak th e m do wn i nto e thyl alco h o l

r

and carb o n dio xide Th e o idatio n o f th e alco ho l r e sults in

. x

th e f o rmati o n o f a ce tical d e h yd e a nd lat e r ac e tic acid Th e .

ace t c aci d is stil l fu r th e r o xidiz e d to carb o n di o xid e and wat e r

I .

Th e fo ll o wi ng r e acti o ns o n d e xtr o se will s e r ve to e lucidat e

th e pr o c e ss

C sHmOe HgO 2 C H3 C H2 0H 2 C02 HgO (e thyl

alc o h o l ca rb o n di o xide and wate r)
, ,
.

—
C Hs CHzOI I C H, COH H2 0 thylald hyd
O (e e e or

a ce ti ca l d e h yd e )

CH C OH C H3 C OOH ( ace ticacid )

3
O .

C Hs COOH 4O 2 H2 0 2 C02 ( wate r a nd carb o n di

o xide )
 ZYM O GENI C B ACT E R I A 2 09

M any alco ho lic liquids may b e Oxidi ze d to f o rm ace tic acid ,

carb o n di o xide and wat e r o may b e o xidi z e d di r e ctly into

, ,
r

wate r and ca rb o n di o xid witho u t a ny acid fo rmatio n e .

M axi mum ac e tic a cid f e rm e ntati o n tak e s plac e a t a te m

ra tu e O f ab o ut 3 5 C a nd c e a se s at 4 5 C a nd 5 C t
° ° ° °

p e r .
5 C . . o
—
.

I n a c o nce ntrati o n o f f o urt e e n p e r c e nt ( 4 p e r ce nt) ac e tic 1

acid inhi bits all bact e rial g ro wth .

Ac e tic f e rm e ntati o n i v e ry pr o mi ne nt in th e vi ne gar indus

s

tri e s in so m e co untri s Th e alco h o lic liquids su ch as cid e r

e .
, ,

wine e tc a e b r o ught in co ntact with ace tic acid bacte ria

,
.
,
r

and al so with a fre e supply Of Oxyg e n ( ai r) and by thi s pr o ce s s ,

a e o xidiz e d
r .

Ac e tic acid is f o rm e d in practically all f e rm e ntiv e pr o c e sse s

(lactic butyric ac e tic

, ,
Th e sp e ci e s o f bact e ria whi ch
,

a e abl e to p r o duc e ac e tic acid a re no t S O num e r o us a s th e

r

lacti c acid bact e ria B acil lus ace tici is a co mmo n f o rm Th e

. .

pro ce s s is ae ro bic Th e bacte ria cann o t p ro duc e acid in th e

’

abs e nce Of alco h o l I t is pro babl e that th e carb o n o f th e

.

alco ho l s e rve s as a fo o d .

B u tyri c Aci d F rm e ntati o n e— Th e f o rmati o n o f butyric .

acid by th e e nzym e s o f bacte ria is a pro ce s s whi ch is quite

wide ly di stribute d Th e fe r m e ntatio n 1s car rie d o n o nly
.

unde r strictly anae r o bic co nditio ns At o ne time it was .

tho ught that lactic a nd butyric f e r me ntatio n s w e r e d ue to

e nzym es o f th e sam e bact e ria ( Amyl o bact e r b u ty icus) It r .

h a s si nc e b e e n pr o ve d that sp e cific bact e ria a re th e ca u s e o f

th e maj o r p o r ti o n o f th e s e f e rm e ntati o ns Ab o ut tw e nty .

2 10 B I OL OGY OF S P E CI AL I ZE D GROUP S
diffe r e nt sp e cie s o f butyric acid bact e ria have b e e n iso late d .

Usually in th e butyric acid f e rm e ntati o n as in all Of th e ,

e nzym e pr o c e ss e s o f this typ e hydr o g e n carb o n di o xid e

,
and , ,

trace s o f fatty acids are pro duce d in additio n .

S o m e o f th e butyric acid o rga nisms b e sid e s p o ss e ssing ,

e nzym e s which will f e rm e nt practically all th e carb o hydrat e

co mp o unds will also cause a fe rme ntatio n o f such substanc e s

,

as glyce rin e gum arabic e tc B acill us o rth o b utyricus is a

, , .

bacte rium o f thi s typ e .

S o m e butyric acid bact e ria fo r e xampl e B acil l us b utyri cus

, , ,

p o sse ss in addi tio n to zymo ge nic e nzyme s also sapr o ge nic

e nzym e s capabl e o f disint e grating pro t e ins C o ns e qu e ntly .
,

it is no t unco mmo n to find butyric acid in putr e fying sub

stanc e s B utyric acid can also b e pro duc e d by th e splitting
.

o f c e rtain pr o t e in co mp o unds .

I t is quit e c o mm o n fo r butyric acid b a cte ri a to infe ct s o ur ~

milk containing lactic acid and di sinte grate this acid and
what lact o s e may b e re maini ng Th e fo ll o wing re actio ns .

S h o w what tak e s plac e wh e n lact o s e ( a) and lactic acid ( b)

a re attack e d

( a) C mHmOu H2 0 4 C PL' C H (OH) C OOH (lactic acid) .

( b) 2 C Hs CH (OH) COO H 2 O C H, CH2 C H2 COOH

(butyric acid) 2 C02 2 H2 0 .

B utyric acid bacte rial e nzym e s a re claim e d by so m e write rs

( F isch e r) to b e th e indir e ct caus e o f sympt o matic anthra x

and m alignant oe de ma two anae ro bic di s e ase pro ce sse s

,
.
 ZYM O GENI C B AC TERI A 2 I I

M e th ane ( OH4) and M ucil a gi no u srF e m e nta ti o n Thi s

— .

fo rm o f fe rme n tati o n is d ue to th e actio n o f e nzyme s which ,

a re se cr e t e d by s e v e r al sp e ci e s o f bact e ria o n c e ll ul o se Sp iril ,

.

lum rugul a is o ne o f th e bacte ria whi ch is co nce rn e d in th e

pro ductio n o f m e than e ( CI L) I t is no t a n inf r e q u nt ccu r

. e o

r e nc e fo r wine b e e r ve ge tabl e s and e sp e cially m ilk to b e

, , , ,

co m e slimy and app e ar mucilagino us in characte r Thi s .

co n di tio n is d ue to th e bact e rial e nzym e s fe rme nting carb o

hydrate co mp o unds Th e mucilagino us gumlike sub stance
.
,

( galactan ) is v e ry similar to c e ll ul o s e T h e f e rm e nt e d .

mate rials abso rb a gre at amo unt Of wate r and thus fo rm a

sticky mass .

Th e bact e rial e nzym e s inv e rt th e c e llul o s e o r lact o se int o

de xtr o s e and thi s in tu r n is br o ke n up into m e thane carb o n

, ,

di o xide hydro ge n a nd fatty acids Th e re actio n s are unce r

, ,
.

ta in afte r th e inve rsio n but it s e e m s p ro babl e that th e

,

m e than e r e sul ts fro m th e spl itting o f alco h o l s whi ch a re

fo rme d in th e pro c e ss Th e fo llo wing re pre se nts th e re actio n
.

( C6Hm05 ) n a O C
s 02
a
3 C
.. HI ( m e thane ) .

M e tha ne is o ne of principal co nstitue nts o f p e tro l e um

th e

gas (4 p e r ce nt)
0 . Co al yi e lds as hi gh as 80 o r 90 p e rc e ntag e
o f m e tha ne .

F e rm e nta ti o n of th e Hi gh e r Al co h o l s , Ar o m ati cs , and

F atty Aci d s Glyce rin e and manni te (pro p e nyl alco ho l)

.

o n f e rm e ntati o n by th e e nzym e s o f B a cillus e th ace ticus and

o th e rs may f o rm e thyl alc o h o l ace tic acid fo rmic acid , , ,

2 12 B I OL O GY OF S PE CI AL I ZE D GROUP S
succinic acid carb o n di o xide a nd hydro ge n Ac e tic lactic
, , .
, ,

butyric p ro p rio ni c malic citric succini c a nd o the r a cids o n

, , , , ,

f e rm e ntatio n are e ach co nve rt e d into varying amo unts Of th e

o th e r s and e v e ntually int o ca rb o n di o xid e a nd wat e r Th e .

M pn e um o ni a al o fe r m e nt s manni t e
. s .

F i sch e r r e f e r s to th e vari o u s d e xtr o and l e vo r o tato ry -

acids whi ch may b e pro duce d in f e r me ntatio ns F o e xampl e . r ,

th e e nzym e o f B c o li cha ng e d e xtr o se i nt o lactic acid s whi ch

s .

di ff e rin th e di r e ctio n and th e e xt e nt to whi ch p o lariz e d light

is r o tat e d Th e so urc e o f th e nitr o g e n supply s e e ms to b e a
.

d e te rmi ni ng facto r in thi s ca se I f ammo nium ph o sphat e is .

suppli e d l e vo —r o tato ry la ctic acid r e sults ; whil e if p e pto n e

,

i s u se d a s a s o urc e o f nit r o g e n d e xt r o r o tat o ry acid is p ro

,
-

d uce d . L a ctic acid pr o duc e d by o th e r f e r m e ntati o n s is

o ptically inactiv e ( l e vo a nd d e xt r o r o tato ry) -

.

Z ym o ge ni c B a cte ria i n M il k a nd i ts P r o d u cts — Th e .

lact o s e o r milk sug r whi ch is no rmally pre se nt in milk unde r

a

go e s d e co mp o sitio n by th e actio n o f th e e nzym e s o f vari o u s

sp e ci e s o f ba ct e ria aft e r th e milk h as b e e n s ta ndi ng fo r so m e

tim e at a wa r m t e mp e r atu re Th e acidi ty is e vid nt to th e. e

s e nse o f tast e wh n it r e ach e s th e e quival e nt o f

e
p e r c e nt
lactic acid As th e a cid is p ro duc e d th e cas e in o ge n o f th e
.
,

milk is act e d up o n by bacte rial e nzym e s S imilar to r e nne t ,

and as a r e sult th e case in f o r m e d is p r e cipitat e d thu s p o

, , ,
r

d ucing a cl o t o r curd Thi s tak e s pla ce wh e n th e a cid r e ach e s

.

a co nce nt r ati o n o f p e r c e nt to p e r c e nt T he r e s ult .

ing co agulum o r curd b e co me s mo r e s o lid as th e acid incr e ase s .

 ZYM OGENI C B ACTERI A 2 13

A large numb e r o f sp e ci e s o f bacte ria which are pr e se nt in

mil k pro duce in th e pro ce s s o f f e r m e ntatio n o the r acids b e
side s lactic a cid a nd la rge amo unts o f ga se s such a s carb o n
,

di o xid e me than e ni t r o g e n and hydr o g e n

, , , ,
B act e ria d o no t
.

pro duce as much acid in cre am a s in milk .

I n th e pr o c e ss O f ch e e se ma nu factur e th e p r e s e nc e o f
,

gas pr o duci ng bact e ria may cau se th e curd to flo at and

-

pr o duce a p o o r ch e s d ue to th e h o le s fo rme d by th e gas

e e ,
.

A cu r d d ue to th e r e nn e t e nzym e s o f bact e ria may als o

b e p r o duc e d with o ut th e fo rm a tidn O f any acid O cca sio n .

ally mil k do e s no t so ur rapidly a nd pro duc e s a so ft slimy ,

cu r d in a w e ak a cid s o lutio n Th e co ndi tio n is kno wn to

.

dai yme n as sw e e t curdling

r Th e bact e rial e nzyme s
.

whi ch pr o duce thi s c o ndi tio n are ve ry simil ar to re nn e t

e n zym e s Th e curd pr o duc e d may h e e v e ntually so fte n e d
.

a nd dige ste d by th e actio n o f th e bacte rial p ro te o lytic

e nzym e s Ducl a ux call s thi s dig e s ting e nzym e case ase ”
. .

Th e ch e mical r e acti o ns in th e s e zym o g e ni c pr o c e s se s a re v e ry

co mpl e x .

B utyri c acid f e rm e ntati o n s usually o ccur in milk which

h as b e c o m e so ur d u to th e f o rmati o n o f lacti c acid
e This .

pro ce ss h as b e e n di s cusse d ab o ve Th e fo rmatio n o f butyric

.

acid may tak e place h o w e ve r in a n e ut r al o r faintly alkali ne

, ,

m e di um I n butyric acid f e rme ntati o n o f milk th e bacte ria

.
,

s e c re t e th e ir e nzym e s a nd pr o duc e th e che mi cal change s in

th milk aft e r th e
e xyg e n h as b e e n ab st r act e d by lactic acid
o

bacte ria and o th e r ae r o bic sp e ci e s B utyric acid fe rm e nta

.
 14 B I OL OGY OF S PE CI AL I ZE D GROUP S
tio n is a distinct anae ro bic pro ce ss B utyric acid is also .

f o rm e d by th e de co mp o sitio n Of cas e in and fats in additio n ,

to milk sugar .

I n additi o n to th e vari o us o rganic acids whi ch may b e

pro duce d in mil k so m e o f th e high e r alco h o ls may b e

fo rm e d .

M ucilagin o us f e rm e ntati o n which has b e e n discuss e d p re

,

v io u sl y as o ccurri ng in milk giv e it a thick slimy c o nsi s t e ncy

.

, ,
.

Th e bact e ria in milk a nd th e vari o u s dairy pro duct s such as

che e s e butte r e tc will r e ce ive m o re de tail e d co nsid e rati o n

, ,
.
,

in a subs e qu e nt chapt e r .

B acte ri al F e rm e nta ti o ns in th e Arts — This inte r e sting

.

subj e ct can b e o nly brie fly co nside r e d S o m e fe rme ntive .

pr o ce sse s are Of gre at e co n o mic imp o rtance whi l e o th e rs ,

a re de cid e dly antag o ni stic to c e rtain industri e s .

I n th e r e tting o f flax ( li ne n) jut e h e mp and co co anut

, ,

fib e r fe rme ntatio n s similar in a m e asu r e to putre factive

, ,

pr o c e sse s are imp o rtant in th e s o fte ning O f th e ce llul o se

,

walls o f th e plants so that th e fib e rs may b e se parate d .

I n th e pr e parati o n o f in di g o fro m th e indig o pla nt (I ndig o

f e ra tincto ria) and in th e curing o f hide s and t o bacco bac ,

te rial e nzym e s are imp o rtant adjuncts to th e pro c e ss e s .

B acte rial e nzym e s are also o f co mm e rci al imp o rtance as ,

b e fo r e state d in th e pre paratio n o f vi ne gar ( ace tic acid)

, ,

butyric and lacti c acids and in many dairy pro ce sse s such
, ,

as cr e am rip e ning ch e e s e and butte r making

,
.

B acte rial fe rm e ntive e nzym e s are r e sp o nsibl e fo r th e un

 Y
Z MO GENI C B ACT ERI A 2 1
5

pl e a sant flavo rs and O do rs which are fr e qu e ntly de ve l o p e d in

fo o d pro ducts Th e y are al so partially r e sp o n sibl e fo r th e
.

und e sirabl e flavo rs o cca sio nally de ve l o p e d in alco ho lic liqui d s .

Z ym o g e nic bact e rial e nzym e s are u se ful in c o njunctio n with

y e asts in br e ad making L actic acid f e rm e ntati o n o ccurs

.

in th e f o rmati o n o f e nsilag e. I n br e ad making th e ba ct e r ial

e nzym e s pr e par e th e s tarch ( fl o ur) by th e acti o n o f amyl o lytic

e nzym e s fo r th e sub se qu e nt acti o n o f th e zym o g e nic e nzym e s

o f th e y e a s ts a nd by the lactic a nd ac e tic acid which th e y

,

g e n e rate give a pl e asant flavo r to th e bre ad .

 CHAP T ER XX
THE SA PR O GENI C AN D SAPR O P HI LI C B AC TE R I A

I ntro d ucti o n . t
De fi ni i o ns
.
— I n th e pr e vi o u s chapt e r it

was p o inte d o ut that e nzym e s p ro duc e d by bact e ria a re re

sp o nsibl e fo r all th e change s o ccurri ng in th e m e di a in whi ch
th e o rganisms a re gr o wing and no t th e bact e rial c e ll i n t t o o .

I t is a w e ll k no wn fact that plants a s a wh o l e s e cr e t e o nly

-

a small am o unt Of ca rb o n dio xide afte r th e natural abso rpti o n

o f carb o n and o xyg e n in this f o rm ( C 0 ) and n e v e r lib e rat e
2 ,

and ve ry rar e ly ab so rb any ni tr o ge n fro m th e air N itr o .

g e n which has b e e n tak e n up fr o m th e nit r at e s o f th e so il is

r e turn e d th e r e to o nl y wh e n th e plant di e s and its pro t e in

co mp o unds a re d e co mp o se d Th e sam e ca n b e said o f th e
.

gr e at e r am o unt o f th e nitro ge n co ntaine d in animal ce lls I t .

is lib e rat e d o n th e d e co mp o sitio n o f th e animal Th e r e is

.
,

h o we v e r in th e animal a small am o unt o f nitr o ge n s e cre t e d

,

in th e milk and e xcr e t e d in th e u rine (ur e a u ric acid hi ppuric

, ,

acid ,
Th e se co mp o unds a e e ve ntually bro k e n do wn
r

into amm o nia and fr e e nit ro ge n by th e actio n o f bact e rial

e nyz m e s .

Tho se bacte ria which bring ab o ut th e pr o fo und d e co m p o

sitio u o r putr e factio n Of nitro g e n o us co mp o unds are call e d
2 16
 TH E SA PRO GENI C AN D S AP R O P HI LI C B AC T ERI A 2 17

sa p ge ni c bacte i a B vulgaris M ig is a typical e xam pl e

ro r . . .

o f thi s cla ss Al l putr e factiv e pr o ce ss e s a e br o ught ab o ut

. r

by th e a ctio n o f ba cte rial e nzym e s Th e se e nzym es are in .

,

th e main p r o t e o lytic
,
Th e re a e still o th e r ba cte ria which
. r

de rive nutrim e nt fr o m and p e rhap s I n so m e cas e s aid in b r e ak

ing up still fu r th e r th e c o mp o u nds f o r m e d by th e s apr o g e nic
bact e ria Th e se o rgani sm s a e call e d s p o phi l i c bacte i a
. r a r r .

B c o li Sp und ul a a nd B subtilis a e e xampl e s o f bact e ria

.
,
.
,
. r

whi ch b e l o ng to thi s la st m e nti o ne d class -

.
.

Th e s apr o g e nic bact e r ia a e v e ry wid e ly distribut e d

r Th e .

e nzym e acti o n o f th e s ap r o g e ni c sp e ci e s u nd ubt e dl y s e rv e s o

the se o rgani sm s as a so urce o f e ne rgy I t was supp o se d dur .

ing th e e arly days o f bact e ri o l o gic l r e s e arch that d e co m p o a

si tio n was d ue to o ne sp e ci e s o f ba ct e r ia w hi ch was call e d

B acte rium t e rm o I t is no w kn o wn that putr e fa cti o n is d ue

.

to a la r g e numb e r o f S p e ci e s o f s ap r o phytic b ct e ria Th e se a .

bacte ria are no t always fo und und r th e same co nditio ns e .

N att e mpt has b e n mad e to cla ssify th e p u t r e factiv e bac

o e

t ria I t se e m s that s o m e sp e ci e s a re pa r ti cula rly co mmo n

e .

in de co mp o si ng animal mat e rial o the r s in d e co mp o sing ,

v e ge tabl e mat e rial and o th e r s in de co mp o si ng u re a

,
.

P ro te o l yti c Enzym e s — T h e p r o t e o lytic e nzym e s o f bac

.

t e ria r e s e mbl e in th e i r a cti o n th e tryp sin o f th e pancr e atic

juice o f th e animal b o dy in many particula rs Th e s e e nzym e s .

a ct b e tt e r i n a n alkali ne than in an a cid m e dium Th e r e is a .

p r o te o lytic e ny m e whi ch is se cr e te d by a l rge va ri e ty o f

z a

bacte ria which disso lve s o r p e pt o niz e s ge lati n co mp o unds .

2 18 B I O L O GY OF S PE CI AL I ZE D GROUP S
This e nzyme will r e sist a te mp e ratur e o f 1 C fo r fifte e n 00
°
.

minute s We ll s r e fe r s to th e stat e m e nt o f S chm ail o witsch

.

that so me bacte ria pro duce an e nzym e which acts in an acid

m e di a o n ge latin but no t up o n album e n and change s it ,

into pro t e o s e s and p e pto n e s He state s that tho se acting

.

in an alkalin e m e dium carry th e pr o ce ss still furth r to e ,

th e p o i nt wh e r e amin o acids such as l e ucin and tyr o si n a re

, ,

pr o duce d .

Th e pr o t e o lytic e nzym e s o f path o g e nic bact e ria in c e rtain

cas e s a e pro min e nt in putr e factive pr o c e s se s The r e is no

r .

r e lati o n b e twe e n th e pr o t e o lytic e nzym e and th e patho g e nicity

o f a sp e ci e s o f bact e ria . Th e r e is a p o ssibili ty that th e bac
te rial e nzym e s d o play a min o r part in th e di g e s ti o n o r d e

co mp o siti o n Oi th e e xudat e s in suppuratio n .

Th e r e a re also o th e r pro te o lytic bacte rial e nzyme s which

pro duce a co agulatio n o f case ino g e n and lat e r a dige stio n O f
th e cas e in o f milk . O th e r sp e ci e s O f bact e ria pr o duc e p ro

te o l yti c e nzym e s which a e capabl e o f acti ng o n th e fibrin o f

r

bl o o d and th e bl o o d co rpuscl e s changing the m into th e p ro

,

te o se s and p e pt o n e s and lat e r int o th e ami no acids

,
He m .

o l ysi s is cl o s e ly r e lat e d to pr o t e o lytic e nzym e acti o n Al l .

e nzym e s capabl e o f di g e sting g e latin will also di ge st th e

case in Of milk (Eijkman) .

N e arly all th e putr e factive bacte ria pr o duce pr o t e o lytic

e nzym e s whi ch a e abl e to br e ak th e pr o t e in m o l e c ul e d o wn
r

to its simpl e s t c o mp o u nd s A larg e numb e r o f s apr o phytic

.

bacte ria pro duce pro t e o lytic e nzym e s and it is thi s class
,
 THE SA PR OGENI C AND S AP R OP HI LI C B ACT ERI A 2 19

whi ch pro duce th e maj o rity o f putr e factio n Th e f o ll o wing .

tabl e fro m We lls s C he mical P atho l o gy giv e s a pa r tial ’ ”

li st o f so me o f th e mo re imp o rtant bacte ria and o the r mi cro

o rganisms whi ch pr o duc e di g e sti ng p r o t e o lytic e nzym e s

P R OT E OL Y
TI C DI G E S TIN G E N Z YM E S

1 . B a t a nthra i
c . cs

B . co li
M pyo . a ur . e us

S tr . py g n o e es

P S pyo cya ne a
.

B vi la
. o ce us

B m y id
. co es

B p digi
. ro o s us

A p rgill
s nig r ( m ld )
e us e o

A p rgill
s y e us o r z oe

ab o ve tabl e sh o ws that th e r e is a se l e ctive actio n by

Th e
'

th e bact e rial e nzym e s fr o m di ff e r e nt sp e ci e s o n th e vari o us

pro te in co mp o unds Th e dige stio n o f g e latin take s place .

in a de finite mann e r amo ng th e vari o us sp e cie s o f bacte ria

( se e F ig .

bact e rial e nzyme s which might b e co nside re d as

A fe w
ste at lytic have b e e n n o t e d in put r e fying flui ds P s
o . .

p y y e a B p ro di gi o sus M py o g e ne s var au r e u s a e
o c a n ,
.
,
. .
,
r

amo ng th e b e st kno wn bacte ria which pro duce fat splitting- -

2 2 0 B I O L O GY OF S PE CI AL I ZE D GRO UP S
0

e nzym e s Glyce r ine and th e va ri o u s fatty a cid s a re am o ng

.

th e e nd pr o duct s
-
I t is a difli cul t matt e r to i s o lat e th e par
.

ti cul ar sp e ci e s o f bact e ria in putr e fyi ng fluids which pr o duc e

th e st e at o lytic e nzym e s .

I n o rd e r that putr e facti o n tak e plac e c e r tain o f th e p re ,

r e quisite s o f bact e rial gro wth mus t b e at hand A ce r tain .

amo unt o f m o i stu r e is abso lut e ly n e ce ssary I t is a fam iliar .

fact that pro te in mate rial such as m e at wh e n dri e d do e s no t

, , ,

de co mp o se T e mp e r atu r e is ano the r imp o rtant r e qui r e m e nt

. .

Lo w t e mp e ratur e s pr e ve nt th e acti o n o f th e putr e factiv e

bact e rial e nzyme s F i sch e r r e fe r s to th e fi ndi ng o f th e Sib e

.

rian mammo th s whi ch had b e e n fro z e n in th e ice fo r ce nturi e s

and th e fle sh was still in go o d co nditio n .

Th e bact e r ia co nc e r ne d in th e put r e factive p r o ce s se s are

ae r o bic ana e ro bic a nd facultative a nae ro bic Th e ae r o bic

, ,
.

bact e ria by u si ng up th e supply o f o xyge n a nd fo r ming ca rb o n

di o xid e thu s pro vide th e ne ce ssary co nditio ns fo r anae ro bio sis ,

and co ns e qu e ntly th e rapid g e ne r ati o n o f gas e s and di sin

te gra tio n o f th e c o mp o u nd Ae r o bic bact e ria in p utre fa c
.

tio n a re o n th e who l e o f se co ndary imp o rtance

, ,
.

M e ch ani s m o f P utre f a cti o n P ut r e facti o n is primarily

.

a pro ce ss o f d e co mp o sing and splitting up nitro ge no u s co m

p o unds by th e actio n o f th e p ro t e o lytic e nzym e s o f bacte ria .

I t is in a m e a s u r e a f e rm e ntiv e p r o c e s s I t sh o uld b e r e m e m
.

b e r e d that in e ve ry case wh e n an animal o r plant co mp o und

is d e co mp o sing that th e zymo ge nic o r f e rm e ntive bacte ria
a re acting o n th e ca r b o hydrat e c o mp o u nd s at th e sam e tim e
 THE SA PR OGENI C AN D S AP R OP H I LI C B AC TERI A 2 2 1

that th e sapro g e ni c bacte ria are acting o n th e nitro ge n o us

co mp o unds Th e pr o ce sse s usuall y go hand in hand Th e
. .

pro c e ss o f true putr e factio n ho we ve r tak e s place o nly in

, ,

nitro g e no us co mp o unds A di stinctio n is s o me time s made

.

b e twe e n put r e factio n a nd de cay Th e fo rm e r is h e ld to b e a

.

di stinctly ana e r o bic p r o c e ss whil e th e latt e r i s appli e d to

,

tho se de co mp o sitio n s taking place in th e pre s e nce o f o xyge n

and d ue to ae ro bic bacte ria .

P r o d u cts P o d u ce d
r . We will di s cu s s as bri e fly as p o s sibl e
th e r e sults O f th e d e co mp o s iti o n a nd S O call e d putr e facti o n
-

o f th e p r o t e i n m o l e cul e whi ch c o ntains th e nitr o g e n Of n e arly

all o rgani c substance s .

I t may b e h e r e s tat e d that th e pr o t e in m o l e cul e can b e split

by many age nci e s amo ng which a e sup e rh e at e d st e am acids

,
r ,

a nd alkali e s in diff e r e nt co nc e nt r ati o ns a nd by bact e r ial

,

e nzym e s . T h e cha ng e pr o duc e d by bact e rial e nzym e s a e

s r

th e m o s t pr o n o u nc e d a nd th e s e c o ndary p r o duct s pr o duc e d

,

the r e by are di ffe r e nt f ro m th e cl e avage pro ducts pro duc e d

in o the r ways I n all ca se s th e p ro te in mo l e cul e is chang e d
.
,

into co mpo unds whi ch a re cl o se ly r e late d to th e o riginal

typical pr o t e in mo l e cul e ; nam e ly p t se s a nd p e p t ne s
,
ro e o o .

At thi s p o i nt th e li ne s O f cl e avag e may b e diffe r e nt a nd many

’

e nd pr o duct s r e s ult
-
. B y th e acti o n Of th e bact e r ial nzym e s e

o f s ap r o g e nic bact e ria th e se pr o t e o s e s ( album o s e s ) a nd e

p p
to ne s are split i nto a s e rie s o f simpl e sub stanc e s whi ch se e m
to b e th e usual e l e m e ntary pr o ducts o f th e pr o t e in m o l e cul e .

The y are as fo llo ws :

 B I OL O GY OF S PE CI ALI ZE D GR OUP S

G ly co co ll
i
Le uc n
S im p l a m in a id
e o c s o f th e
Ala ni n fatt y a id ri c se es
Am i n val ria ni a id
o -
e c c

A p arti a id
s c c
Dib as i c
a id l l y r lat dc s c o se e e
G l uta m i ni a id c c
t m nat m i a id
o o o c c s
Ph ny lala ni n
e

T yr i n os
Ar m ati
o n tit u nt f th
c co s e so e

Tryp t p ha n o
pr t id m l ul
o e o ec e

a-pyrr li d i n arb x y li
o c o c (pro l in) Pr babl y fr m m th r
o o so e o e

11 Oxy pyrro lidi n carb o x y li c a ci d

a
a m in a id b y r arra ng
o c e e

m nt
- -
.

e
(o xy p ro lin) -

12 . Se ri n T g th r with
o e e 10 , 1 1,

bri ng pr t i n in l o e c o se re

lati n t arb h y drat

o o c o es

He x o ne ba se s

Su p l h u xi t r e s s in thi s fo m r

Itwil l b e no t e d that th e simpl e amino acids o f th e fatty

acid se ri e s a re glyco co ll l e ucin alanin and amin o val e rianic
, , ,
-

a cid Th e se acid s a re cha r act e rize d by th e pre se nce o f an

.

N H gr o up attach e d to th e carb o n at o m in th e alpha p o siti o n

2 .

Th e co mm o n f o rmula is R CHNH C OOH O n putre facti o n

- -
g .

th e e sub stance s pro duce many co mp o unds Th e vari o us fatty

s .

acids can b e e a sily d e rive d f r o m th e m o n th e splitting O ff o f

th e N H gr o up Glyco co ll o n splitting pr o duce s ace tic acid
2 .
,

and alanin o n Splitting pro duce s pr o pi o nic acid We lls state s .

that in all pro bability ace ti c and butyric acid are m o s t co m

m o nl y pr o duc e d in this way I n th e pr o c e ss o f th e putr e .
 THE PRO GENI C
SA AN D S AP R OP HI L I C B ACTERI A 2 2 3

factio n Of the se amin o a cid s vari o us gase s such a s hyd r o ge n , , ,

marsh gas amm o nia a nd carb o n di o xid e a re e vo lve d Ace J

, , , .

to n e is in all pro babili ty als o fo rm e d fr o m th e se a m in o

, ,

acids .

F r o m th e diamin o acids (h e x o n e bas e s) s o m e v e r y i nt e e st r

ing substanc e s a e f o r me d Th e se sub stance s diamin s

r .

a re fo r med by bacte rial actio n and a e cha r act e rize d by th e ,

r

fact that th e y r e tain th e N H gro up in s te ad o f Splitting it O ff 2 ,

as in th e fo re go ing cas e The s e substance s a re call e d p to m ains .

,

and fr e que ntly o n abso rptio n pro duce s e ri o us into xicati o ns

, ,

in th e animal b o dy The y will b e di scuss e d in Chapte r .

XX I I N o t al l pto mai ns are pro duc e d fr o m thi s gro up One

. .

o f th e pri ncipal o ne s pr o duc e d is putr e scin ( N

Th e ar o matic substanc e s O f th e pr o t e in m o l e cul e a re p h e nl

alani n trypto phan and tyr o sin Th e re a re vario us sub

, ,
.

stanc e s fo rm e d fro m th e se b o di e s Amo ng tho se pro duce d .

a re ind o l cr e so l and ph e no l fro m tyr o sin ; indo l p r o pi o nic

, ,
-

acid indo l ac e tic acid fro m trypto phan and f ro m th e s e skato l

,
-

a nd ind o l I n i nt e s ti nal put r e facti o n th e i nd o l a nd skat o l are

.
,

abso rb e d o xidize d and co mbin e d with sulphuric o r glycuro ni c

, ,

acid and app e ar in th e urin e as th e S O call e d e th e r e al sul

,
-

ph a te s .

F ro m principal sulphur co nstitu e nt o f th e pro te in

th e

m o l e cul e cy s tin m e thyl m e r captan ( C H S H) e thyl m e rcap

, , s ,

ta n ( C H S H) e thyl sulphid e ( ( C H ) S ) a nd larg e am o unts o f

2 5 , g s 2 ,

hyd ro ge n sulphide (11 8) are pro duce d Hydro g e n sulphi de 2 .

,

to ge th e r with i ndo l and skat o l a e large ly r e sp o nsibl e fo r th e ,

r
2 2 4 B I OL OGY OF S PE CI AL I ZE D GR OUP S
di sagr e e abl e O do r atte ndant o n all putr e factive pr o ce s se s .

Th e se substance s are pr oduce d to a large e xt e nt in th e

human inte stin e Th e pro ce ss tak e s place in th e ab se nce
.

o f o xyg e n .

On th e d e c o mp o siti o n O f th e nucl e in o f th e nucl e o pr o t e in ,

pho spho ric acid and th e vari o us purin base s are pro duce d .

Th e vari o us fat c o mp o und s a re al so de c o mp o se d by th e

actio n Of th e e nzym e s o f sapr o g e nic bacte ria Th e change s p ro .

d uce d are v e ry si m ilar to th e f e rm e ntati o n o f carb o hydrat e s .

G as e s are e vo lve d and glyce ri ne and th e many fatty acids

,

CJ I 5 ( O C 15 H3 1C O) 3 3 H2 0 C H (OH) . 3 CB HC OOH
SI

Palm iti n G l y ri n
ce e Pal m iti a id
c c

a re pr o duc e d B utyric acid is o ne o f th e co mmo n fatty

.

acid s which is pro duc e d in fat de co mp o sitio n Th e l e cithin .

( C H 2 4N P
340 ) wh i ch
9 , is a ph o sph o riz e d fat and c o mm o nly ,

c o mbin e d with th e pro t e in m o l e cul e can b e e a sily split into ,

th e pt o main ch o lin and thi s int o a no th e r v e ry t o xic pt o main

,

call e d n e urin Th e se pto main s wil l b e di scusse d e lse wh e r e

. .

As stat e d h e r e t o f o r e in putr e facti o n th e r e is als o s o m e fe r

,

m e ntati o n Of th e carb o hydrat e co mp o unds Th e e nzym e s .

pro ducing th e se fe rme ntati o ns have th e p o w e r o f hydrating

d e xtr o se malto se inul o se sta r ch e tc Th e pro ducts o f thi s
, , , ,
.

pro ce ss includ e practically all o f th e fatty acid s and th e o y x

acids amo ng whi ch are lactic succinic and o xybutyric e tc

, , , ,
.

I n additi o n to th e s e ac e t o n e o xalic acid a nd s mall qua ntiti e s

, , ,

O f e thyl alc o h o l a re f o rm d M e than e carb o n di o xid e and

e .
, ,

hydro ge n are th e principal gas e s pr o duce d “

.
 THE PR O GENI C
SA AND S AP R OP HI L I C B ACT ERI A 2 2 5

Th e s e nume ro u s put r e factive pr o ce sse s may o ccur in th e

inte stinal t r act o f all animals C o n di tio ns th e r e a re p a ticu
. r

l arl y fa o rabl e to th e acti o n o f a nae r o bic bact e ria

v .

Th e ul timat e e nd pr o duct s o f all putr e factiv e pr o c e s se s a e

-
r

fr e e ammo ni a ( NH ) carb o n di o xide ( C 0 ) hyd ro g e n ( H)

3 , 2 , ,

and wat e r (H O) u nd e r ce rtain co nditio ns p ho sph o ric a cid

z

( H P O
3 ) ph o sph
4 , o r e tt e d hydr o g e n ( P H ) F r e e nitr o g3e n. N
( ) is
undo ubte dly e vo lve d in so me case s .

Th e pr o c e s s o f putr e facti o n is g re atly influ e nc e d by th e

pr e s e nce o r ab se nce o f o xyge n Th e di sagre e abl e aro matic

.

O d o rs a re no t pr o duc e d by a e r o bic bact e rial pr o c e s se s ( d e cay ) ,

but a e e vo lve d o nly wh e n o xyge n is e xclud e d Th e pr e se nce

r .

o f o xyg e n a cc o mp l ish e s rapid O xidati o n and th e o d o r i s no t ,

n o tice abl e aft e r a sho rt space o f tim e .

F o r th e purp o s e o f summarizing we can say that th e fo l ,

l o wi ng substance s r e sul t fro m th e putr e factio n and f e rm e nta

ti o n o f o rganic co mp o unds in all co mp o site de co mp o sitio n
pro ce sse s .

1 . P ro te ins

( a) Ar o matic radi cals ph e nylalani n trypto phan and , ,

tyro sin whi ch by Splitting pro duce indo l skato l

, , ,

ph e no l c r e so l
,
.

( b) S ulphur radi cal s hydro ge n sulphide e thyl sul ,

p hi d e e thyl
,
and m e thyl m e r capta n .

(c) F atty acid radical s fo rmic ac e tic pr o pi o nic

—
, , ,

butyric val e rianic pal m itic ste aric acids ace to n e

, , , ,
2 2 6 B I OL O GY OF S PE CI AL I ZE D GR OUP S
and vario us gase s such as carb o n di o xide am , ,

m o ni a m e thane and hydr o ge n P to mains such

, ,
.
,

as cadav e r in e th yl d e ndi am in and putr e scin

, ,
.

( )
d P h o sph o ric acid and purin bas e s ( fr o m nucl e in ) .

2 . F ats
Glyce ri ne and all th e hi ghe r fatty acids P to mains .
,

such as ch o lin n e u rin e tc fro m th e pho spho rize d

, ,
.
,

fat le cithin .

3 . C arb o hydrat e s

Th e f o ll o wing fatty acids have b e e n iso lat e d and p ro b

ably o th e r s o ccur whi ch have no t b e e n i so late d ,

fo rmic ac e tic pro pi o nic butyric val e rianic

, , , , ,

succinic o xybutyric G as e s such a s hydro ge n

,
.
, ,

carb o n di o xid e m e than e in th e cas e o f ce l l u

,

l o se and ace to n e
,
.

M o stplants O b tain th e ir ni tro ge n supply fro m th e ni trat e s

o f th e s o il Th e nitr o g e n o f th e s o il is th e ke y to s o il
.

f e rtility As a r e sult o f th e actio n o f th e e nzym e s o f vari o us

.

S p e ci e s o f sapr o g e nic bact e r ia amm o ni um salt s a e pr o duc e d

,
r

afte r th e ammo ni a whi ch is f o rm e d during th e p ro c e ss Of d e

c o mp o sitio n is co mbin e d with ce rtain ino rganic e l e m e nts such ,

as s o dium and p o tassium Only a small amo u nt if any o f

.
, ,

the se ammo nium salts can b e u se d by plants a s a so u r c e o f

nitro g e n supply B y th e actio n o f th e ni tr o so bacte ria whi ch
.

a re pr e s e nt in th e s o il th e s e amm o nium salts a re br o k e n up

,
 THE P
SA R O GENI C AN D S AP R OP HI LI C B AC T ERI A 2 2 7

and co nve rte d into nitrit e s Th e nitrite s thu s f o rm e d a re

.

agai n acte d up o n by th e nit rifying bact e ria o f th e so il and as ,

a r e sul t nitrate s a re pro duce d I n thi s manne r th e nitro g e n

.

l o cke d in th e prote in m o l e cul e is again made avail abl e fo r

plant use This pro ce ss is kn o wn as nitrificati o n B ac
. .

t e ria whi ch have th e p o w e r Of ab stracti ng th e o xyg e n o f th e

ni t r at e co mp o u nd a e n o t e d in so m e so il s
r . Th e s e o rganism s ,

by agai n b re ki ng up this co mp o u nd which is us e ful to th e

a ,

plant a e o bvi o u sly v e ry d t rim e ntal to so il f e rtility This

,
r , ,
e .

class Of bacte ria is call e d de nitrifying o rganisms .

 C HAP T ER XXI

PATH O GE NI C B ACTE R I A

In tro d u cti o n
. De fi niti o ns Tho se bacte ri a whi ch p ro
.
—

duce m o rbid change s in th e animal o r plant b o dy und e r ,

natural co ndi tio n s o r unde r co ndi tio ns artificially pro duc e d

by in o culati o n are call e d p atho ge ni c bacte ri a Th e t e rm is
,
.

r e lative and applicabl e o nly within ce rtain limits S o me o f .

th e o r di na r ily ha r ml e s bact e ria wh n int r o duc e d int o th e

s ,
e

b o dy o f th e animal o r plant in large numb e rs p ro duc e m o r

'

bid a ff e cts Th e r e is al so no o ne path o g e nic bacte rium

.
, ,

which will pr o duce dis e ase d c o nditi o ns in a l l a ni mal s o r pla nts .

S o m e va r i e ti e s o f bact e ria a e path o g e ni c fo o ne sp e ci e s o f

r r

animal o r plant and no n patho ge nic fo r o th e rs

-
.

Th e t e r m p a tho g ni c h as c o m e to b e u s ually appli e d to a

e

bacte rium o r o th r m icro Og ani sm which pr o duce s di se ase

,
e r ,

und e r no rmal co ndi tio n s .

M a ny sapr o phyti c bact e r ia a re path o g e nic fo r man and fo r

s o m e S p e cie s o f animal s and plant s Th e se bacte ria may ac
.

cid e ntall y gai n e ntra nc e i nt o th e ti s su e s o f th e b o dy o r may

b e i ntr o duc e d fo r e xp e rim e ntal purp o se s B act e ria O f this .

clas s are call e d facul t ti ve p a asi te s Th e re are s till o th e r

a r .

bacte ria which give all th e e vid e nc e s o f b e ing parasit e s no r ,

mally but o ft e n fo r varying p e ri o ds o f tim e may live as

, , ,

2 2 8
 PATH O GENI C B AC TERI A 2 2 9

sapro phyte s As e xampl e s o f thi s last me nti o n e d class o f

.
-

bacte ria M sp co mma and B typh o sus may b e give n B act e ria
, . . .

o f thi s cla ss a r call e d facul tati ve sa pro p hyte s

e .

The re are o th e r S p e cie s o f bacte ria which are no t abl e to

FI G 4 6
. . Ba t t b r l i
c . u e cu o s s in h l g
um an un . Af t rM
e uir and Rit hi
c e .

e xist fo r any l e ngth o f tim e o ut side o f th e b o di e s o f th e animal

whi ch the y I nfe ct Th e e bacte ria are calle d st i t o r o bl igate
. s r c

p asi t
ar es B act tub e r cul o si s and B act l e pra a re e xampl e s
. . .

o f thi s cla ss I t S h o uld b e no t e d in this c o nn e cti o n that

.

bacte ri o l o gists have b e n abl e to cultivate so m e o f th e s e strict

e

parasit e s o n sp e cial artifi cial m e dia whi ch is made to r e se mbl e ,

2 3 0 B I OL O GY OF S PE CI AL I ZED GROUP S
as n e arly as p o ssibl e th e pabulum o n whi ch th e se micro
,

o rga ni sms gr o w naturally .

I t is supp o s e d that at o ne p e ri o d o f tim e all path o g e nic

bacte ria w e r e sapro phyte s but having th e ir bi o l o gical charac

, ,
~

FI G 4 7. Film pr parati
.
— e on of di harg fr m a w
sc e o di a a
o un n c se of t tan
e us,

h wing B t tani i dr
s o . e n um ti k sp r f rm ati n
c S o e o o . Aft r M ir
e u an d R it hi
c e .

t e rs m o difi e d th e y have b e co m e parasitic o n acco unt o f natu

,

ral s e l e ctio n and e nvir o nm e nt .

E xp e rim e nts hav e b e e n mad e with vari o us sp e ci e s o f bac

te ria and it has b e e n sh o wn that it is p o s sibl e to m o dify bac

,

t e ria to such an e xte n that many O f th e strict para sit e s ca n

’

b e made to b e co m e sapro phyte s and c o nv e r s e ly S o m e sap ,

.
 PATHO GENI C B ACTERI A 2 3 1

ro ph ytic bacte ria may b e indir e ctly p atho g e nic by r e aso n o f

th e fact that th e y may fr e qu e ntly multiply in vari o us f o o d

pro ducts such a s m e ats milk ch e e s e e tc and by th e ir sap

, , , ,
.
, ,

ro ge ni c pr o c e s se s pro duce p o i so no us pro t e i n sub stance s

,

FI G 48 —
. . Trypan o so m a br i f ag ana Th trypa
uc o n . e no so m e is arri d by th
c e e

t t fly Aft r William
se se . e s .

calle d pto mains Th e s e pt o mains wh e n take n int o th e b o dy

.
,

and ab so rb e d by th e lymphatics and bl o o d give ris e to se ri o us ,

into xicati o ns The s e substance s will b e co nside re d m o r e in

.

d e tail lat e r .

Th e inj e cti o n o f c e rtain sp e ci e s o f true sapr o phytic bact e ria

und e r th e skin o f s o me o f th e e xp e rime ntal animal s is O ft e n

atte nde d by th e pro ductio n o f ab sce sse s n e cro sis and g e n e ral
, ,
2 3 2 B I O L O GY OF S PE CIAL I ZE D GR OUP S
All ab o ve m e ntio n e d facts and e xp e rime nts s e rv e to
th e -

de mo nstrate th e co nclusio n whi ch can b e de fi nite ly st te d ; a

name ly that th e limits o f th e patho g e ni c bacte ria a e no t

,
“
r

de fini te .

Di se a s e s I nf e cti o us and C o n tagi o u s

. All dis e as e s p ro .
-

FI G .
40
—. L i hm a D v n b di f kala azar I t i pp d that at
e s n -
o no a o es o -
. s su o se ce r

tain s tag th b di
es e sefl g l l t d im ila t F i g r 4 8
o e s are a e William
a e s r o u e . s .

d uce d by bact e ria and o th e r micr o o rgani sm s are call e d i h

f e cti o ns d i se as e s M any d e finiti o ns hav e b e e n s ugg e st e d by
.

patho l o gis ts fo r th e t e rm inf e ctio n Acco rding to Z i gl e r . e ,

i nfe ctio n is th e e ntrance o f bact e ria i nt o th e b o dy and th e ir

incre ase the r e As M cF a rl and s ays this u sually m e ans an
.
,

e ntranc e int o th e tissu e s O f th e b o dy and a multiplicati o n

o f th e bact e rial c e lls in th e ti s su e s S o m e path o g e nic bac .

 PATHO GENI C B ACTERI A 2 33

te ria pro duce di se as e co ndi ti o ns witho ut e nt e ring th e tissue s

o f th e b o dy F o r e xampl e M sp c o mma p r o duc e s p o is o n o u s
.
,
.

e ff e cts o n th e muc o us m e mbran e o f th e int e stin e s with o ut

e nt e ring th e ti s su e s .

Th e m e r e e ntranc e o f th e bact e ria into th e b o dy o r th e tissu e s

O f th e animal o r plant is o nly a part o f th e pr o c e ss o f inf e cti o n .

I n additi o n th e r e mu s t b e so m e inju r y to th e b o dy c e lls As .

h a s b e e n m e nti o n e d ab o v e th e harml e ss sapro phytic bact e ria

,

may b e pr e se nt in th e b o dy o f an animal and ne ve r pro duce

any mo rbid e ffe cts until fo r s o m e r e a so n o r o th e r th e r e sis t
, ,

ance o f so me o f th e b o dy ce lls is r e duc e d Th e t e rm in

.

fe ste d has b e e n applie d to tho se co n ditio ns wh e re S o call e d -

harml e ss bact e ria a re pr e se n t in th e b o dy n o rmally Th ese .

bact e ria may e nte r th e tissue s o f th e b o dy unde r ce rtain co n

ditio ns and pr o duc e an infe cti o n Ada m i calls thi s pr o c e ss
.

subinfe ctio n .

I t has b e e n stat e d that all dis e as e s d ue to bact e ria a re i n

f e ctio us A c e rta in pr o p o rtio n, o f th e s e infe cti o us dise ase s are

.

al so co nt gi o us ; that 5 the y may b e tr ansmitte d by di r e ct co n

a 1 ,

tact Of th e infe ct e d indi vidual with h e althy indi viduals o r by ,

indir e ct co ntact thro ugh th e age ncy O f f o m ite s Co ntagi o n .

h as to d o with th e manne r o f tran s mi ssi o n o f a di s e as e and ,

i nfe cti o n with th e natur e o f th e cau s e O f th e dis e as e Fo r e

. x

ampl e smallp o x and diph th e rI a are infe ctio us co ntagi o u s d is

, ,

e as e s whil e t e tanus a nd anth r ax are pur e ly inf cti o u s di se as e s

,
e .

I nfe cti o us di se a se s may b e arbitrarily di vi de d i nto sp e cific

and no n sp e cifi c dis e ase s S p e cifi c i nfe cti o us di se ase s are tho se
-
.
2 34 B I O L O GY OF S PE CI ALI ZE D GRO UP S
which are pro duce d by a de finite bacte rium o r micro o rganism .

Th e sympt o mat o l o gy o f th e s e di se as e s is e qually sp e cific .

Th e d e fi nit e bact e rium o r micro o rga nism i s the o nl y e xcitin g

cause o f thi s class o f di s e as e s Diphthe ria typho id fe ve r .

, ,

bub o ni c plagu e a e typ e s o f sp e cific infe ctio us dise ase s Th e

,
r .

FI G .
50 .
— Tr pe o ne m a pall id um o f s yphili s in ti s s ue . William s .

eti o l o gical age nts in th e se di se ase s a re B act diphth e ria .

,

B typho sus and B p e stis and natural inf e cti o n o f a b o dy with

.
,
.
,

th e se bacte ria usually pro duce s a de fi nite patho l o gical co n

ditio n and sympto mato l o gy S o m e i nf e cti o us dise as e s in
.
,

which th e tio l o gical age nt is no t kn o wn a re call e d Spe cific

e ,

infe ctio us di se ase s Hydro pho bia smallp o x and chi ck e n

.
, ,

a re e xampl e s o f thi s cla s s

p o x .
 PATHOGENI C B ACTERI A 2 35

N on p e cific i nfe cti o us di se ase s are tho se whi ch a e no t

-
s r

cause d by any o ne de fi nite bact e rium Th e vario u s infl am .

m a to ry p r o c e sse s whi ch charact e r iz e so many di se as e s ( s e e

tabl e b e l o w) a e e xampl e s o f patho l o gical co nditi o ns whi ch are

r

no t sp e cific and may b e d ue to a vari e ty o f nn crOOrg -

a ni sm s .

F o r e xampl e M p ne um o ni a B c o li S tr p yo ge ne s and many

‘

,
.
, .
,
.
,

o th e r ba ct e ria may p r o duc e i nflammati o n in va r i o u s pa r t s

o f th e b o dy I nf e cti o ns f r e xampl e with S tr py o g e n e s

.
,
o ,
.
,

may pr o duc e d e p e ndi ng up o n th e ave nu e o f inf e cti o n di s

, ,

e as e s s uch a s e r y sip e la s pue rp e ral s e pti ce mia se ptice mia ’

, , ,

t o nsill itis abs ce sse s p e rito nitis e tc Th e se di se a se s are wide ly

, , ,
.

di ff e r e nt in th e ir sympt o mat o l o gy Th e f o ll o wi ng i nc o mpl e t e .

tabl e s will s e rve to give an o utline o f th e vario us co mmo n in

fe ctio us di se as e s

P AR TI AL LI S T OF S P E CI F IC IN F E CT I OUS DI S E AS E S DUE TO
BA T C E RI A

Di se a se E ti o l o gi ca l B acte ri um

G o norrh oe a M ir g n rrh
c o co ccu s o o oe a

C ha n r id c o s B a ill h n id m alli
c us c a cro es s

M ni ngiti
e r br — p inals , ce e o s ,
Mi r c o co ccus m ni ngitidi e s intra ce l l u
d mie c lari s

T yp h id f v ro e e B a ill us t yp h u
c os s

R lap ing f v r
e s e e S p irill um b rm i ri o e e e

M alta f v r e e Mi r m lit n i
c o co ccus e e s s

Dy nt ry ba illary
se e , c B a ill d y nt i S hig
c us se ar a a

T u b r ul i
e c os s B a t ri um t b r l i
c e u e cu o s s

G la nd r e s B a t ri m m all i
c e u e

T ta nu
e s B a ill t tani
c us e
'

I nfl ue nza B a t ri um infl u nz
c e e a
 B I O L O GY S PE CI AL I ZE D GROUP S
Di se as e E ti o l o gi ca l B acte ri um

An thra x B a t ri m a nthra i
c e u cs

L pr y
e os B a t ri m l pr
c e u e a

Di p hth ri e a B a t ri m di p hth ri
c e u e a

A iati
s h lr c c o e a M i r p ira
c os mma co

B ub ni p lagu
o c e B a ill p ti
c us es s

S wi n p lag
e ue B a ill h l ra i
c us c o e su s

M alig na nt d m a oe e B a ill u d m tism l igni

c s oe e a a

S ym p t m ati a nt h rax
o c B a ill h
c i
us c a uv a

C hi k n h l ra
c e c o e B a t ri m h l ra gallinarum
c e u c o e

B t uli m u
o s s B a ill b t li n
c us o u us

S tra ngl es Mi r q i
c o co ccus e u

P AR TI AL LI S T OF S P ECI F IC INF E CT I O US DI S E AS E S N OT DUE To

BA CT E R I A

Di se as e Eti o l o gi cal B acte ri um

ti n m y i
Ac o co s s tr p t thrix a ti n m y
S e o c o ce s

My t ma ce o S tr p t thrix m ad ur
e o a M o l ds
F a rcin d u B oe uf S tr p t thri x f
e ini u
o a rc c s

B la t m y i
s o co s s O di um id id co cc o es
Y a t
Odio m y co sis ( T h
r us h) Od i um l b i n a ca
e s s

M alaria Pla m di um m alari

s o a

Dy nt ry
se e Am ba li
oe co

T xa f v r
e s e e Pir p la m a big m inum
o s e

I nf ti u ja und i
ec o s ce Pi p la m a m m n
ro s co u e

S yp hili s T p n m p allid um
re e e a

Naga na T r yp a n m a br i o so uc

D uri n
o e T r yp a n m a q uip d i um
o so e er

M l d C ad ra
a e e s Tr yp a n m a q ui numo so e

S urra T r yp a n m a va n i o so e s

S l p ing i k n
ee s c e ss T r yp a n m a g m bi n i
o so a e e s

R t di a
a se se T r yp a n m a l wi i o so e s

K ala a z ar
-
T ryp a n ma X o so

F il aria sis Filaria a ng ui ni h minis s o s,

nd th r w rm
a o e o s
 PATHO GENI C B ACTERI A 2 37

P AR TI AL LI S T OF NON S P E CI F IC I NF U S DI S E AS E S
E CT I O AND THE

BA TC E RI A WH I C H M AY P R OD U C E T H E M

Dis e as e Eti o l o gi ca l B acte ri um

Pn e um o n ia ( P n um niti e o )
s M ic r p ne um o ni a
o co ccu s

Co n j un c tiviti s S tre p t o co ccus pyo g e ne s

Iriti s B a cill us t yp h o sus
R hi niti s B a ill
c li us co

Pharyngit i s B a ill pr t v lg ri
c us o e us u a s

L aryngiti s B a t ri m w l hii
c e u e c

Ent riti e s B a t ri m t b r l i
c e u u e cu o s s

T n illiti
o s s B a t ri m p n m ni Fri dla nd r
c e u eu o a e e

O titi s Mi r py g n var a r
c o co ccu s ot e es . u e us , e c.

Ad niti e s Mi r int
c o co ccus l l l i m ni ngiti d i
ra ce u ar s e s

P l u iti
e r s P d m na p y y n
se u o o s oc a ea

M y ardi ti oc s Mi r t t g n
c o co ccus e ra e a

Arthriti s Mi r g n rrh
c o co ccus o o oe a

End arditi oc s

O titi
s nd p ri
s a titi e os s

C y titis nd ur thriti
s a e s

N p hriti
e s

End m tri ti o e s

S al p i ngiti nd o p h it i
s a o or s S a pr g
o e ni c b ac t ri
e a in so m e co nd it i o ns

V agi niti s

Or hiti c s

M ni ngit i
e s

App ndi iti se c

P rit niti
e o s

B ur iti s s

D rm ititi
e s

N uriti
e s

M y iti os s

S t m atiti
o s

Etc .

ab o ve m e ntio ne d infl a m m atio ns may b e e ith e r acut e o r

Th e -

chro nic and may b e atte nde d by suppuratio n (pus f o rm ati o n )

2 3 8 B I O L O GY OF S PE CI AL I ZE D GROUP S
It sho uld b e r e m e mb e r e d that al l o f th e no n sp e cifi c di se ase s -

ab o v m e ntio ne d a e no t always pr o duce d by m i cro o rga ni sms

e -
r .

M e cha nical physical and ch e mical facto rs may b e r e sp o nsibl e

, ,

i n c e rtain rar e i ns tanc e s .

Th e di s e a se s ( no n sp e cifi c) tabulat e d in th e f o r e g o ing li s t -

may b e cau se d by a va rie ty O f bacte ria N o t e ve ry dise a se .

m e n i o ne d is cau e d by al l th e ba ct e r ia tabulat e d S o me
‘

s .

di se as e s a e caus e d by a la rg e vari e ty o f bact e ria ( te n sp e cie s

r

in s o m e case ) whi le o th e r m o rbid c nditio ns a e cause d by

s ,
o r
‘

o nly a fe w sp e ci e s S o m e o f th e bact e r ia m e nti o n e d als o

.

pr o duce sp e cifi c i nf e cti o ns wh e n attacki ng th e b o dy at d e finit e

p o i nt s e g M go n rr hoe a i n th e g e nito u rinary tract B
,
. . . o -

,
.

typh o su s in th i nt e sti n s e tc Th e y h o w e ve r no t unco m

e e ,
.
, ,

m o nl y pr o du ce no n sp e cifi c i nf e cti o ns -
.

P A T AL
R I LI S T OF I NF T U D A EC IO S I SE S ES OF UN KN O WN ET I OL O GY ,

SP F A DN
E CI IC N ON ,S P E
-
C F C
I I

P rt u i ( Wh p i ng gh )
e ss s oo co u

Par titi p id m i (M um p )
o s, e e c s

S p tt d f v r R ky M untain
o e e e ,
oc o

Y ll w f v r
e o e e

R h m ati f v r ( R h um ati m
eu ) c e e e s

T yp h f v r us e e

S arl t f v r ( S arlati na )
c e e e c

Ap hth u f v r ( F t nd m u th d i a )
o s e e oo a o se se

Hy dr p h bia ( R abi )
o o es

S m all p x ( V ari la ) o o

C w p x V a i nati n ( V a i na ) a m
o o , rgani cc o cc ,
s e o sm

C hi k n p ( V ari l la)
c e ox ce
 PATHOGENI C B AC TERI A 2 39

M e a sl e s R i nd r p t e es

Rub lla ( R th l in)

e o e R hi n lrm o sc e o a

D ng e (Br akb n
ue e o e fe v r)
e Sh p p ee ox

B ri b ri
e -
e Di t m p r
s e e

Tra h m a c o S wi n ry i p la
e e s e s

Th e M e ans ofr ansm i s si o n o f B acte ria fr o m O ne I nd ivi d

T

ual to Ano h e t r — .Th e t r an s mis sio n o f path o g e nic bact e r ia

and o th e r mi cro o rgani sms f ro m o ne indi vidual to ano th e r

take s place in diffe r e nt ways We wil l bri e fly call att e nti o n .

to a fe w w e ll r e c o g niz e d fa ct s in thi s co nn cti o n

-
,
e .

I t has b e e n kno wn fo r y e a r s that a pati e nt Su ff e ring fr o m

di e as e s s uch a s go no rrh oe a chanc r o ids syphi li s l e pro sy o r

s , , , ,

e xt e r nal s uppu r ati ng l e si o ns co uld tra n mit th e dis e as e by s

di re ct co nta ct with th e abrade d skin o r muco us m e mbrane o f

th e di se a se d p e r so n o r so m e a r ti cl e having b e e n in co ntact

with th e di se a se d in di vidual I n the se cas e s th e bacte ria pass

.

di r e ctly by co ntact fro m One inf e cte d w o und to ano th e r and

i nfe ct it Tub e r cul o u s l e sio ns have al so b e e n n o t e d to b e
.

d e ve l o p e d in th e ski n by dir e ct ino culati o n fro m o p e ratio ns

and ne cro p si e s p e rf o r me d o n tub e r cul o u s i ndividuals Th e n .

again the r e a re o th e r di se ase s fo r e xampl e such as scarl e t , ,

f e ve r a nd smallp o x whi ch can b e t r ansmitte d by th e age n cy

,

o f f o mi t e s ; that is a r ticl e s which have b e e n in co nta ct

,

with th e di se as e d in di vidual such as e ating ute nsils b e d , ,

cl o thi ng b o o ks e tc
, ,

I t h as b e e n r e c e ntly p o i nt e d o ut that typh o id f e v e r whi ch ,

wa s h e re to fo r e supp o s e d to b e t r ansmitt e d o nly thr o ugh th e

age ncy o f wate r whi ch has b e e n co ntaminate d by s e wage o r
2 4 0 B I OL O GY OF S PE CI ALI ZE D GR OUP S
inf e ctio us substance s co ming in co ntact with wat e r may b e ,

carrie d by n o rmal p e rso n s wh o have had an atta ck o f typh o id

f e ve r at s o me tim e pr e vio us C ase s are o n r e co rd wh e re B
. .

typho sus has b e e n ca r ri e d fo r e ighte e n y e ars afte r a n infe cti o n by

a do m e stic and the n tran smitte d by p e r so nal uncl e anlin e ss to
s e ve ral m e mb e r s o f th e fam ily in which sh e was e mpl o y e d .

B typho su s is f r e q u e ntly Ob se rv e d in th e g all bladd e r in p e r

.

s o ns having had th e di se as e fo l o ng p e rio ds F lie s may al so

r .

carry B typh o sus f ro m f e ce s to va ri o u s f o o d pr o du cts and

.
,

o th e r individuals may th e n b e inf e ct e d .

I n bub o nic plagu e th e di se a se i s tra nsmitt e d am o ng th e

rats which a e v e ry e asily i nf e cte d with B p e ti s by m e ans

,
r . s

o f th e bit e o f th e rat fl e a A fe w Ob s e rvati o ns are o n r e co r d

.

sho wing that th e s e rat fle as may al so bite and infe ct man .

F r e qu e ntly th e fl e a d e p o sit s th e bact e ria o n th e skin a nd by

scratchi ng th e o rgani sm s are in o culat e d So me di se as e p ro .
-

d ucing bact e ria a e larg e ly wa te r b o rn e such as B typho sus

r -

, . .

T h e sam e i al so tru e o f B d yse nte ia and th e M sp co mma

s . r .

o f Asiatic ch o l e ra .

I n di se as e s li k e mala r ia y e ll o w f e v e r a nd s o m e f o rms o f
, ,

fi l ari a sis m o sq uit o e s act a s th e ca r ri e r s f th e i nf e cti o n o In .

malaria th e h e mat o zo an unde rgo e s ce r tain p e rio ds Of i ts lif e

cycl e in th e mo squito I n y e ll o w f e ve r this is p ro bably th e
.

ca se also but a s th e e ti o lo gical o rgani sm is no t kn o wn we

,

canno t fo rmulat e de finite info r mati o n in thi s r e gard .

P r o t o z o an di se a se s su ch as T e xa s f e v e r African r e lap si ng
, ,

fe ve r are carri e d by m e an s o f th e ti cks a nd th e inf e ctio us

, ,
 PATHO GENI C B ACTERIA 2 4 1

o rganism is transfe rr e d in so m e case s fr o m m o the r to yo ung .

I n tr o pical pr o t o z o an di se a se s su ch a s nagana and sl e e pi ng

,

FI G .
51 . S p iro ch a ta o b rm i ri f r lap i g f v r William
e e e O e s n e e . s .

sickne s s which a e d ue to t r ypan o so me s th e tse tse fli e s are

r ,

facto rs in th e tran smissio n Of th di se a se Th e se pro to z o an e .

di se ase s a re transmitte d by bite s o f th e fli e s .

B e dbug s c o mm o n fl e a s and li ce may al so b e capabl e o f

, ,

tran smitting ce rtain i nfe cti o us di e a se s B act tub e rcul o sis s . .

has b e e n f o und in th e b e dbug I t has also b e e n state d .

r e ce ntly that th e l o use transmits th e micro o rganism o f

typhus f e ve r .

I t is al so a w e ll kn o wn fa ct that h e althy p e rs o ns may

-

Carry o n th e ir muc o us m e mbran e s such o rganisms as th e

R
2 4 2 B I OL O GY OF S PE CI AL I ZE D GR O UP S
M pn e umo nia and B act diphth e ri a witho ut pro ducing
. .

e ffe ct . The se same o rgani sm s may b e co me vi rul e nt

tak e n into a susce ptibl e indi vidual Th e o rganisms may
.

b e tran s fe rr e d fro m o ne to th e o th e r by di re ct o r indi r e ct

co ntact .
 CHAP T ER XXI I
P ATHO GE NI C B ACTE R IA (C o nti nue d)

M o de of Acti o n of te ri a t P atho g e nic

P a h o ge ni c B a c .
—

bacte ria pro duce mo rbid co ndi tio ns in th e ani mal b o dy by th e

f o llo wing ge n e ral m e tho ds F r e que ntly the re is a co mbina
.

tio n o f two o r mo re o f th e m e tho ds in o ne dise ase co nditio n

in th e b o dy .

a( ) B y m e chanicall y cl o gging th e capill ari e s o f th e bl o o d

ve sse l s thus pre ve nting th e o xyg e n and nut rime nt f ro m r e ach
,

ing th e vita l ti s su e c e lls I n anthrax whi ch is cau se d by

.
,

B act anthracis a large p o rti o n o f th e patho g e ne sis is p ro

.
,

d uce d by this m e chani cal m e th o d M yc o tic e mb o li a re O ft e n

.

fo rm e d in th e cap illari e s .

( b) B y th e p ro ductio n O f so lubl e to xins o r p o is o ns by th e

actio n o f bacte ria inside th e b o dy whi ch wh e n ab so rb e d by ,

th e bl oo d lymphatic s o r ne rv e tissu e and ca r ri e d to th e

, ,

vari o us c e ll s o f th e b o dy pro duc e mo rbid change s withi n

th e m I n diphth e ria fo r e xampl e th e to xins ge n e rate d by
.
, ,

B act diphthe ri a are r e sp o n sibl e fo r th e dise ase

. .

(6) B y th e d e s tructi o n o f ti ssue ce ll s at th e fo cus Of th e

'

inf e ctio n thu s incapa citati ng th e se c e ll s fr o m p e rf o r ming th e ir

,

n o rmal functio ns I n tub e rculo si s th e de structio n Of ti ssu e

.
,

by B act tub e rcul o sis is o ne imp o rtant part Of th e dis e ase

.

2 43
2 44 B I OL O GY OF S PE CI AL I ZE D GROUP S
pr o ce ss P ro gre ssive gangr e ne is also d ue in part to th e
.

de structi o n o f tissue b y bacte ria

‘

Dis e as e d c o nditi o ns O f thi s natur e may als o b e pr o duc e d in

th e b o dy by bact e ria which cann o t di r e ctly b e call e d path o

g e ni c .

(d) B y th e pro ducti o n o f p o i so ns o r to xi ns by sapr o g e nic

bact e ria acti ng o n fo o d p ro du ct s o utside th e b o dy Wh e n .

take n into th e b o dy thro ugh th e dige s tive t r act and ab so rb e d ,

th e s e p o iso no us sub stanc e s pr e se nt give ris e to s e ri o us in

t o xi cati o ns S imilar m o rbid co nditio ns may ari se o n acco unt
.

o f th e acti o n o f sapr o g e nic ba ct e r ia withi n th e int e s tin e s .

Th e p o i so n o us sub s tanc e s pr o du ce d a e call e d pt o mains r .

M e at p o i so ning milk and ice c r e am p o is o ni ng a e e xampl e s

, ,
r

o f di se as e d c o nditi o ns ari i ng fr o m th e acti o n o f pt o mains

s .

O ccasi o nally aft e r parturiti o n a p o r ti o n o f th e plac e nta is

,

r e tain e d with i n th e ute rus and it may b e act e d up o n by

,

sapro ge nic bact e ria P to mai ns ge n e rat d and abso rb e d in

. e

thi s r e gio n u sually pr o duce pro no unce d into icatio ns x .

Two principal fact o r s a re invo lve d in th e pr o ducti o n O f a di s

e as e by bact e ria ( )
I T h e e ntra n c e O f bact e ria i nt o th e

tissue s o f th e b o dy and the ir multiplicatio n Thi s facto r it .

,

will b e re call e d co nstitute s i nfe cti o n 2 ) Th e e lab o rati o n o f

,
.

p o iso no us pro duct s by th bact e ria which may act o n th e tis

e

sue s imm e di ate ly sur r o undi ng th e m o r o n th e e ntir e b o dy .

Th e latt e r fact o r c o nstitut e s what is k no wn as p o i so ning o r

i nto i ca ti n
x oI n m o s t m o rbid c o n di ti o ns b o th O f th e se fact o rs
.

a re c o nce r ne d I n o nly a fe w cas e s i s o ne fact o r al o n e i nvo lve d

. .
 P ATHO GENI C B AC TERI A 2 45

I n th e case o f th e inf e ctio n Of th e b o dy and th e multiplica

tio n within th e b o dy o f B act anth ra ci s th e path o ge nic
.
,

e ff e cts a re pr o bably pri ncipally m e cha nical and d ue in th e

main to th e rapid mul tipli catio n o f th e bacte ria I n thi s case .

the re is al so so me fo r m Of to xin p ro duc e d by th e bacte rial c e lls .

Th e pr o ducts o f th e m e tab o li s m o f th e path o g e nic bact e ria

will re c e ive co nside ratio n late r .

P o o f o f Eti o l o gi al R e l a ti o nsh ip
r c . I n o rd e r to pr o ve that

a de finite micro o rgani sm is th e e tio lo gical facto r o f a sp e cific

'

di se a se R o b e r t K o ch d e vi se d th e f o ll o wing p o stulat e s o r
,

rul e s whi ch if fulfill e d e stabli sh e s th e co nn e ctio n b e twe e n a

micr o o rgani sm and a di e ase
s .

( a) Th e m icro o rgani sm mu s t b e fo und in th e tissu e s o r

b o dy o f th e animal o r plant with th e di se as e .

( b) Th e micr o o rgani m mu t b e i so lat e d in pur e culture

s s

fro m th e b o dy o f th e di se as e d ani mal o r plant .

( c) I t mu s t b e gr o wn succ e s sfully o n culture m e dia .

(d) I t must b e abl e to pro duc e th e di se a se in a h e althy

animal o r plant in appr o ximat e ly th e sam e fo rm and giving
th e sam e sym pt o mat o l o gy as in ( a ) I t must als o again b e
.

f o und in th e ti ssu e s o f th e e xp e rime ntal animal o r plant .

T h e maj o rity O f path o g e nic bact ria fulfill th e se laws but

e ,

th e re a e s o me micro o rgani sms whi ch undo ubte dly pro duce

r

sp e cific inf e cti o u s di se a se s which d o no t fulfill th e p o stul at e s

O f K o ch .Th e f o ll o wi ng tabl e giv e s a partial li s t o f th e mic ro

o rgani s m s whi ch pr o duc e S p e cific i nf e cti o us di se as e s but whi ch

are unabl e to f ul fill K o ch s laws

’
2 46 B I O L O GY OF S PE CI AL I ZED GR OUP S
B acte rium l e pra (Le pr o sy) .

Tr e p o n e ma pallidum ( S yphi li s) .

Spiro ch a ta o b e rm e i e ri ( R e lapsing f e ve r) .

Am oe b i c o li (Am oe bic dy se nt e r y) .

P lasm o di um malari a ( M alaria) .

Th e re are o th e r bacte ria a s fo r e xampl e th e M go no r

, , ,
.

rh oe a which f ul fill all but o ne o r two p o stulat e

,
I n thi cas e
s . s

M g o n o rrh oe a will no t pro duc e th e di se as e wh e n i no cul at e d

.

into animals Th e r e are stil l o th e r bacte ria which have b e e n

.

pro ve d to b e th e e tio l o gical age nts in di s e ase s which d o no t

c o mply with all th e laws Th e c o n s tant ass o ciatio n o f d e fi
.

nite micr o o rganisms with dise as e s S ho wing th e sam e sym pto m

a to l o gy has l e d to th e s e o rga ni s m s b e ing r e gard e d s sp e cific
a .

Eff e cts o f P a th o ge ni c B a cte ri a o n th e B o d y Th e fo l .

l o wing tabular o utline is give n by M uir and R itchi e and po r

tray s in a ve ry g e ne ral way th e actio n o f th e patho g e ni c bac
t e r mo n th e animal b o dy .

A . Tissu e C hanges .

( ) Lo cal chang e s I n r e gi o n o f th e bact e ria

1 . .

P o siti o n ( a) At primary l e si o n .

( b) At s e co ndary fo ci .

C haract e r ( a) Ti ssu e re acti o ns .

( b) De ge n e r ati o n acute o r chro ni c

, .

( 2 ) C hang e s pr o duc e d at a di s ta nce fr o m th e bact e ria

dire ctly by th e ab so rptio n o f to xins .

( a) I n sp e cial tissue s
 P ATH OGENI C BACTERI A 2 47 .

( ) a r
1a e sult
As o f damag n e rve ce lls a nd
e —

fib e rs se cre ting ce ll s v e sse l wall s o r

, , ,

( )
2 a C ha ng e s o f a r e activ e natur e in th e bl o o d
f o rmi ng o rgans ( B o n e s sple e n , ,

( )
b G e n e ral anat o mical c hang e s d u e to m al n u

tri ti o n o r incr e a se d wast e .

B Ch ange s in M e tab o lism

. .

O ccu rr e n ce o f fe ve r e rr o rs o f assimilati o n and

,

e li minati o n .

Th e e ff e cts
bacte rial acti o n in th e b o dy bri e fly state d
o f , ,

a re as fo ll o w s ( )
a D e g e ne rati o n a nd n e cr o s is (b) r e ge n
e ra tio n r e activ e and de f e n siv e chang e s
,
De g e n e rati o n a nd
.

n e cro sis a re d ue to th e susce ptibility and vul ne rabili ty Of th e

tissue s I n th e latte r pro ce s s ( b) in all p ro bability th e pha
.

go cyte s and th e b o dy flui ds are b o th r e sp o nsibl e .

Th e p o siti o n o f th e l e si o ns in th e bo dy is no t e a sily e x

plain e d by any m e chanical m e an s With ut do ubt th e . o

mi c r o o rga ni s m s o r th e ir t o xi ns and sp e cial va ri e ti e s o f tis su e

’

ce ll s e nte r into che mical co mbinatio n with e ach o the r The r e .

is a se l e ctive actio n o n th e part o f th e bacte ria o r the ir to xins

fo r th e se sp e cial tissu e s Th e afli ni ty O f th e t e tan o spas m in
.
-

O f th e t e tanu s t o xin fo r n e rve c e ll s i s a cas e in p o int .

I n acut e i nf e ctio ns by ba cte ria th e r e is u sually a l o cu s o f in

fl am m atio n at th e fo cus Of th e infe cti o n He mo rrhag e oe d e ma

.
, ,

and in s me case s s uppura tio n may b e atte ndant co nditio ns

o .

I n chr o ni c inf e cti o ns as fo r e xampl e th e inf e ctiv e g r anul o

, , ,
 4 8 B I OL O GY OF S PE C I ALI ZE D GR OUP S
mata (glande rs tub e rcul o sis acti no myco sis syphili s and
, , , ,

l e pro sy) the re is o nly a small am o unt o f vascular change but

,

a co mparative ly larg e amo unt o f tis sue hyp e rplasia This .

hyp e rplasia (incr e ase in ti ss ue ce lls) is principally n o te d in th e

co n ne ctive ti ssu e s .

Th e g e ne ral l e si o n s in th e b o dy d ue to th e t o xi ns o f bac

t e ria are usually tho se o f cl o udy swe lling and so me tim e ,

s,

fatty de ge ne ratio n o f par e nchymato us o rgans such a s th e ,

live r kidne y e tc C apillary h e mo rrhage s d ue to hyali ne

, ,
.

chang e s in th e a r t e rial capil lari e s are al so n o te d .

M ark e d chang e s in th e m e tab o li sm o f th e b o dy a re n o t e d

in bact e rial infe ctio ns F e ve r may b e d ue e ithe r to th e d e
.

structio n o f tissue o r th e fo rmatio n o f antib o di e s e tc P a ,

.

ral ysis c o ma spa s m s a re no t unc o mm o n in inf e cti o ns wh e r e

, , ,

th e n e rv o us syst e m i s mark e dly a ff e ct e d . Di s turbanc e s o f th e

s e cr e to ry functio ns o f th e vario us glands o f th e b o dy a re also
n o te d .

Th e P o i s o no u s P r d u cts o f B acte ri a
o . Th e p o i so n o us

ch e mical substanc e s pro duce d by bacte ria may b e gr o up e d

int o f o ur classe s as f o ll o ws
( )
a P r o ducts o f th e d e c o mp o siti o n o f th e m e di a o n wh i ch
th e bact e ria a re gr o wi ng P to m ai ns and s o m e o th e r b o di e s a re
.

pro duce d in this way .

( )
b S o lubl e p o is o ns synth
,
e tically p r o duc e d by th e bact e rial
ce lls and s e cr e te d by the m int o th e surr o undi ng m e dia Th e .

true to xi ns are pro duce d in thi s way .

()
c P o i s o n o us pr o ducts sy n th e tically pr o duc e d by th e bac
 PATH O GENI C B ACTERI A 2 49

t e ria which a re no t se c r e t e d o utside th e ce ll wall The se .

p o i so n a re sp e cific sub stance s On a cco unt o f b e ing intra

s .

c e llula r th e se t o xic sub stance s a re call e d e nd o to xi ns

,
.

( )
d P o is o no u s pr o t e in s which a e c o ns titu e nt
r s o f th e
p o r

to p l asm o f th e bact e rial c e ll Th e se pro t e ins are no t in any

.

se nse so lubl e o r sp e cific and are no t de finit e ly kn o wn to b e

r e sp o n sibl e fo any m o rbid co nditio n in th e b o dy Th e y may
r .

b e c o nce r ne d in tub e rcul o sis The s e p o iso no us pro ducts are

.

call e d t xi c bacte i al p o te i ns
o r r .

P to m ai ns . Th e s e sub stanc e s w e r e th e fi r st bact e rial pro d

ucts to b e r e c o g ni z e d Th e y w e r e a ssume d to b e alkal o idal
.

in natu r e and o n a cc o u nt o f th e fact that p o is o no u s plants o we

,

th e ir e ffe cts to alkal o idal sub stance s so m e o b se rve rs w e re in ,

cl ine d to b e li e v e that pt o main s w e r e r e sp o nsibl e fo r dis e as e .

P t o mains are ba sic ni tr o g e n o u s s ub s ta nc e s a nd a re f o und in

th e m e di a o n which c e r tai n sp e ci e s O f bact e ria a e gr o wi ng r .

I t wa s s o o n d e t e rmi ne d that pt o mai ns f r o m path o g e nic bac

t e ria w e r e no t alo ne abl e to pr o duce th e m o rbid e ff e ct s n o te d

in e xp e rim e ntal animals Th e m o s t hi ghl y p o i so no us pt o mains
.

a e pr o duc e d by th e S O call e d no n path o g e ni c bact e r ia ( s ap

r - -

ro ge nic bact e r ia) O nl y a s mall amou nt o f r e se arch w o rk has

.

b e e n do n e o n th e subj e ct o f pto mains duri ng re ce nt y e ars .

P t o main s are f o rm e d by th e a cti o n O f vari o u s sp e ci e s o f

sap ro ge nic bact e ria n p ro te in mate rial Th e y o we th e ir

o .

basic nit ro ge no us cha r act r to th e a m i no gro up s which th e y

e

co ntain I t h a s b e e n de te rmin e d that pto mains may b e p o

. r

d uce d by th e acti o n o f s apro g ni c ba ct e ria o n d e ad path o g e ni c

e
2 5 0 B I OL OGY OF S PE CI AL I ZE D GR OUP S
bacte ria o r o th e r bacte ria I t is no t kn o wn whe th e r o r no t
,
.

any pto mains a re p ro duce d by intra—ce llular bacte rial pro ce sse s .

Th e pt o mains are cl o se ly r e lat e d to th e amin o acids a nd We lls ,

state s that in all pro babil ity the y are pro duce d by s e co nda ry
chang e s (e nzym e actio n) o n th e s e acids P to main s are th e m.

s e lve s susce ptible to th e actio n Of sapro ge nic and sap ro phi l ic

bacte ria Th e y are n o te d to de cr e ase in amo unt in so me cul
.

tur e s and thi s fact is acco unt e d fo r by r e as o n o f th e ir susce p ti

bil ity to th e acti o n o f o th e r bact e ria P to mains a re distinct
.

fro m t o xins as will b e p o int e d o ut lat e r

,
.

P to mains o nly p ro duc e dis e as e in th e animal bo dy wh e n

th e y a re intr o duc e d in th e fo o d pro ducts in which th e y have
b e e n fo rm e d by th e pro ce ss o f d e co mp o siti o n The y a re also
,
.

pr o duce d in th e b o dy by th e r e t e nti o n and th e d e co mp o siti o n

o f pr o t e in mat e rial such as a p o r ti o n o f th e plac e nta in th e
,

ute rus afte r partu ritio n Th e to xic co nditi o n r e sulti ng fro m

.

th e abs o rpti o n o f pt o mains is kn o wn as sapr e mia .

Th e r e are many di ff e re nt pto mai ns which have b e e n iso

lat e d divid e d into gro ups and name d Al l th e pto mains
, ,
.

have ve ry c o mpl e x fo rmulas Th e re is o ne gro up Of pto mains

. .

which is o f Sp e cial i nte re st and this is th e ch o lin gro up This

,
.

gr o up will b e brie fly co nside r e d Th e gro up include s ch o lin

.

and r e late d substance s as fo llo ws

C HgOH C H2 N ( C H3 ) 3 OH .

C H2 = CH —N ( C H3 ) 3 — OH .

C H( OH) 2 — C H2 — N C H3 ) 3 —O H
( .

OOO H C H2 — —
N ( C H5 ) 3 OH .
 PATH OGENI C B ACTERI A 2 5 1

C h o lin is a substance which is pr e se nt n o rmallyin all ce lls .

I t f o rm s th e nitr o g e n p o rti o n o f th e l e cithin m o l e cul e Th e .

so urc e o f thi s pt o main i n d e c o mp o sing pr o t e in mat e rial is

the r e fo re e vide nt I t is sta t e d by va rio u s write r s th at it is

.

bro k e n do wn in th e anI m al b o dy e sp e cially wh e n n e rvo us tis

s ue,
whi ch is rich in l e cithin is di sint e grat e d Ch o li n is said
,
.

to b e p r e se nt i n th e ce r e br o spi nal fluid and in th e bl o o d in

-

ce rtain d e g e ne rative n e rvo u s dise as e s F o r e xampl e it has .

,

b e e n r e p o rte d in th e bl o o d in ce re bral syphi lis e pil e psy and , ,

de m e ntia paralytica C h o lin is o nl y: slightly to xic but th e

.
,

n e xt m e mb e r o f th e gr o up n e urin int o whi ch ch o lin can b e

, ,

tran sfo rm e d is v e ry t o xic We lls sugge sts that into xicatio ns

,
.

in th e gastr o int e stinal tract whi ch a re O ft e n ascrib e d to

-
fo o d
into xi catio ns may b e d ue to ch o lin b e ing split o ff fro m th e
”

l e cithi n o f th e f o o d by th e acti o n o f sapr o g e nic bacte ria and

th e ch o lin thus f o rm e d b e i ng cha ng e d int o n e urin which wh e n ,

ab so rb e d give s rise to int o xicatio n

,
.

C h o lin a nd n e urin a e cl o s e ly r e lat e d in structur e and physi

r

o l o gi cal acti o n to muscarin an alkal o idal sub stance firs t

,

de rive d f ro m mu sh ro o m s A pto main ide ntical with musca rin

.

h as b e e n is o lat e d fr o m d e c o mp o sing fi sh a nd th e nam e mus ,

ca rin h as also b e e n applie d to it N e uri n a nd muscarin are

.

v e ry p o iso n o u s pto mains B e tai n th e las t m e mb e r o f th e

.
,

gro up is o nly slightly to xi c I t is al so d e rive d fo r th e mo st

, .
,

part fro m th e l e cithi n o f th e ce ll s

, .

To xins . As h as b e e n s tat e d ce r tai n bact e rial c e lls hav e

,

th e p o w e r O f se c r e ti ng p o i so n o u s sy nth e ti c p r o ducts int o th e

2 5 2 B I OL OGY OF S PE CIAL I ZE D GR OUP S
surr o unding m e dia Th e s e substanc e s are call e d s o lubl e t o xins
.

o r t o xin s Th e r e a e v e ry fe w bacte ria which have th e p o w e r

. r

O f p r o duci ng th e se to xi ns B act diphthe ria B t e tani . .

,
.
,

P S p yo cya ne a and B b o tuli nus a re p r actically th e o nly

.
,
.

bact e ria which pro duce to xi ns Of thi s cha r a ct e r with th e p o s ,

sibl e e xc e pti o n o f so m e sp e ci e s whi ch p ro duc e t o xi n lik e h e -

m o l yticp r o du ct s Th e se sub stanc e s are r e f e rr e d to b e l o w

. .

C e rtain animal s s u ch as snak e s e e l s lizard s sc o rpi o ns e tc

, , , , ,
.
,

by activity o f a particular gr o up o f gland c e ll s pr o duc e s o lubl e

t o xi ns v e n o ms e tc I mmunizati o n with th e se animal to xins
, ,
.

pro duc e s antib o di e s I n so m e t o xins a s fo r e xampl e t e ta nus

.
, , . ,

to xi n two di stin ct to xi ns ca n b e se pa r at e d t e tano spa smi n

, ,

whi ch h s a n affi nity fo r ne r ve tis su e and t e tano lysi n (h e mo

a ,

lysi n) wh ich de s t ro y s th e e d bl o o d c e ll s I n diphth e ria in all r .

,

p r o bab ility th e re a e al s two t xi ns g e ne r at e d th e princi

,
r o o

pal to xi n to which m o t o f th e diphth e ritic i nfe ctio n is d ue and

,
s ,

ano th e r t o xic sub stance whi ch is f o rm e d sl o wly by th e bac ,

t e ria and call e d by Ehrli ch a to xo n Th e paraly sis n o te d lat e .

i n s o m e ca se s O f diphth e r ia i s a s crib e d to th e se to o ns x .

Th e ch e mical c o n s tituti o n o f t o xins is u nkn o wn A gr e at .

amo unt o f r e s e arch w o rk has b e e n do ne and is b e ing do ne at

th e pr e s e nt tim e al o ng thi s li ne T o xi ns a e v e ry labil e sub . r

stance s and a re e a ily d e stro y e d by co mparativ e ly l o w m o i st

s

he at light ch e mi cals and by o xyge n e v e n in th e amo unt fo und

, , , ,

in th e air Al l o xidizing ag e nt s including th e e nzym e s O f th e

.
,

b o dy a nd o f bact e ria de str o y to xi ns Th e y are no t affe cte d

,
.

by chl o r o fo r m to luo l phe no l e tc and are pr e cipitate d by

, , ,
.
,
 PATHOGE NI C BAC TERI A 2 53

th e maj o rity o f th e h e avy m e tals On acco unt o f the ir ma rke d

.

lab ility ch e mical analy se s cann o t b e mad e with accuracy I t

,
.

is p o ssible that as physical ch e m i stry is de ve l o p e d m o re in ,

f o rmatio n will b e gain e d in r e gard to th e se substance s I n .

v e s tiga tio ns a re b e ing ca r ri e d o n at th e pr e s e nt tim e by s o m e

o f th e no t e d phy si cal ch e mist s M any fact s have b e e n l e ar ne d

.

ab o ut t o xin s by m e an s o f animal e xp e rim e ntatio n .

I t is h o w e ve r e stab l ish e d that t o xins a re no t tru e pr o t e ins

, ,

as th e y give no ne o f th e r e actio ns fo r the s e sub stance s The y .

s e e m to b e m o r e clo se ly r e lat e d in the ir vari o us r e actio ns to

th e e nzym e s F o r e xampl e it is imp o ssibl e to i so late th e m
.
,

pu r e and th e y are no t lik e pro te ins but se e m like e nzym e s to

,

b e co l l o di al substa nc e s o f hi gh m o l e c ul ar c o mp o siti o n B o th .

to xi ns and e nzyme s pass th r o ugh p o r ce lain filte rs but have ,

th e ir st r e ngth much r e duce d Th e y rar e ly pass a di alyzing

.

m e mb r ane Th e y a re b o th d e str o y e d by 8 C m o ist he at

. 0
° '

.
,

but will with stand 00 C dry h e at and l o w t e mp e r ature s

1
°
.

fo r s o m e tim e Th e y r apidly de te rio rate o n standing and thi s

.
,

se e ms to b e d ue to th e de structio n o f o ne part o f th e e nzyme

o r t o xin m o l e cul e T h e t o xin and e nzym e m o l e cul e s are
.

made up o f two distinct ch e mical parts o g ro ups One r .

part co mbine s with th e sub s tance attacke d (hapt o pho r e ) and ,

th o th e r p o r ti o n pr o duc e s th e t o xi c o r f e rm e ntive e ff e cts and

e

th e d e s tructi o n o f th e substanc e ( to x o ph o r e o r zym o ph o r e

gro up) .

Th e part o f th e t o xi n o r e nzym e m o l e cul e whi ch is aff e ct e d

b y th e vari o us ag e nts is th e t o x o ph o re o r zym o ph o r e gro up ,

2 54 B I O L O GY OF S PE CI AL I ZE D GR O UP S
and th em o l e cul e s thus b e co me what are call e d to xo ids o r fe r
m e nto id s E nzym e s act e qually w e ll o n lib e ratio n aft e r b e i ng
.

co mbine d with substance s o n which th e y have acte d whi le ,

to xi ns di ff e r in that th e y are r e nd e r e d in e rt afte r co m bin

ing with any sub stance An e nzym e can th e re fo r e as far as
.
, ,

a m o unt go e s pr o du ce m o r e e ff e ct tha n th e t o xi n B o th t o xin

°

.
,

and e nzym e are to xic o r p o i so no us to animal s The ir inj e ctio n .

pro duc e s a r e actio n in animals a nd c e rtain antib o di e s are p ro

d uce d Th e b o di e s a re c o n sid e r e d b e l o w
. .

I t can b e r e adily s e e n f r o m th e fo r e g o i ng that th e r e a re

m a ny p o ints o f r e se mblanc e b e tw e e n th e nzym e and th e t o xin

e .

To xi ns di ff e r fro m pto mains in t hat th e y are sp e cific sub

stance s pr o duce d by d e finite ba cte ria whi l e o ne di stinct p to
,

main can b e pro duce d by th e acti o n o f s e ve ral sp e ci e s o f bac

t e ria Antib o di e s can b e pr o duc e d again st t o xins but no t
.
,

agains t pto mains To xi ns a e s e cr e tio n s o f th e bact e rial c e lls

. r ,

whi l e pto mains are p r o duc e d by cl e avage o f th e m e di a o n whi ch

th e bact e r ia a re g r o wing P to mains a e c o mparative ly s tabl e
. r

substance s and can b e subj e ct e d to ch e mi cal analysis To xins .

a re v e ry labil e and can no t b e analyz e d ch e mically .

l E n d o to xi ns T h e maj o rity o f bact e ria pr o duc e e nd o t o xin s

.
,

and it is to th e se p o is o no u s ch e mical sub stance s that th e

maj o rity o f dis e as e s a e dir e ctly d ue I n all pro bability
r .
,

th e e nd o t o xin i s a s e cr e ti o n o f th e ba ct e rial pr o t o pla s m

withi n th e c e ll wall I t is no t thro wn o ut thr o ugh th e wall into

.

th e su r r o un di ng m e dia lik e th e s o lubl e t o xins but i s sto r e d up ,

within th e ce ll The se e ndo to xins are lib e rat e d by th e actio n

.
 PATH OGENI C B ACT ERI A 2 55

of th e nzym e s in th e b o dy which the y attack o r by auto lysis

e ,

( s e lf
-
d e s t r ucti o n by e nzym e s) o f th e bact e ria th e m se lv e s T h e .

va ri o us sp e ci e s o f pyo g e nic (pus pro ducing) mic ro co cci M -

, .

pn e umo nia B typho su s M sp co mma and many o th e r bacte ria

, ,
.
,
.
,

p r o duce e ndo t o xins Th e filtrate (p o rce lain filt e r) o f cultu re s

. .

O f th e se bact e ria i s o nly slightly t o xic Th e b o di e s Of th e .

bacte rial ce ll s a re e xtre m e ly to xi c whe n inj e cte d i nto animals ,

pro ving that th e p o i so no u s sub stance is co ntaine d withi n th e

c e ll wall Endo to xin s can b e e xtracte d by pre ssing o ut th e
.

int r a ce ll ular sub stan ce s by m e ans Of a high p o we r hyd raulic

-

pr e s s o r by g rinding th e bacte ria afte r the y have b e e n re n

,

de re d b rittl e by liquid air I mmuni zatio n Of animals with .

bact e ria pro ducing e ndo to xins yie lds in th e b o dy fluids Of th e

-

animal i nj e cte d no t antito xin s but in cr e ase d bacte ricidal ,

s ub s tanc e s agglutinins pr e cipiti ns O ps o ni ns e tc

, ,
Th e failu r e
, ,
.

to s e cur e a ntib o di e which will co mbat th e acti o n o f th e e nd o

s

t o xins dire ctly (a ntie nd o to ins) h a s s e rio u sly i m pe d e d th e

x

p ro g re ss o f se rum th e r apy .

Th e f o ll o wi ng t o xic sub s tanc e s a re pr o duc e d by s o m e o f th e

highe r plants The y a e no t unlike bacte rial e ndo to xi ns

. r

e xc e p t that a ntib o di e s may b e p r o duc e d against th e m by

immunizati o n The y are call e d phyto to xins

. .

R icin fro m th e se e ds o f R icinus co mmunis ( casto r Oil b e an) .

Ab r i n fr o m th e s e e d s o f Abrus p re cato i us (j e qu e rity b e a n) r .

C r o tin f r o m th e s e e ds o f C ro ti n tiglium .

R o bin f r o m th e l e av e s a nd bark o f R o bina ps e ud o acacia

(l o cust) .
 CHAPT ER XXI II
P ATHO GENI C B ACTER I A ( C o nti nue d)

r
To xi c B a cte i al P ro te ns i .h as b e e n f o und that aft e r
—I t

bact e ria hav e b e e n wash e d fre e o f all so lubl e t o xin and have
b e e n fr e e d o f all e n do to xin that th e r e maining c e ll wall and ce ll
,

pro to plasm p o sse sse s s o me t o xic pro p e rti e s Wh e n inj e cte d .

into an e xp e rime ntal animal th e s e pro ducts pro duce infl amm a
tio n n e cro sis and p us f o rmatio n Th e e ffe cts are no t in any
, ,
.

s e ns e sp e cific a nd inj e ctio n in diffe r e nt a nimals may pro duce

,

diff e re nt r e sults Th e to xic substance s Of thi s nature are no t

.

co nfin e d to patho ge nic ba ct e ria al o n e but it is no t infre que nt

,

to find that no n patho g e ni c bact e ria w hi ch hav e b e e n tr e at e d

-

as m e ntio n e d ab o ve p o s se ss th e sam e pro p e rti e s (B p r o digi o

, .

sus ,
Th e se to xic substance s a e s e pa r ate and di stinct
r

fro m e ndo to xin s and s o lubl e to xi ns as is sho wn in th e fact

that th e y r e sist a t e mp e rature o f C fo r te n mi nut e s wh e n
1 10 .

in o ne p e r c e nt sulphuric acid s o lutio n Th e bact e rial p ro .

t e ins can b e iso late d in this way Th e substance s are un .

d o ubt e dly pr o t e in in natu r e and in all pro bability s o m e fo rm

O f nucl e o p r o t e in
-
. We lls thi nks that th e y may p o ssibly b e
d e rive d fro m th e e ndo to xin s by so m e indire ct ch e mical pro ce ss .

Th e r e are s o m e dise a se s n o tably tub e rcul o sis and anth r ax

, ,

l e avi ng o ut o f c o n sid e r atio n th e m e chanical e ffe ct o f th e

bacte ria in th e latte r cas e which d o no t se e m to b e d ue to any
,

2 56
 PATHOGENI C B ACTERI A 2 57

s o lubl e to xin s o r any e ndo to xi ns whi ch th e r e sp e cti ve o rgan

isms might p ro duce I t h as b e e n sugg e st e d that the re may b e
.

s o m e r e latio n b e tw e e n th e i r to xi c e ff e ct s a nd th e pro te i n sub

sta nce s whi ch a e c o ntain e d within th e i r ce ll pro to plasm
r .

Anaph yl axi s to B acte ri al P r te i ns — I t has b e e n sh o wn

o .

by va rio us wo rke rs that an animal b o dy may b e made to b e

co m e hyp e rse nsitive to m o s t all p ro t in s includi ng th o se e ,

d e rive d fro m bact e ria Th e co nditio n Of hyp e rsusc e ptibility

.

to p r o t e i ns is d e sig nat e d as a naphyl xi s ( Gr again s t p o te c

a . r

tio n) I f a n ani mal r ce ive a s m ll inj e cti n o f a de finit e

. e s a o

p ro te in l ike e gg whi te ho r se s r um e tc and the n again afte r

, ,
e ,
.
,

at l e ast 8 to 3 days r c e ive a so me what la rg e r inj e cti n o f th e

1 e s o

same p ro te in th e animal u sually die s o is se rio u sly into xicate d

,
r .

Th e sam e sympt o m s r e sult wh e n bact e rial p r o t e i ns a re us e d .

The r e is a p o ssibility that an in di vidual may b e h yp e se nsi r

ti e d to a bact e rial pr o t e i n and lat e r wh e n i nf e ct e d with th e

z ,

h o m o lo go us bacte ria in larg e r numb e r may Offe r no r e sistance

whate ve r Anaphylaxis may b e tran smitte d fro m m o th e r to
.

y o ung i n ut o I n thi s way th e individual may se e m to b e

er .

natu r ally hyp e rse n sitive to th e pr o t e in c o n ce rne d .

B a ct ri al H e m o l ys ins
e I t is a w e ll e s tabli sh e d fa ct that
.
-

many bact e ria patho ge nic and no n path o ge ni c pro duce to xic
,
-

sub stance s whi ch de t ro y re d bl o o d c rp uscl e s Th e filtrat e s

s o .

o f c e rtai n bact e r ial cultur e s such as M py o g e ne s va r s au r e u s

,
. .

and albus S t pyo ge ne s B typho su s P S p yo cya ne a e tc

,
r .
,
.
,
.
,
.
,

c nta in th e se h e m o lyti c sub tanc e s B acte rial he m ly sins

o s . o

a re cl o s e ly r e lat e d to th e t o xic bact e rial pr o t e ins but a e p e r ,

r ,

8
2 5 8 B I O L O GY OF S PE CI AL I ZED GR OUP S
hap s m o r e cl o se ly r e late d to th e so lubl e t o xins The y can
, .
,

fo r all practical purp o s e s b e r e gard e d as m o difi e d t o xins

,

which have a sp e cial afli nity fo r re d blo o d co rpuscl e s Th e se .

h e m o lysi ns are di vide d i nto di sti nct ch e mical mo l e c ul e s a ,

co mp l e m e nt o r th e rm o labil e sub s ta nc e a nd an a mbo ce p to

,
or r

th e rm o stabl e substa nce as have b e e n n o te d in th e bact e ricidal

,

substanc e s in th e s e rum .

He m o lysi ns a re c o ntain e d in th e s e cr e ti o ns o r e xtracts o f

animal and plant ce lls such as snake ve no m e e l s e rums a nd
, , ,

th e vari o us phyt o t o xi ns ab o ve m e nti o n e d Th e m e cha nism

.

o f h e m o ly si s ( bl o o d de structi o n) is v e ry simi lar to bact e r i o lysis

( bact e rial di s tracti o n) .

Anti to xi ns . P ractically all c e lls O f th e b o dy p o ss e ss

physi o l o gi cal r e ce pt o rs ( side chain s) co mp o se d o f de fi nite

,

che mical radical s co n ne ct e d with a ce nt r al gro up o f ato m s o r

m o l e cul e s Th e s e side chains hav e o p e n val e nci e s and a e
. r

capabl e o f uniting with va rio us fo re ign sub stanc e s such as ,

fo o d p o iso ns e tc Th e classical e xampl e us e d to de mo nstrate

, ,
.

a ce ntral g ro up a nd its sid e chains is th e b e nze n e ring and

th e substanc e s wh ich may b e f o r m e d fro m it :

H GO O H

\ \
C C
,
f ,
f
H C c H H— C C — H

H — c C — H H— c C— H
\
v7
C

H
B nz at
e o e of So da
 PATH O GE N I C B ACTERI A 2 59

Th e ab o ve graphi c re pr e se ntatio n sh o ws th e b e nzo l m o le

cul e having a c e ntral g ro up o f C with late ral side chain s o f H
6

c o nn e cti ng with e ach ato m o f C Wh e n o ne o f the se side

.

chai ns o f H is re place d by C OOH b e nzo l is co nve rte d into

,

b e nzo ic acid I f to this acid radi cal N a is unite d supplanting

.
,

an H in th e OOH o f thi s radical so di um b e nzo ate is p ro duce d

, .

Whe n to xins are ge ne rate d in th e b o dy by path g e nic bac o

t e ria o r wh e n th e y a re i nt ro duce d artificially fo r th e purp o se

o f pr o duci ng active immu nity th e to xin m o l e cul e c o mbi ne s
,

with th e vari o us S id e chains Of th e ce ll s Th e r e is fi r st a che mi

.

cal co mbinatio n b e twe e n that che mical gro up o f th e to xin

m o l e cul e whi ch is kn o wn as th e hapto pho r e with a co rre
, ,

spo ndi ng gr o up ( S id e chai n) i n th e ce ll kn o w n as th e hapt o

phil e L at e r th e so call e d ch e mical to xo pho r e gro up O f th e

.
-

to xi n mo le cul e unite s with th e co rre spo nding g ro up (S ide chain)

in th e ce ll call e d a to xo phi l e gro up Th e unio n Of th e to xin
.

with th e ce ll s is a de fi nite ch e mical unio n which is stabl e in

so me cas e s ( t e tanu s) and no t so stabl e in o the rs ( diphth e ria) .

Th e c e ll is u sually stimulat e d o nly wh e n th e hapto ph o r e gro up

unite s with it and is injur e d o nl y wh e n in addi tio n th e to xo

, , ,

pho r e gro up unite s with it Th e unio n o f th e to xin mo l e cul e s

.

with th e va ri o us c e lls o f th e b o dy fo r whi ch it p o sse sse s an

affi nity cause s a r e actio n and an inc r e a e d ge n e ratio n O f
,
s

che mical side chains no t in th e ce ll injure d as was first

, ,

tho ught but by ce lls which have b e e n indi re ctly injure d by th e

,

re mo val f ro m active w o r k o f tho se ce ll s i njure d by th e to xin

mo le cule I n o the r wo rds the re has b e e n a co mp e n sato ry
.
,
 60 B I OL O GY OF S PE CIAL I ZED GR OUP S
hyp e rplasia o f ch e mi cal side chains jus t as we have a hyp e r
,

pla sia o f ti ssue c e ll s in chro nic infl am m atio ns Wh e n a ce r tain .

numb e r Of c e ll s have b e e n s timulate d by th e to xi n th e ch e m ,

ical S ide chain s fr o m th e o th e r c e ll s b e gin to b e se cre te d into

th e bl o o d lymph a nd b o dy flui ds
, ,
Th e s e s e cre te d side
.

chains co n stitute th e antit o xin m o l e cul e s .

Th e e xact ch e mical natur e o f th e antit o xi n m o l e cul e I s no t

k no wn Thi s is als o true o f th e to xin mo l e cul e I t h as b e e n

. .

sh o wn that th e antito xin o f diphthe ria is clo se ly a sso ciat e d

with th e se rum glo bulin but thi s is no t p i m f ci e e vid e n ce
-

,
r a a

o f th e a ntit o xin b e ing a gl o bulin Th e antit o xin m o l e cul e is

.

e vid e ntly O f v e ry larg e dim e nsi o ns M o st antit o xic m o l e cul e s

.

d o no t pass thr o ugh a p o rc e lain filt e r O f th e small e st calib e r .

Th e m o l e cul e s b e hav e lik e c o ll o ids b e i ng e l e ctr o n e gative

,
-

a nd th e r e f o r e m o v e t o wa r d th e a no d e o f th e e l e ctrical fi e ld .

Th e a ntit o xin m o l e cul e s a re larg e r than th e c o rr e sp o nding

to xi n m o l e cul e s I n the ir r e actio ns a ntito xins r e s e mbl e th e

.
,

kn o wn pro te in sub s ta nce s m o r e clo se ly than to xi ns The y a re .

no t as labil e as th e t o xin but may b e alt e r e d a nd d e str o y e d

,

with c o mparative e a se .

I n th e c o mm e rcial pr e parati o n o f antit o xins h e althy h o rs e s

a re immu niz e d to i ncr e asi ng d o se s o f t o xi n o r mixtur e s o f t o xin

and antito xin Afte r a c e rtain p e ri o d varying fro m two to

.
,

six m o nths th e h o rse s are bl e d to th e e xte nt o f fi v e o r six lite rs

,

o f bl o o d and th e bl o o d s e rum which s e parat e s o ut o n s tan di ng

,

c o ntai ns th e antito xin (di scharge d side chain s) This anti .

to xin is standardize d against to xin which has b e e n pr e vi o usly

 PATHO GENI C B ACTERI A 2 61

standardi ze d against standard antito xin Dri e d standard .

antito xin is stabl e .

T h e u ni o n b e tw e e n th e t o xin m o l e cul e and th e antito xin

m o l e cul e is a d e finit e che mical co mbinati o n Antito m 15 . x

pro duce d in th e large s t quantitie s in th e S pl e e n and b o n e

ma rro w o f all sus ce ptibl e animal s .

Wh e n a b o dy is na turally inf e ct e d by a path o ge nic bac

te rium ,
o r is immuniz e d with livi ng o r d e ad path o g e nic

bacte ria whi ch pro duce e ndo to xi ns th e r e is de ve l o p e d in th e

,
.

bl o o d se rum and o th e r b o dy fluids Of th e infe cte d animal a

vari e ty o f substance s As h as b e e n pre vi o usly state d th e
.
,

b o dy fluids co ntain no di s tinct antito xins Th e fl uids co n .

ta in th e fo ll o wing substance s .

Anti b acte ri al S u b s ta nce s — The se pro p e rtie s are mani

.

fe ste d by th e abil ity o f th e bl o o d s e rum and b o dy fluids to

d e st ro y th e vitality o f th e bact e ria witho ut d e stru ctio n Of th e

ce ll sub stance (bact e ricidal) o r to di sinte grate in additio n th e
,

bact e rial c e ll (bact e ri o lysi s) B acte ri o lytic actio n may b e

.

d ue to aut o ly sis o f th e bact e ria th e ms e lv e s o r to th e di re ct

acti o n o f th e s e rum Two imp o r tant m o l e cul ar gro ups are

.

co nce rn e d in a bact e ri o lytic and bact e ricidal re actio n Th e .

acti o n o f the se ch e mical gr o ups may b e sho wn by th e fo llo w

‘

ing e xp e rim e nts.

I f an antibact e rial s e rum b e it bact e ricidal o r ba ct e ri o

,

lytic is h e ate d to 60 C fo r twe nty m inute s th e n place d in

,
°
.
,

co ntact with th e sp e cific bacte ria i n viva o r i n o itr th e ba ct e ria o,

will no t b e affe ct e d On th e o th e r hand if s o me n o rmal

.
,
2 62 B I OL OGY OF S PE CI AL I ZED GR OUP S
bl o o d se rum b e add e d th e bacte ria will b e ma rk e dly e ff e cte d
,

and kill e d and in bacte rio lytic s e rum s will b e di sinte g r at d

,
e .

I n additi o n th e e xp e rim e nt S h o ws th e p r e s e nce o f an e xt re m e ly

labil e sub s tanc e in th e bact ricidal se r um whi ch was r e place d

e

by th e fre sh n o rmal se rum Thi s substanc e is call e d th e

.

co mp l e m e nt The re is pre se nt in th e s e rum al so a s tabl e sub

.

stance whi ch r e sists h e at and all e xte rnal ag e nts Thi s sub .

stance is call e d th e a mbo ce pto o r immun e b o dy Th e se

r .

che mical substance s can b e d e fi ni te ly s e parat e d fro m e ach

o th e r . I t h a s b e e n S h o wn by e xp e rim e nt that th e ch e mical

gro up which is call e d th e amb o ce pto r mus t first uni t e with

th e bact e rial c e ll b e f o r e th e c o mpl e m e nt can c o mbi ne and

de stro y it Anti co mple m e nts and anti amb o c e pto rs may b e

.
- -

pro duce d o n immunizing with the s e b o di e s Co mpl e m e nts .

may also b e chang e d into co m pl e m e nto id s by he at Th e am .

b o ce p to r c o mpl e m e nt r e acti o n will b e m e nti o n e d lat e r in th e

-

co nsid e rati o n o f o th e r immun e s ub s tan ce s .

Aggl uti ni ns .Aggluti ni ns are substanc e s which are f o und

in th e bl o o d s e rum o f s o m e no rmal and immunize d ani mal s ,

a nd which hav e th e p o w e r to pr o du ce a clumping Of th e

sp e cific micro o rganism causing so me infe ctio ns G ro up .

agglutinins that is agglutinins fo r bact e ria r e late d to th e

, ,

sp e cie s pro ducing th e agglutinin may al so b e pr e se nt in small

,

am o unts in th e se rum Th e se may b e r e m o ve d by diluting

.

th e s e rum . Th e agglutinin in th e s e r um e nt e r s int o ch e mical

co mbinati o n with th e so call e d agglutin o ge n whi ch is pr e se nt

-

in th e bact e rial c e ll and th e r e s ul ti ng ch e mical r e acti o n p ro

,
 PATHO GENI C B AC TERIA 2 63

duce s an agglutinate o f th e bact e ria Agglutini ns a re pro .

d uce d u nd e r similar co ndi ti o ns to th e a ntito xi ns The y are .

di scha rge d ch e mical side chain s Aggl utino id s may b e p o

.
r

d uce d by h e ati ng o r ad di ng c e rtai n ch e mical s to th e aggl u

ti ni n and anti aggluti nins may b e p r o duce d by im m uni a

,
-
z

tio n with agglutini n Th e ph e no m e no n o f aggluti nati o n is

.

made use o f in th e so call e d G rub e r Widal r e acti o n fo r ty

- -

p h o i d f e v e r and al s o i n t h e diag no si s o f gla nd e rs .

O p s o ni ns
.
— I n s o m e inf e cti o n s ( py o ge nic e tc ) d ue to bac ,
.

t e ria which pro duce e ndo to xin s bact e ricidal sub sta nce s are
,

no t alway s e a sil y pr o duc e d Witho ut do ubt phago cyto sis

.

by th e p o lym o rph o nucl e ar and m o no nucl e ar l e uco cyt e s a nd ,

o th e r c e ll s o f th e b dy is O f gr e at imp o rta nc e in c o mbati ng

o

c e r tain i nf e cti o ns Th e re a re pre s e nt in n o rmal se r ums

.

sp e cific sub stance s call e d Opso ni ns Th e s e ps o nins are in

. o

cre ase d during an inf e ctio n and by th e pro ce s o f im m uni a s z

tio n by vacci natio n Th e phago cyte s d uring an inf e cti o n

.

ca n o nly i ng s t th e
e e ti o l o gical micr o o rgani sm afte r th e y
have co me in co ntact with th e Op so nins Of th e blo o d s e rum .

Thi s h as b e e n p o sitive ly d e mo ns trat e d Th e natur e o f th e .

r e acti o n b e twe e n th e o ps o nin and th e bact e ria and o th e r

i nfe ctio us micro o rganism s is no t kn o wn Th e o p so nins p re .

pare th e bacte ria fo r ph go cyto sis o r to use th late st te r

a ,
e

m ino l o gy th e bact e ria a e

,
s e n sitize d Th e m o r e Op so nin s
r .

th e m o r e phag o cyti c acti o n will tak plac e and th e r e f o r e th e

e ,

mo r e i nfe cti ng ba cte ria will b e d t r o y e d Th e Op so nin s

es .

have no di re ct e ffe ct o n th e l e uco cyt e s o r o th e r phago cyte s .

2 64 B I O L O GY OF S PE CI AL I ZE D GR OUP S
Th e rmal Op so nins may b e inc r e ase d by immunizing th e
no

a nimal b o dy with kill e d bacte ria thu s causing th e ce ll s f th e ,

o

b o dy to r e act and se cr e te mo r e o pso nin a s is th e cas e in anti ,

to xin fo rmati o n .

P r e ci piti ns — I f th e bl o o d se rum Of an animal whi ch has

.

r e c e ntly had an infe ctio n o r b e e n a r tificially immunize d is

, ,

bro ught in co ntact with a filtrate o f bacte rial cul tur e o f th e

same sp e ci e s o f bact e ria which caus e d th e i nfe ctio n o r th e ,

sp e ci e s whi ch was us e d in th e pro ce ss o f immu nizati o n a ,

d e finite ch e mical r e acti o n r e sults Thi s r e acti o n i s char .

a cte ri e d by th e pr e cipitati o n O f c e rtain o f th e pr o t e i n sub

z

stanc e s in th e filtrat e whi ch are e vide ntly de riv e d fro m th e

bacte rial ce lls Th e substance s in th e s e rum a re call e d p re
.

cip itins Th e r e acti o n is n e arly a S p e cific o ne ; that is p re

'

.
,

cip i ta tio n is rar e ly ac co mplish e d by a s e rum wh e n plac e d in

co ntact with a filtrate pro duc e d by bacte ria o the r than tho se
us e d in th e pro c e ss o f immunizati o n o r pr e se nt in th e infe c ,

tio n O ccasio nally fo r e xampl e th e se rum o f an animal

.
, ,

immuni z e d to B typh o sus will p r e cipitate th e filt r ate s o f

.

o th e r m e mb e rs o f th e so call e d Typh o id C o l o n G r o up
-
such -

as B co li B p a atyph o sus e tc This fact i ndicate s that

.
,
. r ,
.

th e r e is a cl o se ch e mical r e latio nship b e twe e n th e se bacte ria .

S p e cific p r e cipitins hav e b e e n Obtain e d fo r practically all

pro t e in sub stance s such as bl o o d milk and e gg album e n , ,

with s e rums pro duce d by immunizing animals with th e s e

fo r e ign substance s P re cipitin s are al so sp e cific fo r th e bl o o ds
.

o f vari o u s animal s P r e cipiti ns a e distinct fro m aggluti ni ns

. r
 PATH O GENI C B ACT ERI A 2 65

I n th e h mi cal r e acti o n th e pre cipitin o f th e s e rum mu st

c e

e nt e r i nt o ch e mi cal c o mbi nati o n with a sub s ta nc in th e fi l e

t r ate d e rive d f ro m th e bacte ria call e d pr e cipitino ge n and by , ,

thus co mbini ng a d e fi nite ch e mi cal pr e cipitat e is f o rm e d

, .

P r e cipitati o n l ik e agglutinati o n d o e s no t tak e plac e in th e

, ,

ab se n ce O f salts ( e l e ct r o lyte s) N o thing d e fini te is kn o wn in

.

r e gard to th e ch e m ical st r uctur e o f th e pr e cipitins Th e y .

a re p r o bably p r o t e in s ub s tanc e s Th e y are p r e cipitate d in

.

th e e ugl o b ul in part o f th e se r um l ike m o st antib di e s , a nd ,

o

cann o t b e s e parat e d fr o m th e s e rum pr o t e ins On h e ating .

s e rums co ntaini ng pre cipiti ns th e y are change d into p re cipi

to i d s . Th e c o mbi ni ng gr o up is u ninjur e d but th e a ctiv e ,

gr o up ( zym o pho r e ) which caus e s th e pr e cipitatio n is d e

, ,

s tr o y e d .

Anti e nzym e s S o m e path o g e ni c bact e ria pr o duc e e n

.

zym e s (M pyo ge ne s var au re u s e tc ) in small amo unts I m

. .
,
. .

m uni a tio n o f a ni mal s with th e s e e nzym e s pr o duc e s a nti e n

z

zyme s in th e se rum o f th e animal Littl e is kno wn in r e gard to

.

th e a ntie n ym e s z Th e r e is a p o ssibility that in ce rtain cas e s

.

the y may i nhi bit bacte rial gro wth Th e subj e ct o f bacte rial .

e nzym e s has b e e n c o nsid e r e d in an o th e r chapt e r ( C hap X V) . .

Ta bl e f
o Anti bo di e s whi ch may be pro duce d by I mmuni zati o n

bacte rial to xins o f

Antito xin s fo r
B act diphth e ri a ( Diphth e ria)
. .

B t e tanus (Te tanus)

. .

B b o tulinus ( B o tul ism)

. .
2 66 B I O L O GY OF S PE CI ALI ZE D GROUP S
P S p yo cy a ne us ( B
. lue gr e e n p us)
-
.

B fe se ri
.
( S ympt o matic anthrax ) .

Anti cyto to xi ns fo r th e vario us to xic pare nchymato us ce lls of

th e b o dy .

Antih e m o l ysins fo r th e he m o lysins Of bact e ria .

Antil e uco cidi n fo r th e l e uco cyte p o is o n o f th e pyo ge nic micro

co cci ,
e tc.

Antit o xins fo r animal t o xins .

Antiv e nin again s t ve n o m o f s nak e s .

Antit o xin fo r p o i so n o f th e sco rpi o n .

Antit o xi n fo r th e p o i so n o f c e rtain sp e ci e s o f spid e rs .

Antit o xi ns fo r s e rum p o i so ns o f s o m e sp e ci e s o f fish ,

e el,

turtl e salamand e r a nd wa sp
, ,
.

Antit o xi ns fo r plant t o xins .

Antiricin fo r p o i so n o f cast o r Oil b e an .

Antiabrin fo r th e p o is o n o f th e j e q ue i ty b e an r .

Antir o bin fo r th e p o is o n o f th e l o cu s t l e av e s and bark , .

Anticr o tin fo r th e p o i so n o f th e c r o ti n Oil b e an .

Antip o l l e n fo th e p o is o n in th e p o ll e n o f vari o us plants

r .

Antis e rums fo r
M sp c o mma (Asiatic ch o l e ra)
. .

B typh o su s ( Typ ho id fe ve r)
. .

M p ne um o nia ( Pn e um o nia)
. .

B p e stis ( B ub o nic plague )

. .

Anti e nz ym e s fo r th e e nzym e s of vario us zym o ge nic bact e ria .

 PATHO GENI C B AC TERI A 2 67

Vacci ne s a nd Age nts i nfl ue nci ng the Op s o ni ns

Vacci ne s have b e e n pre pare d o f th e e ti o lo gical age nts

S mallp o x .

Tub e rcul o sis .

Anthrax .

S ympt o matic anthrax .

Hyd r o ph o bia .

P ractically all bact e ria which pro duc e e ndo t o xins such as , ,

M sp co mma (Asiati c cho l e ra)

. .

B typh o su s (Typh o id fe ve r)
. .

M pyo g e n e s var aure u s ( P us i nfe cti o n s)

. . .

It sho uld b e r e m e mb e re d that o nly a fe w o f th e antib o die s

ab o ve m e ntio ne d a re Of use as the r ap e uti c ag e nts .

F acto r s whi ch i nfl u e nce a nd m o d ify I nfe cti o ns Th e .

co nditio n s whi ch mo dify an inf e ctio n in th e b o dy by bacte ria

o r o th e r micr o o rgani sm s may b e bri e fly s tat e d as f o ll o ws
’

( )
1 Th e virul e nce o f th e i nf e cting ag e nt .

( )
2 Th e numb e r Of th e i nf cting o rganisms e .

(3 ) Th e ave nue o f th e i nfe ctio n .

(4) Th e s ubj e ct o r i ndividual i nf e ct e d .

( I ) Vi rul e nce thi te r m is m e ant th e ability Of an

. By s

o rga ni sm to p ro duce di se ase Thi s p ro p e r ty va rie s and may

.
,

b e i ncre a se d o r d e cr e as e d T h e vi r ul e nc o f a path o g e nic

. e

bact e rium may b e incre ase d by pa ssi ng it th ro ugh a s e ri e s o f

susce ptibl e animals o r gro wi ng it in ce ll o idin sacs place d in
,
2 68 B I OL O GY E
OF S P CI A I ZED G OL R UP S
th e abdo mi nal cavity o r unde r th e skin Of a susce ptibl e anim al .

Th e virul e nc e may b e d e cr e as e d by gr o wing th e bact e r ia o n

cultur e m e dia a nd by g ro wing th e cultur e s at a c mparative ly

,
o

high h e at Virul e nce de p e nds up o n th e ability o f th e ba cte ria

.

o r micr o o rga ni s ms to g r o w in th e b o dy and to subs e qu e ntly

f o rm to xic sub sta nc e s .

( 2 ) N umbe — Th e numb e r o f bact e ria o r o th e r m ic r o

r .

o rganisms n e c e ssary to pr o duc e an inf e cti o n in th e b o dy and

o v e rc o m e i ts r e si s tanc e va r i e s with th e sp e ci e s F o r e xampl e .

,

it has b e e n fo und that th e intr o ductio n O f o ne singl e B act .

anthracis is all that i s ne ce ssary to p ro duce anthrax This .

o ne bact e rium o f c o urs e s o o n m ultipli e s v e ry rapidly in th e

, ,

b o dy Obs e rvati o ns have b e e n mad e o n tub e rcul o si s and

.
,

it has b e e n d e mo nstrat e d that at l e ast e ight hundre d and

twe nty o f B act tub e r cul o si s a e n e ce ssary to pro du ce th e
. r

dise ase e xp e rime ntally in a gui ne a p g I n all pro bability I .

,

in m an a nd animal s it r e quir e s o nly a fe w bact e ria to start a

tub e r cular i nfe cti o n natu r ally .

Watso n Ch e yn e f o und o n e xp e rime ntati o n with B vulgaris

-
.

that si millio n b acilli inj e ct e d und e r th e S kin p ro duce d no

x

l e sio n e ight milli o n p ro duc e d an ab sc e ss fi fty si millio n a

, ,
-
x

phl e gm o n and that two hundre d and twe nty fi v e millio n

,
-

w e r e ne ce s sa ry to pro duce a fatal r e sult in two h o urs .

I t h a s b e e n al so d e m o ns trat e d that two hu ndr e d and fifty

millio n o f M pyoge n e s va r au r e u s w e re r e qui re d to p r o du ce

. .

an ab sce ss a nd o ne billi n w e r e n e c e ssary to caus e de ath in

,
o

an e xp e rim e ntal animal such as a rabbit ,

.
 PATH O GE NI C B ACT ERI A 2 69

Th e s e fe w e xp e rim e nts S h o w that th e int e rnal r e sistance o f

th e b o dy is g r e at e r again s t s o m e i nf e cti ng o r gani sm s than

o th r s
e . The r e is ho w e ve r a limit to th e numb e r o f micro
, ,

o r g ni sms which may b e intr o duc e d int o th e b o d y w

a i th o ut ,

pro ducing any patho lo gical co nditio n .

I t sh o ul d b e r e m e mb e r e d that th e phago cyt e s o f th e b o dy

aid mat e rially in Co mbating inf e ctio ns The y a re able to .

d e stro y o nly c e rtain sp e cie s o f bacte ria and ce rtain numb e rs

O f bact e r ia with o ut th e b o dy b e c o mi ng i nf e ct e d
,
I t will b e .

r e a dily se e n th e r e fo r e that th e numbe r o f bacte ria influe nc e s

, ,

and m o difie s th e inf e ctio n .

(3 ) The Ave nue f I nf cti o n o eB y thi s e xpr e ssi o n is m e ant

.

th e p o i nt and cha nne l o f i nf e cti o n I t mak e s a d e ci d e d di ff e r e nc e

.

in th e sympto mato lo gy and subs e que nt r e sults if th e ave nue

o f inf e cti o n di ff e r s wh e n a d e fi ni t e micr o o r gani s m is th e e ti

o l o gical fa ct o r .F o r e xampl e B act tub e rcul o si s wh e n e nt e r

,
.

ing th e b o dy thro ugh th e re spirato ry o r dig e stive tract may

p ro duce tub e rcul o si s o f th e lungs o tub e rcul o sis o f th e in r

te stine s and acc e sso ry o r ga ns a nd furth e rm o r e u sually te r , ,

m i na te s fatally I f B act tub e r cul o sis i nf e ct s th e ski n a dis

. .
,

tinctl y l o cal di se a se k no wn a s lupu s r e sul t s whi ch may p e r

sist inde fi nite ly witho ut e nding fatally Th e S t pyo ge ne s . r

.

wh e n it e nt e r s th e sup e rfici l ti ssue o f th e b o dy p r o duce s

a

e r y sip e las a nd ab s c e sse s a nd wh e n pr e se nt in th e ci r c ul at o ry

, ,

sy s t e m p r o duc e s s e ptic e mia

,
M pn e um o ni a may p r o duce
. .

p ne um o nia absc e sse s co nj unctivitis o titis and m e ni ngitis

, , , , ,

de p e nding up o n its p o int o f e nt r ance and th e ti ssue attacke d .

2 7 0 B I O L O GY OF S PE CI AL I ZE D GR OUP S

(4 ) The S ubj e ct I nfe cte d susce ptibility o f indi

. Th e

vi d ual s to i nf e cti o ns vari e s with th e ( 1) sp e ci e s ( 2 ) with rac e ,

and individual i di o syncraci e s and (3 ) with th e age o f th e in di

,
~

vidual Th e sp e ci e s Of animal whi ch is attack e d by a path o

.

ge nic micro o rgani sm may b e r e sistant (immun e ) o r sus

ce p tib l e An animal may b e naturally immun e t o inf e cti o n
.

o r may hav e acquir e d immunity by having a di se as e o r by

vacci nati o n with a patho ge ni c nu cro o gam sm I n a m o difi e d r

fo rm F o r e xample ani mal s a re immune to such dise as e s as

.
,

go n o rrh oe a l e pro sy e tc which aff e ct man Th e y d o no t

, ,
.
,
.

naturally co ntract such e ndo to xic di se ase s as typh o id f e ve r ,

Asiatic ch o l e ra but may b e ar ti ficially inf e ct e d by th e b ac

,

t e ria wh e n intr o duce d artificially Tub e rculo sis is m o re .

co mm o n am o ng th e ne gro rac e than amo ng o th e r r ace s .

Dimi ni h e d vitality o f th e b o dy d ue to e xp o su r e use o f d r ugs

s , ,

injury l o ss o f sl e e p o ve rwo rk and di so b e di e nce o f o th e r no r

, , ,

mal hygi e nic co ndi tio ns influ e nce th e r e sult s o f an infecti o n .

The r e are c e rtain di se a se s fo r e xampl e diphth e ria whi ch is

, , ,

mo r e co mm o n in yo u nge r life than in adult lif e Tub e rcul o sis .

is a di se a se which is m o r e co mmo n in e a rly lif e ( 5 3 0) than— 2

at any o the r p e rio d o f life Th e re is e vide ntly mo r e susce p

.

tib il i ty to inf e cti o n at c e r tai n p e r i o d s o f lif e .

I n r e gard to s o urc e o f th e inf e cti o us micr o o rga nisms it ,

may b e bri e fly stat e d that path o g e nic bact e ria and so me

o th e r inf e cti o u s o rganisms e tc may b e n o rmally p r e se nt in
,
.
,

th e n o se m o uth i nt e s tin e s r e spi r at o ry tract o n th e S ki n and

, , , , ,

may al so b e i ntro duce d thr o ugh th e age ncy Of f o o d and d rink ;

 PATH O GENI C B ACTERI A 2 7 I

I m m uni y to t r I mmuni ty is r e si s t
t
P a h o ge ni c B acte i a .

ance to di se e Th e ab e nce o f such r e si stance characte rize s

as . s

s u c e ptibility
s Th e s ubj e ct o f immuni ty in all its pha se s is
.

b e i ng car e f ully inv e stigate d at th e pr e se nt time by many

sci e ntific w rk e r s I t is o ne o f th e m o st inte re sting o f th e
o .

c o mpa r ative ly un e xpl o r e d fi e lds o f sci e nce Onl y a ve ry .

b ri e f o utline o f th e subj e ct is give n b e lo w .

Th e fo ll o wi ng tabl e wi ll b e s e rvic e abl e a s an o utlin e

R a ial
c

Nat ural
I m m un it y
I mm un it y
Ac q r
ui e d

N atural immunity is th e immunity which c e rtain sp e cie s o r

race s o f ani mal s and plants p o sse ss against patho ge ni c bac
t e ria a nd o th e r di se ase p ro ducing mic ro o rgani ms F o r -
s .

e a m pl e m a n i s immun tO chi ck e n a nd h o g ch o l e ra and th e

x
h
e ,

ho r e is immune to typho id f e ve r a nd Asiati c cho l e r a Th e

s .

lo we r animal s d o no t have mala ria y e l l o w fe ve r scarl e t f e ve r , , ,

or me a sl e s Th e ne gro as a rac e is mo re immun e to y e llo w

.
, ,

fe ve r than th e whi te race .

Acqui r e d immunity is that r e si s tanc e which is acquir e d by

havi ng a di e ase by b e i ng va ccinate d o by i nj e ctio ns with
s , ,
r

antise rums That immunity whi ch is acqui re d by havi ng th e

.

di ase o by vaccinati n is call d ti v and that acq ui r e d

se ,
r o ,
e ac e ,

by b e ing i no culat d with an anti se r um a f e xampl e an

e ,
s, or ,

antito xin is call e d p assive im munity Vac cinatio n is th e

,
.
2 7 2 B I OL O GY OF S PE CI AL I ZE D GR OUP S
inj e ctio n i nto th e b o dy O f th e m o di fi e d a nd atte nuate d bac
t e ria o r virus o f a di se ase so a s to p ro duc e th e di se ase in a much
l e ss s e ve r e f o rm than no rmally I n active immunity th e r e is
.

a r e actio n o n th e part o f th e b o dy ce lls wh ile in pa ssive ,

immuni ty th e c e lls d o no t r e act and th e antib o dy e xis ts in

active until co mbine d with th e to xic sub s tance s Of th e micro
o r ga nisms . Acquir e d immunity to c e rtain i nfe cti o ns may ,

to s o m e e xt e nt h e i nh e rit e d acc o rdi ng to th e lat e st w o rk o n

,

thi s subj e ct .

I mmunity may b e e ith e r antibact e rial that is again s t , ,

bacte ria (e ndo to xin s) o r antito xi c that is agai nst th e to xi ns

, , ,

s e cr e te d by th e bact e ria Th e t e rm immunity is a r e lative

.

o ne .L arg e numb e r s o f path o g e nic mic r o o rga ni sms o r larg e ,

amo unts o f t o xin always pr o duce so m e e ff e ct o n th e animal

,

b o dy whe n inj e cte d fo r th e purp o se o f e xp e rim e nt .

The The o i e sr f I om m u ni ty ( ) E. xhausti o n Th

1 e o ry .

I t was claim e d at o ne tim e by P a s t e ur that in acqui r e d im

munity th e bacte ria o r infe cti ng ag e nts pre se nt in o ne infe c

ti o n us e d up all th e pabulum n e ce ssary fo r th e vital activity Of
th e o rganisms and that in natural immunity th e se substance s
,

w e re n e ve r pr e se nt I nf e cti o n is th e re fo re imp o ssible

.
, ,
.

( )
2 N o xi o u s R e t e n ti o n Th e o ry T hi s th e o ry. h e ld that
s o me sub stance s o f a n o xi o u s charact e r w e re p r e se nt in th e
b o dy n o rmally in so me case s o r w e r e r e tain e d in th e b o dy as
,

a r e sult Of a pr e vio u s infe ctio n A s e co nd inf e ctio n was .

,

the r e f o r e impo ssibl e fo r thi s r e a so n

,
.

(3 ) P hag o cytic Th e o ry M e.tchnik o f

f cla i m s that th e
 PATH O GENI C B ACTERI A 2 73

activity o f th e phago cyte s ( l e uco cyte s e tc ) in th e b o dy a e

,
r

r e sp o nsible fo r th e immuni ty I t is v e ry p r o babl e that

.

immunity to ce r tain di se a se s de p e nds o n th e phago cyte s .

(4) C h e mical Th e o r y Eh rli ch a nd o th e rs h o ld th e Vi e w

.

that ce rtain che mical sub stance s which a re pre se nt to a

limit e d e xt e nt in all no r mal se rum s and ce lls are i ncr e a se d
aft e r having a n infecti o n o r vaccinati o n and a re r e sp o nsibl e
fo r b o th natural and a cquir e d immunity o f th e b o dy . Th e se
substance s a re bacte ricidal substanc s agglutinins p re
e , ,

cip i tins
,
o p so ni ns
,
e tc
. and ntito xi ns
,
a .

I mmunity can b e m o s t sati s fact o rily e xplai ne d o n th e ba sis

O f th e phag o cyti c acti o n o f th e b o dy c e ll s a nd th e ch e mi cal

actio n Of ce rtain substance s in th e b o dy flui ds ( N o s 3 ) and

.
 CHAP T ER XXI V
B ACTER I AL DI S E AS E S OF NTS
PL A

In tr o d ucti o n M o s t o f th e S CI e ntifi c r e se arch o n th e

.

path o ge nic bacte ria has b e e n do n e o n tho se sp e cie s which

pr o duc e dis e ase co nditio n s in man and animals F e w im .

p o r tant inve stigati o ns o f any co ns e qu e nce w e r e made pri o r

to th e la s t d e cad e o n th e subj e ct o f bact e rial di se as e s o f plants .

R e c e ntly bact e ri o l o gi s t s hav e b e e n paying m o r e att e nti o n to

thi s subj e ct Th e Unite d S tat e s go ve rnme nt h a s e stablish e d

.

in th e De pa r tm e nt o f Ag r i cultu r e a B ur e au o f P la nt I ndustry .

Th is bure au is active ly e ngage d am o ng o th e r thi ngs in mak

ing path o l o gical and phy si o l o gical i nv e s tigati o n s o f pla nts .

I n lab o rat ri e s in o th e r parts o f thi s c o untry and abr o ad

o

r e se a r ch is no w b e ing carri e d o n al o ng th e lin e s Of plant

patho l o gy .

E a l y C o nce pti o n
r I t was h e ld fo r s o m e tim e by c e rtain
.

writ e rs , and is stil l h e ld to a limi t e d e xt e nt by so me inv e sti

gat o rs (F isch e r Wa r d) that bact e ria d o no t cau se p rimary
, ,

i nf e cti o ns o f plants Th e o nl y e xce pti o n to this co nditio n o f

.

aff air s it is h e ld is in th e case o f th e bacte rial no dul e s whi ch

, ,

de ve l o p o n th e ro o t o f th e l e gumi no u s plants F i s ch e r fo r
s .
,

e xampl e ,
i n hi s Vo rl e su ng e n iib e B akte ri e n mak e s th e
r

fo ll o wing state m e nts He call s att e ntio n to th e anato my o f

.

th e pla nt a nd sh o ws that ba ct e r ia can o nly e nt e r thr o ugh th e

,

2 74
 B ACT ERI AL E E
DI S A S S OF PLAN T S 2 75

st o mata and l o dg e in th e i nt r a ce l l ul a sp ce s und r natural

‘

-
r a e

c o ndi tio n s He argu s that th e bact e ria can de rive no nutri

. e

m e nt f ro m th e mate rial co ntaine d in th e spa ce s b e twe e n th e

ce ll s and th e r e fo r e canno t pro duce any di a se in th e plant
,
se .

He h o ld s th e o pini o n that path o g e nic bact e r ia ca nno t di s

s o lve th e ce llul o se wall Oi th e plant ce ll F urth e rmo r e th e .
,

s tat e m e nt i s mad e that th e o nly sp o r s which ca n p o ssibly e

g e rmi nate in th e aq ue o us mat e rial

in th e int r a ce llula r space s a e tho se
-
r

o f th e para siti c fung i ( Hyp h o m y

c e t e s e tc ) since th y p o sse s re
,
. e s

s e rve fo o d mat e rial Th e se fungi .

,

it is claime d may di sso lve away th e

,

c e ll wall p e ne trate th e ce ll with th e

,

hyp ha and thus p ro duce a m o rbid

,

co nditio n the r e in ( rust s ,

F i sch e r and so m e o th e r s thi nk

that th e bact e rial infe cti o n o f th e

plant is se co nda ry to i nfe ctio n by
Cr ti f t r ip
fungi He stat e s that th e uni n l af i lat d with B a t m
.
F 5 IG

e
. 2 .

no cu
o s s se c

e
on o

c .
u n

ca

jur e d plant is imp e rm e abl to th e D tr (P r l t r i la e e s e u e cu u e no cu

ti with dl ) Aft r Sm ith on ne e e e

attacks o f bacte ria and th t e ve n
. .

,
a

in th e inju r e d plant o nl y th o s e c e ll s whi ch are atta ck e d by

th e hyph o myc e t e s e tc a e availabl e

,
f r bact e rial acti o n
.
,
r o .

Th e fa ct that ba ct e ria are p r s e nt in di se a e d pla nt s in e s

e n r m o u s numb e r s i s O f co u r e
o r e co gni ze d but F i sche r nd
,
s , ,
a

o th e rs thi nk that th e y are e xi ti ng o nl y m e ta tro p hi cal l y s .

2 7 6 B I OL O GY OF S PE CI AL I ZE D GROUP S
r e s e nt C o n ce pti o n Again st th e Opini o n o f the se fe w
P .
—

inv e stigato rs will b e fo und th e maj o rity o f patholo gists

and bact e ri o lo gi sts I t is no w agre e d by th e l e ading au
.

th o i ti e s that th e r e a e di s tinct ba ct e rial di e a se s o f pla nts

r r s .

FIG 5 3
. . So f t ro t o f gre e n cucum br e s i no cu lat d with B
e . caro to vo rus .
( Pr
u e

cul t r in
u e o culati on .
) Aft r S m ith
e .

Th e se di se ase s can b e pro duce d in th e vari o us sp e ci e s o f

h e althy plants by ino culati o n with de finite sp e cie s o f bac
t e ria ( Fig .

S p e ci e s o f bact e ria have b e e n r e p e at e dl y is o lat e d fr o m di s

 B ACT ERI AL E E
DI S AS S OF PLANT S 2 77

e as e d plant s and r e in o culate d into h e althy plants pro ducing

‘

th e id e nti cal di e a se The se e xp e rime nts are co nclusive

s .
,

a nd no sub s tantiati o n e xi s t s fo r th e f o r e g o i ng stat e m e nt s Of

Fisch e r e t al N o pr e inva si o n by a fungu s is n e c e s sary I t

, . .

is no t unco mmo n ho we ve r to have a distinct fungus di se as e

, ,

in plants ( rust s ,

I t must b e r e c o g ni z e d that th e signs o r sympto ms o f bac

te ri al di se a se s in plants

a re in th e main q uit e S im

il a r Thi s is particularly
.
'

true o f th e bact e rial r o ts .

S o m e tim e s car e ful s t u dy

is ne ce ssary in o rde r to di s
c riminate b e twe e n the se
va ri o u s r o t s ( Fig
“ ”
.

S o m e O f th e bact e rial d i s

e as e s O i plants hav e b e e n

p r o ve d to b e t r ansmitt d e

fro m plant to plant by

m e ans Of th e bit e s Of i n F 5 4 B a t ri m am p tr i t r ip
“

IG . .
-
c e u c es e n u n

S e cts .
r t ( V a lar l ) Aft r S m th
oo . scu o cc usro n . e i .

I t has b e e n no t e d that th e bact e ria w hi ch p r o duc e th e

va ri o us plant di se ase s s e c r e t e an e nzym e which di sso lve s th e

c l lul o se O f th e ce ll wall Th e a ctio n o f the se e nzym e s is
e .

sl w in co mpa riso n with th o se o f th e bacte ria patho ge nic to

o

anim als Ab o ut o ne hund re d a nd twe nty fi e s parate

.
-
v e

di e ase s o f plants have b e e n r e c gnize d thus far

s o .
2 7 8 B I OL O GY OF S PE CI AL I ZED GR OUP S
M e th o d ti o n I n plant s as in ani mal s bact e ria
of I nf e c .
, ,

u su lly cann o t e nte r th e b o dy unl e ss it h as b e e n i njur e d in

a

so m e way and an av e nu e o f inf e cti o n thu s p r o vid e d Th e .

h e althy uninjure d animal o r pl nt b o dy is u ually imp r e g a s

nabl e as far as ba ct e ria are c o nc e rn e d Th e r e sis tance o f th e .

FI G 5 5
. . B act e ri um pruni in gr een pl um .
(C avity f rm ati o on .
) Af t r Sm ith
e .

b o dy must b e l o w e r e d b e fo re inf e ctio n is p o s ibl e I n th e s .

case Of plants th e r e sistan ce o r vitality o f th e plant is u sually

,

l o w e re d and th e ave nue Of infe cti o n pr o vide d by th e sting o f

an ins e ct a f r o st bit e o r a brui se Thi s is pa r ti cularly n o tic e
, ,
.

abl e in th e cas e o f th e fruits o f plants whi ch have b e co m e

inf e cte d A fe w case s f bacte rial di se a e a e o n r e co rd wh e r e
'

. o s r

no l e si o n o f th e pla nt co uld b e f o und .

F IG .
56
. Ol iv kn t ( t um r )
e o s o s .
( Cu lt ure lati
ino cu on .
) tr
Af e Sm ith .
2 80 B I O L O GY OF S PE CI ALI ZE D G R OUP S
Oft e n
sp e cific sys te ms in a plant are affe ct e d a nd the se ,

al o ne F o r e xampl e th e vascular syst e m may b e o cclude d

.
,

( g
F i A t
. o th e r tim e s th e bact e r ia invad e th e i ntra
c e llular spac e s and f o r m cavitie s ( Fig At o th e r tim e s .

the re is a co mp e nsato ry hyp e rplasia o f ce l ls and as a r e sult , ,

tumo rs are fo r m e d ( Fig .

Th e fo ll o wi ng pa r tial li s t giv e s a fe w o f th e bact e rial d is

e as e s pr o duc e d in pla nt s and s o m e O f th e bact e r ia which

p ro duce th e m Th e bact e ria are in s o me in s tanc e s no t

.

sp e cific and ce rtain sp e cie s Of bacte ria may cause dise as e

in s e v e ral S p e ci e s o f plants .

To ba
wilt cco P d m na
se u o o s so l a nce a ru m

C m b r wilt
u cu e B a ill u i d
c s ar o ea

M l n wilt
e o B a ill t h
c us ra c e ip h i l us
T m at wilt
o o P u d m na
se o o s so l a nce a ru m
B light p t n gr or s o s o een a pp l e s,
q u n i
ce s ,

p r n p ar p l m
u e s, e s, u s, e tc . Ba c ill u m yl
s a o v o rus

P tat t
o o ro P se u d m na l n
o o m
s so a ce a ru

T r ni p
u t ro P se u d m na a m p tr
o o s c es e

C arr t t o ro Ba c ill t
us ca ro o v o ru s

Di f th l av
s e as e s o e e es a nd s e tm f t s o co Pse u d m n
o o l m
as va scu a ru

t n w t rn br
o ,
s ee co ,
oo m co rn abbag
,
c e ,
Pse u d m na a m p tr
o o s c es e

e tc . P se u d m na m l
o o s m a va ce a ru

B a t rial liv kn t ( t m r )
c e O e o s u o s Pse u d m na t w ti
o o s s e ar

Di a se f wal t fl wr
se O nu s , o e s o f p la nts , Pse u d m na hy a inthi
o o s c

e tc . Pse u d m n a j g l nd
o o s u u a e s
 P A RT VI I . DI S T R I B U T I O N O F .

BACT E R I A

C HAP T ER XXV
B AC T E R I A OF TH E S OI L

Di s tri b u ti o n a cco rdi n g to H a b ita t

me plants are . So

l o calize d a nd app e a r o nly unde r ce r tain co nditio ns whil e ,

o th e r s are m o r e co s m o p o litan a nd what i s t r u e o f pla nt s a nd

,

animal s is tru e al so Of b cte ria I t i a matt e r o f so me co n

a . s

sid e rabl e imp o rtance to kno w th e r e lati o n o f bact ria to ce r e

tai n l o catio ns At fir s t tho ught it might b e supp o se d that

.

bacte ria w e r e e ve nly di stribute d in natur e but thi a s has ,

s,

b e e n said is no t true
,
.

B acte ri a f th e S o il
o Th e bact e ria fi nd c o nditi o ns favo r
.

abl e fo g ro wth in th e so il a nd th e upp r laye r s o f th e sur f ce

r ,
e a

o f th e e arth t e e m with th e m Th e g r e a sy f e e l which th e e a r th

.

h as and with which e v e ry o ne i s familiar i s d ue in larg e ,

me asu re to th e ba ct ria whi ch it co nt ins Of co ur se c e r tain

,
e a .
,

pa r ts o f th e su r fac e so il a e much mo r e abund ntly suppli e d

r a

with bact e ria than o the r parts ; th e law s go ve rning this will
b e di scu ss d a s we pr o c e e d but in a g e n e ral way th e numb e r
e ,

o f o r ga ni s m s in a ny pa r ticul ar o il d e p e nd s Up o n th e am o u nt
s

o f f o o d s ubstanc o r humu s whi ch 5 pr e s e nt

e ,
A gr e at ma ny
,
1 .

2 81
2 82 DI S T RI BUTI O N OF BACTERI A
o rgani sms are asso ciate d with putr e facti o n a nd de cay and ,

whe r e the se p r o ce sse s are active ly go ing o n e n o r mo u s numb e rs

o f bact e r ia are f o und ( F ig .

FI G 5 7
. . Co l ni
o e s o f so il ba t ria
c e . Af t r Lipm n
e a .

Q uanti tativ e Di s tri b u ti o n of r

B a cte i a i n th e S o il

Th e n umb e r o f bact e ria in diff e re nt s o ils is d e t e rmi n e d by

th e cha r act e r Of th e s o il I n s o m e s o ils th e y are c o mpara
.

tiv e l y fe w in o th e rs th e numb e rs are e n o rm o us

, Th e num .
 B ACT ERI A OF THE S OI L 2 83

b e r o f bact e ria in vari o us s o il s b o th in a ve g e tative and sp o r e

,

co nditi o n ,
a re sh o w n in th e f o ll o wi ng tabl e :

BA T C E R I A IN S UR FA CE LA Y ERS OF S OI L

KIND SOIL
B ACT ER A I PE R

G RAM
OF

V irgin so il Wo o d s,

V irgi n so il Gra ss p l o t
,

V irgin so il Pa s t ure
,

M ad S il C ll g Hill
e o ,
o e e

M ad S il H ill id
e o ,
s e

M ad S il C ll g Hill
e o ,
o e e

M ad S o il B a nk f L a k
e ,
o e

C ultivat d S il R ad id
e o ,
o s e

C ltivat d S il Grai n fi l d
u e o ,
e

C nta m i nat d S il S tr t
o e o ,
ee

C nta m i nat d S il B arny ard

o e o ,

C nta m i nat d S il H t l y ard

o e o ,
o e

Th e re is a marke d ve r ti cal di stributi o n o f th e bacte ria in

th e s o il Th e gr e at e s t numb e r a re in th e fi r s t S i i nch e s a nd
. x

afte r a de pth o f o ne f o o t th numb e r fall s Off v ry r apidly e e

C mparativ e ly fe w ba ct e ria ith e r in th e v g e tativ e o r sp o re

o ,
e e

stat e a e f o und at de pths o ve r fi e o r si f e e t

,
r v x .

Th e S p e ci e s Of B cte ri a — Th e bact e ria o f th e s il a e

a . o r

mainly sapro phytic in natu re and are a sso ciat e d with th e

p ro ce sse s o f p ut re factio n a nd de cay A g r e at many are al so .

m e re ly sap Ophi l ic that is th e y r e quir e d ad o rganic matte r

r ,
e ,

but are no t large ly co nce rne d in its de co mp o sitio n The re .

a re a fe w path o g e nic o r di s e ase p r o duci ng bact e ria w hi ch -

2 84 DI S T RI B U T I O N OF B AC T ERI A

have th e ir h o m e in th e so il such a s th e B t e tani and B

,
. .

fe se ri (sympto matic a nth r ax) A g r e at many m o r e di se a se

.

pro ducing bacte ria simply pa ss a Sho rt but va riabl e time in

th e s o il in an inactive c o nditi o n
,
The re h as b e e n a gr e at de al
.

o f di s cu ssi o n as to wh e th e r o r no t path o g e nic bact e r ia can

gro w in th e so il a nd it is still an Op e n que stio n h o wl o ng th e se

,

bact e ria can p e rsist in th e so il if the y d o no t g ro w The r e is .

,

p e rhap s no go o d r e aso n a pri o i wh y dis e ase pr o ducing bac

,
r -

t e ria might no t gro w in th e so il I n ce rtain lo caliti e s at l e as t

.
, ,

th e r e wo uld b e an abundanc e o f fo o d and at c e rtain s e aso ns o f

,

th e y e ar th e r e is suffi ci e nt h e at S o m e path o g e nic f o rms will

.

gro w in me di a made fro m so il e xtract but th e e vid e nc e that ,

bacte ria can gro w in th e natural s o il is v e ry S light and th e ,

chi e f r e a so n why th e y d o no t gro w it is e asy to mag n e 15

,
I i ,

b e cause o f th e antag o ni sm o f what may b e te rm e d th e so il bac

t e ria Di se a se bacte ria a re pr e s e rv e d in th e so il fo r a co n
.

sid e rab l e p e ri o d o f tim e a nd t h is i s e sp e cially tru e o f th o s e

,

bact e ria whi ch have sp o r e s S o il which h a s b e co m e co ntami

.

na te d with th e B act a nthraci s is da ng e r o u s s o il fo r ma ny

.

y e a r s Th e sp o re l e ss fo rms live fo r a much sho r te r time and

.
,

o rdi na r ily th e y ca nn o t c o p e with th e u nfav o rabl e c o n di ti o ns

whi ch surro und th e m e xc e pt fo r a bri e f time Th e p r actical .

imp o rtanc e o f this pr o bl e m co m e s o ut in th e que tio n o f th e s

p e rsi ste nc e o f di se a se bact e ria in huma n o r animal cadave r s .

Th e p o pular n o ti o n is that a c e m e t e ry is a da ng e r o us plac e

and many e pid e mics have b e e n trac e d in th e p o pul ar mind , ,

to cl o s e p r o ximity to s uch plac e s o r to co ntaminati o n o f

,
 B ACT ERI A OF THE S OI L 2 85

wat e r fr o m the m e tc Wh e n e v e r a grave is Op e n e d that

,
.

co ntains th e b o dy o f a p e r so n d e ad o f an i nfe ctio u s di se as e o r ,

an att e mpt is mad e to m o ve a c e m e te ry f e ar is invariably ,

aro use d am o ng p e o pl e livi ng in th e n e ighb o rho o d The re is .

ve ry littl e e vide nce h o we ve r to S ho w that any o f th e bact e ria

, ,

O f human di se a se can liv e fo r any l e ngth o f tim e in th e s o il .

B act a nthraci s wi th ut S p o r e s liv e o nly a fe w days B

. o ,
.

typh o sus and M sp c o mma o nly a fe w w e e k s a nd th e B act

.
, .

tub e rcul o sis no t o ve r a fe w m o nths Th e antago nism o f .

o th e r f o rm s is und o ubt e dly a p o t e nt fact o r in th e ir e a r ly d e

structi o n I t is e asily s e e n th e n that with th e e xc e ptio n o f

.
, ,

th e B t e tani wh o s e di stributi o n in th e so il i s pra ctically w o rld

.
,

wid e that th e dis e as e pro ducing bact e ria hav e o nly a ve ry

,
-

l o cal di stributio n and th e se l o caliti e s b e ar a de finite r e latio n

,

to s o m e pr e e xi s ti ng ca se o f di se a se O th e r so il bact e ria o f a
.

ve ry gre at impo rtanc e a re th e l e gume and nit rifyi ng bacte ria .

A di s cu s si o n o f th e bi o l o gy o f th e s e is f o u nd e l se wh e re .

Th e Le gu m e o N i tro ge n fi xi ng B a cte ri a
r -
Thi s gro up o f .

s o il bact e ria h as alre ady b e e n r e fe rr e d to a nd th e i r bio l o gy

discu sse d but it is de sirabl e in thi s co nne ctio n to e mphasize
,

th e ir imp o rtanc e as nitr o ge n gathe re rs Nitro g e n is an e ss e n .

tial plant f o d Wh e n th e supply is limite d in a so il it is

o .
,

fo und ne ce ssary t r e pla c it N it ro ge n f rtilize r s a e at th e

o e . e r ,

p re se nt tim e ve ry e xp e nsive I t h s b e e n kno wn fo a ve ry l o ng

,
. a r

tim e that c e r tain l e gumi no us pla nts w e re abl e t incr e a s e th e o

nitro ge n supply O f th e o il a nd it was lat e r fo u nd that thi s

s ,

activity was as so ciate d with th e pre se nc e o f n o dul e s o n the se

2 86 DI S TRI BUTI ON OF BACTERI A
l e gume s ( Fig . It

is no w kn o wn that th e
nitro ge n fixatio n is
b ro ught ab o ut by th e
bacte ria whi ch pro duce
th e se n o dul e s Th e
.

kn o wl e dge whi ch has

b e e n gaine d in r e ce nt
y e ar s has Op e n e d up a
numb e r Of pro bl e m s ,

th e s o luti o n o f which

w o ul d b e o f th e g re at
e st imp o r ta nc e to ag r i

cultur e . So me s o il s

s e e m to lack th e pr o p e r
ki nd o f n o dul e bact e ria ,

and it is n e c e ssary that

fi e ld s whe r e l e gumi n o us
cro p s a e to b e gro wn
r ,

such as cl o v e r o alfalfa r ,

S h o uld be i no culat d e

with th pa r ti cular b ac
e

t e ria ( Fig . At
t e mpts have b e e n mad e
FI G .
58 .
— T brl
u e ce s on th e ro o t s o f an o l d
to put up cultur e s O f
alfalfa pl nt
a .

th e se bact e ria in a fo r m
whi ch o r th e fa r m e r Up to th e
.
 B AC TERI A OF THE S OI L 2 87

pr e se nt time ho w e ve r e ffo r ts in thi s di r e ctio n have large ly

, ,

fail e d and r e co u rse must b e had to th e t r ansf e r e nce o f th e s e

,

bacte ria th ro ugh th e natur al so il in whi ch the y a re f o und

rathe r than by m e ans O f a r tifici l cul tur e Ano th e r pro bl m a s . e ,

p e rhap s uto pian in natu re is to ado pt th e se l gume ba cte ria

,
e

to a lif e o n th e r o o ts o f

o th e r kind s O f pla n ts .

F r i ns tance
o if we
,

co uld te ach th e l e gume

bacte ria to g ro w o n
th r o o ts o f c o r n th e r e
e ,

s e e ms no r e a so n why
we co uld no t rai e s

thr e e hund r e d bush ls e

of c o rn to th e acr e — B a t r id fr m l g m
F 5 IG .
9 .
( ) Fr m c e o s o e u es . I o

Wh e r e we HOW rai e M lil t

s alba ( 3 d 5 ) fr m M di ag
e o us 2 , , an o e c o

ativa ; (4) fr m Vi ia vill a (Harri

s d o c os so n a n

B arl w) Aft r L ipm o . e an .

S o m e bact e ria a e r

abl e to fix th ni t r o g e n o f th e ai r inde p e nde nt o f any sym

e

bio tic r e latio n with th e l e gum e s I t is qui te lik e ly that it .

will de ve l p that the s e o rganisms a e o r may b e co me ve ry im

o r , ,

p o rtant age nts in th e mai nte nance o f s o il f e rtility Al re ady .

atte mpts have b e e n made to cultivate th e se bact ria and put e

the m up in co mm e rcial fo r m I f it sho uld tu rn o ut that thi s

.

p
i s o ss e ibl th
,
e e livi
s n g n it r o g e n gath e r e r s
co uld b e b o ught
in th e mark e t and s wn o n th e fi e ld whe re th e y w o uld fix fo r
o ,

plants th e ni tro g e n fro m th e ine xhaustibl e supply in th e ai r .

2 88 DI S TRI BUTI ON OF B ACTERI A
r r
Th e Ni t i fyin g B a cte i a The y are e xce e dingly imp o r
.

tant o rganisms fro m th e standp o int o f th e cycl e o f nit ro g e n

in nature . Th e sapr o g e nic a nd sap ro p h ili c bact e ria br e ak

do wn th e o rganic matt e r to th e fo rm o f ammo nia Thi s .

is availabl e to a li mit e d e xt e nt if at all to th e gr e e n plants

, ,
.

Th e c o nv e r si o n o f th e amm o nia fo rm e d during th e pr o c e ss

o f putr e facti o n i nt o th e nit r at e s is a matt e r o f gr e at e st im

p o rtance in s o il f e rtility This chang e is e ffe cte d by th e

.

nitrifying bact e ria I n o rde r that the y may d o th e ir wo rk

.
,

c e rtain c o nditio ns must e xi s t Th e so il mus t b e r e adily

.

p e rme abl e to air sinc e th e se bact e ria are o xidi zi ng o rganisms

,
.

Drainag e and m e th o d s o f c ultivati o n are o f ve ry g r e at imp o r

tance in thi s co nn e ctio n I n o rde r that nitrificatio n may
.

pr o ce e d succ e ssfully it is n e ce ssary that th e acids pro duc e d

,

by the m b e n e utralize d by base s A so il to e nco u r age nitrifi

.

catio n mus t the n have suitabl e base s Th e que stio n o f s o il

, ,
.

f e rtility is the n in its las t analysis a bacte rio l o gical pro bl e m

, ,
.

De ni trifying B acte ri a —
Th e d e nitrifying bact e ria a re
.

fr e que ntly fo und in th e s o il The i r actio n is just th e r e ve rse

.

o f th e nitrifying bact e r ia ; that is th e y r e duc e th e nitrat e s ,

back to ammo nia I n thi s way th e nitro g e n is lo st as fo o d

.

fo r th e high e r plants S uch bact e r ia a e th e n v e ry und e

. r , ,

sirable fro m th e agricultural s tandp o int I t is ve ry fo rtu .

nat e that th e co ndi tio n s whi ch a e de mande d by th e nitrify r

ing bacte ria a re tho se whi ch a e m o s t unfavo r abl e fo r thi s

r

class ; h e nce co nditio ns which favo r nitrificatio n i nte rf e r e

,

o r pr e v e nt th e activity o f th e d e nit ifying bact e ria r .

 C HAP T ER XXVI
B ACTE RI A OF THE AI R

t n Ai r practi cally always co ntains bacte ria

I ntro d uc i o .
—
.

I n cro wd e d citi e s dusty str e e t s and e l se wh r e th e numb e rs

, ,
e ,

may b e ve ry la rge On high a w capp e d m o untai ns and

.
,
-

o v e r th e se a at so m e di s ta nc e f r m th e sh o r e th e bact e ria a re
o

ve ry fe w T h e bact e ria a e no t co n stant in the ir p re se nce in

. r

th e air but vary at di ff e r e nt tim e s o f th e y e ar and u nd e r

,

d iff e re nt co nd i tio ns I t h a s b e e n s upp o se d in th e pa s t that

‘

th e bact e ria o f th e air fr e qu e ntly includ e th o se Of gr e at dis e a se

pro ducing p o we r a nd it h as b e e n suppo se d that ce r tain di s

,

e a se s c o uld b e ca rri e d fo r lo ng di stanc e s thro ugh th e air .

M o de rn vi e w s h o w e ve r se e m to indicat e that th e bact e r ia

, ,

o f th e air a re far l e ss da ng e r o u s tha n wa s f o rm e r ly th o ught .

C o nd iti o n i n Ai r — T h e bact e ria o f th e ai r are no t i n a

.

state o f active gro wth but a e in a do rm nt co nditio n while

,
r a

in th e air A m o me nt s re fle ctio n will S h o w that thi s is o f

.
’

n e ce ssity so B a cte ria wo uld b e quite unabl e to ge t th e ir

.

fo o d in a dri e d co nditio n F urth e r mo re th e bacte ria a re u u

.
,
s

ally attache d whil e in th e ai r to so me du st pa r ti cl a nd ve ry e ,

fr e que ntly th e y ride o n th e se dust particl e s in small masse s

and no t a s in d ividual s .

G ri gin o f B acte i ar — Th e air bact e ria a re d e rive d ve ry

.

U 2 89
2 9 0 DI S TRI B UTI ON OF B ACT ERI A

larg e ly fro m th e so il lay e rs b e n e ath A g re at many diff e r e nt .

kinds o f bact e ria fi nd the ir way i nto th e air but a g r e at many ,

o f th e m a e u nabl e to withstand th e d e siccati o n to which th e y

r

a e s ubj e ct e d s o that c e rtai n f o rms di e o ff a nd o th e r s r e main

r , .

Thus th e r e is e stablish e d a mo r e o r l e s s charact e ristic bacte rio

l o gical fl r a Th e e xt r e me minut e n e ss o f th e bact e ria p e rmits
o .

th e m to r e main in th e air fo l o ng p e ri o ds o f tim e

r .

B act e r ia can no t b e di sl o d ge d fr o m a m o ist surfac e Th e y

'

wo uld no t b e bl o wn ihto th e air fo r in s tance fr o m th e su rface

, ,

o f wat e r T h e o nly way that wat e r bact e r ia can ge t i nt o th e

.

air w o uld b e to have th e m sh o t i nto it by th e bur sting f o

bubbl e s o f gas Thi s que stio n is o ne o f co n sid e rabl e practical

.

imp o r tance d ue to th e discus sio n wh i ch h as f re q u e ntly o ccu rr e d

,

in r e ga r d to th e bacte ria in s e w e r air Th e air fro m se w e rs .

i s u sually l o w in its ge rm c o nt e nt d ue to th e fact that th e ,

s e wage a nd th e wall s o f th e se w e r s a e m o i st Th e se co nsid

, ,
r .

e a ti ns
r o a s w e ll as th o e di s cu sse d e l se wh e r e
,
s l e ad to th e ,

b e li e f that s e w e r ai r is no t a s g r e at a facto r in th e pro duc

tio n o f dis e as e as was fo rme rly supp o se d I t is q uit e lik ly . e

that in cas e s wh e re it wa s supp o se d to b e th e ca u e o f di se a se s

it was m e r e ly a co nco mitant circumstanc e F urth e rm o r e .

,

th e bact e ria o f di se a se a re no t lik e ly to o cc u r in th e b r e ath

‘

O f th o se s u ff ri ng fr o m di s e a se fo r th e s am e r e a so n
e Th e y are .

ve ry f r e que ntly sh o t fro m th e mo uth o f di se a se d p e r so ns during

fi ts o f c o ughing and sn e e zing and to l e ss e r e xt e nt duri ng talk
, , ,

ing in littl e dro pl e ts o f sputum Wh e n th e se d ro p l e ts dry ”

,
.
,

th e n th e bacte ria in th e m may ge t into th e air .

 B ACTERI A OF THE AI R 2 9 1

Quan ti ta ti v e Di s tri b uti o n T h e numb .e r o f bact e r ia i n th e

air va ri s with diff r e nt co nditi o ns as h as al r e ady b e e n indi

e e ,

ca te d . Th e y a e fo und to b e ve ry spa r se in th e ai r o ve r sno w

r

co ve r e d mo untains a nd o ve r la rg e b o di e s o f wat e r Th e y a re
, .

co mpa r ative ly fe w in th e c o untry ai r a nd ve ry abu ndant in

th e air o f citi e s o r o th e r part s o f th e c u ntry wh e r e th e r e i s a
,
o

gr e at de al o f dust Th e ir di st ributi o n wil l de p e nd up o n th e

.

charact e r O f th e su r face o f th e so il th e am o unt and di re cti o n ,

o f wi nd th e acti o n o f th e su nlight e tc
, ,
.

S e a s o nal Di s t i b uti o n r S e a so n h a s a n i nflu e nc e up o n

.

th e numb e r o f bact e ria in th e air C o ndi tio ns whi ch fav o r

.

dryn e ss a nd h e nc e dust i nc re ase th e numb e r o f bacte ria

, ,
.

R ai ns no t o nly k e p d o wn th e du s t but th e y wa sh th e air f r e e

e

fr o m bacte ria Th e sam e is true o f sn o w so that wh e n th e

.
,

air is wa sh e d by rain o r sn o w it is much fre e r f ro m bacte ria

than at o the r tim e s .

Th e S pe ci s o f B acte ia i n th e Ai r
e r B act e ria f o u nd in th e
.

air are mo stly sapl pp hyte s and are harml ss Th e r e a e ce r e . r

tain parasitic bacte ria ho w e ve r which may b e ai r b o rn e

, ,
-
.

One O f th e b e s t e xampl e s o f ai r b o rn e di se a se pr o duci ng bac

-

,
-

t e ria is B ct tub e rcul o si s That thi s ge r m ca n b e tran sp o rt e d

a . .

thro ugh th e air fro m o ne individual to ano th e r is b e yo nd

q ue stio n and mo st p e o pl e b e li e ve that thi s di se as e is trans
,

m itte d larg e ly if no t e nti r e ly thr o ugh th e ba ct e ria whi ch ge t

, ,

into th e air fro m dri e d sputum Th e w o rk o f C o rne t and .

o th e r s has sh o wn e xp e rim e ntally that thi s i s p o s ibl e Re s .

ce nt wo rk by a co nsid e rabl e numb e r o f inve stigato rs fo llo wing ,

2 9 2 DI S TRI B UTI ON OF B ACT ERI A
th e le ad o f R ave n e l has sho wn that th e di s tributio n o f tub e r
,

cul o si s may b e acc o unt e d fo r in o th e r ways to a c e r ta i n e xt e nt,

at l e ast B act d iphth e ri a may b e dist ribute d th r o ugh th e air

. .
,

a nd t hi s g e rm can und o ubt e dl y liv e fo r l o ng p e ri o d s O f tim e in

th e air .O th e r di s a s e s that w e r e f o rm e r ly s upp o s d to b e air

e e

b o rn e such as i nflu nza are no w acco unte d fo r so far as th e ir

,
e , ,

di stributi o n is co nc e r n e d in o th e r way s I n a c o nsid e rabl e

,
.

numb e r o f o th e r di se as e s such as th e acut e e xanthe mata

, ,

which include s smallp o x m e a sl e s and s carl e t fe ve r e vi

, , ,

de nce se e m s to p o int to th e air as a ve hi cl e o f trans mis sio n .

I t sh o uld b e n o t e d that in th e pa s t it has b e e n larg ly tru e e

that we have attribut e d to th e air an imp o rtant rOl e wh e n we

have b e e n larg e ly o r co mpl e te ly igno rant o f th e cause and
m e ans o f di stributio n o f th e s e di se ase s As o ur kn o wl e dge .

has b e c o m e mo r e e xact we have fo und that th e air h as had l e ss

,

to d o with it than wa s f o rm e rly supp o se d Wh e th er thi s is.

g e n e rally true o r no t o nly tim e can t e ll B ut we d o no t have

.

at th e p r e se nt tim e any co nv i nci ng e vid e nce that any di s e ase

is e ith e r in wh o l e o r in part large ly spr e ad thr o ugh th e air .

O f th e sapr o phyt e s which p e r si s t in th e air fo r l o ng p e ri o d s o f

tim e and co n s titute v e ry larg e ly th e bact e rial fl o ra o f th e air ,

th e micr o co cci a re th e m o s t pr o m in e nt Am o ng th e s e are

.

many brilliant chro mo g e nic fo rms I n addi tio n to bact e ria

.

th e re is always pr e s e nt a co nside rabl e numb e r o f mo ld Sp o re s

and ye ast ce lls .
 C HAP T ER XX VI I
B AC T ERI A OF WATER AN D S E WAGE

I nt Wate r always co ntain s bacte ria I t was

ro d ucti o n — . .

supp o se d s om e y e ars ago that such wate rs as di stil le d wate r

a nd that fr o m d e e p w e ll s w e r e s t e ril e but sinc e B u b o n S and e r
, r

so n in,
87 sh o w e d that many bacte ria e xi st e d in di still e d
1 1,

wate r it has b e e n r e co gni z e d that th e r e are always bacte ria

,

pr e se nt in a wate r supply M any o f the se bacte ria are p e r

.

fe ctl y harml e s S O far as man is c o nc e rn e d and e xist in th e

s ,

wat e r co mp o sing a natural flo ra N o n e o f th e bacte ria f o und

.

n o r mally in wat e r a e ha r mful to man S o me o f th e se h o w

r .
,

e v e r may cau se tr o ubl e by gr o wing in th e wat e r pip e s fil ling

, ,

th e m up as in th e cas e o f th e C re no thrix C e rtain bacte ria

,
.

that cause a g o o d de al Of difli cul ty may also ge t into th e wate r ,

e sp e cially fr o m human s e wag e One o f th e gr e at pr o bl e ms

.

o f sanitati o n is to pr o t e ct th e wat e r suppli e s fr o m e wag e co n s

ta m ina tio n o r in cas e th e y b e co m e co nta m inat e d to purify

,

the m .

N um b e r o f B acte ri a i n Wate r Th e numb e r vari e s fro m

.

a ve ry fe w bact e ria p e r cubic ce ntime t e r in th e wate r fro m

d e e p we lls o r springs to a ve ry gre at many in a surface wate r ,

e sp e cially if thi s is c o nta m inat e d With s e wag e A go o d ide a .

2 93
2 94 DI S T RI B UTI ON OF B AC TERI A
o numb e r o f bacte ria in di ffe r e nt wate rs may be o bt ai n e d
f th e

fr o m th e fo llo wing tabl e :

N UM B ER OF BA T
C E RI A IN VAR I O U S WATE R S

SO U RE
C N UM B ER OF BA ER A CT I P E R cc .

t id
ou s Pari M iq u l
e o f s, e .

I 9 i n id
s f Pari M iq u l
e o s, e .

3 t 3
o J an w ki
00 , o s .

gla i r in N rwa y S h m
ce o , c e l ck .

M adi n W i n in
so ,
s co s .

B uj wid .

F o unt in a .

to 2 000 , ak
L e Men d ta Wi o ,
sco ns in .

Lak e Ge n va e ,
n ar h r
e F s o e , o l a nd Duna nt .

Lak e Ge n va e ,
m id dl e .

to S e ne i .

R hi n e .

R h on at Ly n e o s .

K h oc .

to 61 hall w w ll R b n r
2 0, s o e ,
u e .

t 4
o d p w ll L nd n Fra nk lin
00 , ee e s, o o ,
.

t 33 d
o p w ll V i nna K wal ky
,
ee e s, e ,
o s .

t 5
o M adi n W i
0, n in so ,
s co s .

0 t 6
o p ri ng Fra nk f rt n th M ain
0, s s, o -
o e -
.

9 t 34 5
o p ri ng Zuri h C ra m r
2 ,
s s, c ,
e .

Th e r o f B a cte ria i n Wa te r
O i gi n S o m e bact e ria c o m e .
—

fro m th e air b e i ng wash e d fro m th e S ky with th e du st Th e

,
.

maj o rity o f bact e ria ge t into s urface wate r fro m so il wa shi ngs .

Th e numb e r which ge t i n d e p e nds up o n th e c o ndi ti o ns sur

r o undi ng a b o dy o f wat e r ; fo e xample a g re at many mo re r ,

bacte ria ge t in fro m a cul tivate d so il whe n thi s make s up th e

 B ACTERI A OF WAT ER AN D S E WAGE 2 95

wate rshe d than fro m a vi rgin so il ; m o r e a nd pa r ticul a rly ,

mo re dange ro us kinds wh e n th e wate rsh e d is i nhabite d than

,

whe n it is uninhabit d M any o f th e bacte ria whi ch ge t i nto th e

e .

wate r a e capabl e o f gro wing th e r e and f o r mingwhat i s kno wn

r

a s th e wat e r ba ct e ria o r th e no rmal bact e rial fl o ra o f th e wat e r

, .

M any o f th e se bact e ria a e abl e to gr o w e ve n in wat e r s that

r

co ntain ve ry littl e o rganic matte r C e rtain sp e cie s o f bact e ria

.

can practically always b e fo und in a wat e r supply I n S pite .

o f th e fact h o w e v e r
,
that th e y se e m to b e we ll adapte d to
,

life in wate r th e y d o no t gr o w b e ydnd ce rtain limits One .

wo uld naturally thi nk tha t if bacte ria e xiste d in wate r fitte d

fo r th e ir d e v e l o pm e nt that th e y w o uld s o o n incre as e to e no r

mo us numb e rs but this is no t true P r o t e cte d fr o m th e

, .

co ntami natio n which co me s fro m th e s o il th e bacte rial co nte nt ,

o f a surfac e wat e r r e main s qui t e c o nstant and w o uld giv e a

co mpa r ative ly l o w c o unt Wh e n the s e same wate rs a e se pa

. r

rate d fro m th e ir natural co nditio n as fo r instanc e whe n th e y

, , ,

a e b o ttl e d r e ady fo r analy si s th e numb e r o f bact e ria in th e m

r ,

inc re ase s r apidl y to an e n o rmo us e xte nt Thi s i ndi cat e s that .

the r e is in natural wat e r s a ge rm icidal sub stance whi ch hinde rs

th e e xc e ssiv e multiplicati o n o f wat e r ba ct e ria J u s t what this .

bacte ricidal pro p e r ty is d ue to is an o p e n que stio n .

Th e S p e ci e s Of B a cte ri a i n W a te r C e rtain bact e ria .

find th e ir Optimum co ndi tio ns fo r de ve l o pme nt in wate r .

Th e se are th e bacte ria whi ch are o rdi narily sp o ke n o f as wate r

bacte ria Th e y have b e e n e xte nsive ly studi e d by a numb e r
.
’

o f inve stigat o rs but are no t imp o rtant ,so far as we kn o w in

, ,
2 9 6 DI S T RI BU TI O N OF BAC TERI A
a ff e cting th e p o table ne ss (drinkabl e n e ss) o f th e wate r Othe r .

kinds o f bacte ria mak e th e ir way into th e wate r and u sually

quickly di sapp e a r Tho s e which are mo st imp o rtant in this
.

co nne ctio n are fro m se wage in g e n e ral a nd human se wage in ,

particular Amo ng th e se se wage bact e ria whi ch ge t into wate r

.

a re B . co li B typh o sus B dyse nte ria and M sp co mma

,
.
,
.
,
. .

B acillus c o li is th e c o mm o n inhabita nt O f th e huma n int e s tin e .

I t i s al so f o u nd in th e i nt e s tin e o f o th e r a ni mal s particularly ,

tho s e which are do me sticate d I ts pre se nce in surface wate rs .

in ve ry small numb e rs fo r e xampl e l e ss than o ne to a cubic

, ,

c e ntime te r may p o s sibly b e acco unte d fo r by th e pr e se nce

,

o f th e e xcr e ta o f animal s no t cl o se ly as so ciat e d with m a n .

B ut whe n pre s e nt in gr e at e r numb e rs than indi cate d ab o v e ,

the y c o nstitute by practically c o mm o n co n se nt e vide nce

, ,

o f dang e r o us s e wag e c o ntaminati o n Th e p r e s e nc e O f B c o l i . .

in wate r may th e n b e re ga r de d a s an i ndicatio n o f p o llutio n

, , .

I n fact its pr e se nc e o r abs e nc e in wat e r a s d e t e rmin e d by an

, ,

analysis is use d ve ry large ly in f o rming an Opini o n in r e gard to

,

th e p o tabl e n e ss o f a wat e r supply N o t infr e qu e ntly th e di s .

,

e as e pr o ducing bact e ria m e nti o ne d ab o v e B typh o s u s B dys

-

,
.
,
.

e nt e ri a and M sp co mma ge t into s e wage p o llut e d wate rs

.
,
-

and ve ry s e rio us e pide mics are fr e que ntly trace d to se wage

p o llute d wate rs I n th e case o f typho id f e ve r pro bably at
.
,

l e ast sixty p e r ce nt o f th e cas e s are wat e r b o rn e One o f th e -

.

first typho id e pide mics to b e care fully s tudi e d wa s that which

o ccurr e d in L a wse n S witz e rland in 187 2 ,
He re fro m Augu s t,
.
,

to O ct o b e r a hundr e d and thi rty cas e s o f typh o id f e ve r

,
 B ACT ERI A OF WATER AN D S E WAGE 2 97

de ve lo p e d in a p o pulatio n o f 7 80 Thi s e pide mic was ve ry

.

de finit e ly trac e d to th e wate r supply co nn e cte d with an unde r

g r o u nd st r e am which b e cam e c o ntaminat e d with th e di s

cha rge s o f typho id fe v e r pati e nts I n 1885 th e P lymouth

.
,

e pid e mic o ccu r r e d whi ch h as b e c o m e a clas sical e xampl e o f a

wate r b o rne typho id e pide mi c I n that y e a r mo r e than o ne

-
.
,

th o usand case s de ve l o p e d in a ve ry sho rt p e ri o d o f time o ut o f a

p o pulatio n o f 8 ( F ig
000 . I t was c o nclusive ly sh o wn that

th e dis e as e was cau se d by th e di s char e s o f a typh o id pati e nt

g
livi ng and nurs e d On th e banks o f th e str e am fro m whi ch th e
wate r supply o f th e city was take n M any o th e r e pide mics
.

have b e e n de finit e ly t r ace d to an infe cte d wate r supply and th e ,

tim e must so o n co me wh e n no city will think o f taking wate r

fo r dri nking purp o s e s whi ch c o m e s fro m a s o urc e that is no t

pro p e rly pro te cte d fro m se wage co ntam inatio n Th e dange rs .

that co m e to citi e s taking th e ir wate r supply fro m an infe cte d

s o urce is w e ll illus t r at e d in tho se citie s situate d o n th e M o hawk
and Huds o n rive r vall e ys .

Asiatic ch o l e ra i s als o a wat e r b o rn e d i se a se

-
F o rtunat e ly
.
,

Asiatic ch o l e ra has b e e n k e pt o ut o f Am e rican and Euro p e an

co untri e s fo r many y e ar s and it is ho p e d that it will n e ve r re
,

turn in e pide mic fo rm I f it do e s ho we ve r it wil l trave l

.
, , ,

as it h as in th e pas t large ly thr o ugh wate r That it is a wate r

,
.

b o rne di se ase is ve ry strikingly sho wn by th e facts de v l o pe d e

in G e rmany in 89 3 1 . H amburg and Alto na a re two co n

ti guo us citi e s . Hamburg t o o k its wate r supply f ro m th e
R ive r Elb e . B e cau se it had its intak e situat e d s o m e what
2 9 8 DI S T RI B U T I O N OF B ACT ERI A

FI G . 60 —
. P lym th pid m i
ou e e c of typh id f v r
o e e .
 B ACTERIA OF WAT ER AN D S E WAGE 2 99

ab o ve ity th e wat e r was u se d in a raw stat e Alt o na

th e c , . .

o n th o th e r ha nd wa s o blig e d to tak e its wat e r if at all f r o m

e , , ,

rive r aft e r th e s e wage o f p e o pl e had b e e n p o u r e d

'

th s ,

i nt it Th e wat e r was th e r e fo re filt e r e d I n 893 during th e

o . . 1 ,

ch o l e ra e pide mic th e ca se s we re ve ry f r e qu e nt in Hambu rg

, ,

whe r e awwat e r was us e d and ve ry fe w in numb e r in Alto na

r , ,

wh e re filte re d wate r was use d Th e cha ract e r o f th e wate r .

supply was th e o nly diffe re n ce b e tw e e n th e se two place s which

c uld acco u nt f th e di t ributi o n o f th e case s To co mpl e te
o or s .

th e p r o o f that thi s Hambu r g e pid e m i C was wat e r b o rne in th e -

wi nte r aft e r th e di e a se had di sapp e a re d f ro m Hamburg

,
s ,

it bro k e o ut e xplo sive ly in Alt o na b e cau se a s was lat e r , ,

a c r tai ne d th e sand filt e r had crack e d o n f r e zing all o wing

s e ,
e ,

th e unfilt e r e d wat e r to pa s s i nt o th e city main .

W a te r An al ysi s T h e e ffici e ncy o f th e bact e ri o l o gical

.

wat e r analysis is b e co ming m o re and m o re e vide nt as its use in

c r e a se s and th e t e ch ni que b e c o me s p e r fe cte d I ts value sup .
,

p l cm e nting that o f a ch e mi cal analy si s was fi r st d e m o n s t r at e d ,

by F rankl a nd s w o r k n th e filt e r e d wate r o f Lo ndo n C h e mi

’
o .

cal analy si s was u n bl e to d e t ct th e pu rific tio n ac co mplishe d

a e a

by a sand filt e r a nd the se filte r s w e r e b u t to b e disca rde d

,
a o

in Lo ndo n a s i ne ffi cie nt wh e n F r ankland sho w e d that so far

as th e re mo val f ba cte ria was co nce rne d th e i r fficie ncy w
o ,
e as

v e ry high i e o v r ni ne ty nine p e c e nt F ilte r s can o nly b e

,
. . e -
r .

e fli ci e ntl y c o nt r o ll e d by p r o p e r ba ct e r i l o gi cal a naly e I n th e o s s .

sanita ry e xami nati n o f wat e r th e bact e ri o l o gi cal de te rmina

o ,

tio n s are b e co ming r e co gniz e d as o f gre at value I n a bacte ri o .

3 00 DI S T I R B UTI ON OF B AC T ERIA
l o gical wat e r analy sis th e numb e r o f bacte ria p e r cc is .

d e t e r mi ne d by plate cultur e s I t is imp o ssibl e to judg e th e

.

cha r acte r o f a wate r by th e numb e r o f bact ria it co ntain s e

al o ne but th e bact e rial co nt e nt is f r e qu e ntly a matte r o f im

,

p o r tance take n in c o nn e ctio n with o th e r facto r Th e numb e r s .

o f di ff e r e nt ki nd s o f bact e r ia pr e se nt in wat e r i s al so imp o rtant ,

but as in th quantitative analysis do e s no t give info rmatio n

,
e ,

whi ch can b e u se d al o n e I t is v e ry ra r e that B typho sus o r

. .

M sp c o mma can b e d e t e ct e d i n th e wat e r T h e r e a so n fo r

. .

thi s is amo ng o th e rs that th e analysis is usually no t at

, ,

t e mpt e d until co n side r abl e tim e has e lap se d si nce th e intr d uc o

tio n o f th e se bacte ria int th e wate r The se bact e ria live fo r

o .

o nly a sh o r t tim e i n wat e r u nd e r o rdinary co nditi o n s a nd ,

th e r e fo r e have pr o bably di sapp e are d o r b e co m e e xtr e m e ly

,

fe w in numb e r at th e tim e o f an a naly sis and agai n it is a ,

ve ry di fli cul t matt e r to se parat e B typho sus fro m o th e r cl o se ly

.

r e l at e d bact e ria What is so ught fo r in a wat e r analy si s is th e

.

p re se nce f B c o li Thi s is usually do n e by i ntro ducing th e

o . .

wat e r to b e e xami ne d in va ryi ng am o unt s f ro m o ne t e nth O f a

cc to te n cc i nt o sp e cial cultu r e m e dia Th e cultu re m e dium
. . .

m o s t g e ne rally u se d is d e xt r o se b o uill o n in f e rm e ntati o n tub e s .

T h e pr e se nc e o f thi s bact e rium is d e t e rmin e d in what is f e r

q u e ntly call d th e pr e s umptiv e t e st by i ts ability to f e r m e nt th e

e

suga r in th e d e tfo se bro th Of th e f e r m e ntatio n tub e s with th e

x
'

pro ducti o n o f c e rtai n amo u nts o f gas o f a d e fi nite f o rmula ,

th e p r o du cti o n O f e d c o l o ni e s o n lact o s e litmu s agar

r th e ,

no nl iq ue fa ctio n o f g e lati n a nd th e p ro ductio n Of indo l

, .
 B ACT ERI A OF WATER AN D S E WAGE 0
3 1

In wat e r a nalys e s an imp o r tant pa r t i th e co ll e ctio n and

,
s

t r an spo rtatio n o f th e sampl s Th e sampl e m ust b e co ll cte d

e . e

in st e ril e b o ttl e s and in s uch a way a s to avo id co ntami

natio n and e xami natio n must b e made e ithe r imm e diate ly
,

o r th e sampl e m u s t b e k e pt pack e d in i ce so a to pr ev e nt th e s

chang e s whi ch o th e rwis e tak e place .

Th eP uri fi ca ti o n o f W te r AS th e p o pulati o n o f th e
a .

c o u nt ry i nc r a s s th e ne ce ssity o f purifi catio n o f wate r

e e ,

supplie s fo r c rtain p o r ti o ns o f th e p o pulati o n at l e as t b e

e , ,

c o m e s imp e r ative T h e pu rificati o n O f wate r i s at th e b o tt o m

.

a ba ct e ri o lo gical p ro ble m I t may b e acco mpli h e d in a num

. s

b e r o f di ff e r e nt way s ; by s e dim e ntati n whi ch may tak pla ce o ,

e

naturally in so m e wat e rs o r a s a r e sult Of th e additi n o f

,
o

ch e mical s in o th e r s I t is se cure d al so and mo st f r e q u ntly

.
,
e ,

by filt r ati o n m th o d s Thi s is a cco mpli she d by what are

e .

kno wn as co nti nuo us sand filte r s u se d la rge ly in E u r o p e , ,

by m e ans o f m e cha ni cal sand filte rs d e ve l p e d and wide ly ,

o

u se d in Ame rica a nd by i nt e r mitt e nt sand filte rs d e ve l o p e d in

, ,

Am e rica and ve r y suc ce ssfully u se d o n g r o s ly p o llut e d wat e r s s .

I n th e c o nti nu u s sa nd filt e r th e e ffi ci ncy d e p e nds up o n th e

o e

fo rmatio n o f a slime lay e r o n th e surface Of th e filt r s that e ,

o

this filt e r is r e ally a livi ng filt e r I n th e m e chanical filte r .

,

th e e fli cie ncy i s d ue to a p r e cipitat e pr o duc e d o n th e sa nd by

th additi o n O f ch e mical s to th e wat e r

e I ts adva ntag o ve r . e

th e co nti nu o u s filt e r c o m e s f r o m th e fact that it w o r k s mu ch

m r qui ckly a nd h e nc e o c cupi s l s s space I n th e inte r

o e ,
e e .

mitte nt sand filt e r th e pu rifi cati n de p e nd s larg e ly up n th e

,
o o
3 02 DI S T RI BUTI O N OF B ACT ERI A

actio n ofitrifying bact e ria which de ve l o p in th e int e r

th e n

s ti ce s O f th e filt e r b e tw e e n th e fl ushi ng wh e n th e mate rial s

is fill e d with air .

S e wage B y s e wag e is m e ant th e liqui d wa s t e fr o m laun

.

dry kit ch e n and wat e r cl o se t I t c o ntains o rganic matt e r

, ,
.

a nd e n o r m o u s numb e rs O f bact e ria Wi nslo w f und . o

p e r cc i n wi
. nt e r and p e r cc in s umm e r M a ny . .

di ff e r e nt sp e ci e s o c cur Thr e e g ro up s a e o f pa r ticular i nt e r

. r

e st One g ro up i s co mp o se d O f th e putr e factiv e ba ct e ria

.
,

B vulga r i s e tc
. Ano th e r g r o up is co mp o se d o f tho s e bac
,
.

t e ria which are characte ri stic o f s e wag e — such fo r ms as B ,

.

co li B cl o ac a B w e lchi i and S tr pyo g e n e s a re in clude d

,
.
,
.
,
. .

Th e t hi rd gr o up a nd fr o m a public h e alth s ta ndp o int th e

,

m o s t imp o rtant is th e di se as e pro duci ng bacte ria as B

,
-

, .

typh o s us M sp c o mma and B dyse nte ia

,
.
,
. r .

The P u ifi a ti n f S e w ger c
— Th e s avag e and barbarian
o o a .

a re littl e co nce rn e d with thi s p r o bl e m Th e ir r e fuse is .

thro wn in a pil e and wh e n it b e co me O ff e nsive the y mo ve

,
s

on . C i v iliz e d m a n m o ve s th e r e fus e In th e case o f singl e .

dw e llings and small co mmuni ti e s thi s h as b e e n do n e by bury

ing o r putti ng it i nt o a c e ssp o o l I n la r g e r co mmu niti e s it .

has b e e n wa sh e d i nto an adjace nt b o dy o f wat e r As th e .

p o pulatio n i ncr e ase s it is fo und that th e se m e tho ds are um

s ati sfact o r y fo r th e car e o f th e gr o wi ng vo lum e O f se wag e .

R e c e ntly bact e r i o l o gical m e th o d s o f se wag e pu r ifi cati o n h v e a

b e e n de ve lo p e d I n o rde r to pu rify s e wag e its o rgani c matt e r

.

must b e de stro y e d and th e path o g e ni c bact e ria which it may

,
 B AC T ERI A OF WATER AN D S E WAGE 0
3 3

co ntain kill e d Al l satisfacto ry m e th o ds a re fundame ntally

,
.

bacte rio lo gi cal Wh e th e r buri e d tr e ate d by th e dry e a r th

.
,
-

sy st e m th e ce ssp o o l th e st re am o b e tte r m e tho ds se wag e

'

, , ,
r ,

i s purifi e d o r r e nde r e d harml e ss by ba ct e r ia Th e m o de r n .

me tho ds mo st f re que ntly u s d a re th e int e rmitte nt fil tratio n

e
,

b r o ad irrigatio n and th e s e ptic tank and co ntact b e d me tho ds

, .

The irimpo rtance is in th e inve rse o rde r fro m that in which

the y a e e nume rat e d
r .

I nte m i tte nt fi l t ti o n is us e d succ e s s fully i n ce rtain l o cali

r ra

ti e s whe re th e co a r se mat e rial n e e de d in the ir co nstructi o n ,

such as g r ave l b ro ke n sto ne s o Cinde rs are available One

, ,
r , .

ac re pu rifi e s th e se wage fo o ne tho usa nd p e o ple Th e mo st

r .

favo r able lo cati o n is alo ng a rive r b e d The se b e ds altho ugh .

,

th e y must be much mo re e xt e nsive than tho se r e qui re d fo r

wat e r may co st l e ss S e wage may b e pu rifi e d by thi s me tho d
, .

so that it is indistingui shabl e fr o m pu r e wat e r e xc e pt p e rhaps ,

by th e amo unt o f so dium chlo ride Th e ae r o bi c bacte ria are .

' '

e sp e cially active in t hi s filt e r B o ad i i ga ti n o s e w ag e

. r rr o r

farmi ng is succ e ssfully p r a ctic e d whe re th e pro p e r so il e xist s

( dr y and p o ro u s) Th e amo unt Of land r e qui r e d is f r o m te n
.

to tw e nty tim e s as g r e at a s is r e quir e d by th e i nt e r mitt e nt

fil te r Th e s e wag e i s run do wn o ne S ide o f th e fi e ld in a di t ch

.
,

and in lat e ral di t ch e s o furro ws it flo ws th ro ugh g ro wing

r

c ro ps to wards a st re am Th e se wage is app l ie d i nte rmitte ntly

.

a s ab o ve Thi s me tho d i u se d in B e r li n and P a ri s a nd also

. s ,

e xt e nsiv e ly in E ngla nd a nd to so me e xt e nt in N o rth Am e rica .

I t is e co no mical sin ce s e ve ral cro ps can b e r aise d in s t e ad o f

,
 0
3 4 DI S T RI BUTI ON OF B ACT ERI A
o ne . It is p e rfe ctly satisfact o ry fro m a sanitary standpo int .

Th e s e p ti c ta nk was fi r st bui lt by C ame ro n o f E ngla n d Thi s .

tank is a tight ce me nt chamb e r thr o ugh whi ch th e se wage

sl o wly fl o ws Th e p o i nt o f e nt r y is b e low th e surface so a s
.
,

no t to di sturb th e sc u m that so o n fo rm s a nd no t n e a r n o ugh

,
e

th e b o tt o m to int e rf e r e with th e se dim e nt Th e r at e O f fl o w

.

fo r th e s e wag e is s u ch that it r e qui r e s ab o ut tw e nty f o ur -

h o urs to pa ss th ro ugh th e tank Th e ana e ro bic bact e ria.

find co nditio ns favo r abl e fo r th e ir a ctivity and g radually

liqu e fy th e s o lid matt e r Thi s r e ducti o n may amo unt to
.

80 p e r c e nt .Th e wo rk o f th e a na e r o bi c o r putr e fa ctiv e bac

t e ria in th e s e ptic tank is suppl e me nte d by th e wo rk O f th e

a e ro bic bact e ria in th e co ntact fi l te be ds Th e se a e fil l e d
r . r

with co arse mate rial such as Cinde rs a nd co ke Th e b e ds a re

, .

co ve r e d with th e e ffl u e nt fro m th e se pti c tank fo a fe w h o ur s r

and are the n drain e d and all o we d to stand a fe w ho urs Thi s .

pr o ce ss is r e p e at e d se ve ral tim e s a day Thi s me tho d o f .

s e wag e di sp o s al is mo st satisfacto ry and is Wide ly use d at th e

pr e s e nt time Th e amo unt o f land r e quir e d is small th e
.
,

att e ntio n r e quire d slight a nd th e d e g r e e o f p u rificati o n high

, .

Th e use o f ch e mical s s uch as chl o r id e o f lim e fo r th e pur

, ,

p o se o f ste rilizing th e e fflue nt Of th e s e ptic tank o r p o llute d

wat e rs is b e ing advo cat e d This m e th o d may b e o f th e
.

gre at e st s e rvice in tre ati ng wat e r suppli e s duri ng e pide mi cs

o f wat e r b o rn e di s e as e s e sp e cially typh o id f e ve r
-
, .
 C HAP TE R X X VI I I

B AC TER I A OF M I LK AN D I TS PR O D U CT S
I ntro d ucti o n . M il k is
m o st imp o rtant o f th e
o ne Of th e

fo o d pro ducts I t is an e xce ll e nt fo o d fo r man and it is just

.
,

a s g o o d f o o d fo r mi cr o b e s a nd si nc e milk i s usually s o ld in i ts
,

ra w s tat e it happ e ns that it f e q he ntl y co ntain s e n o r m o us

r

numb e r s O f ba ct e ria S o m e o f th e se bacte ria pro duce no

.

change what e ve r in th e milk and th e i r p r e s e nc e is a matt e r o f

,

indi ff e r e nce C e r tain o the r bact e ria pro duc e change s in th e

.

milk its e lf causing it to sp o il The se are unde sirabl e al

,
.
,

tho ugh th e y may in no way a ff e ct th e h e alth o f th e co nsume r .

S till an o th e r clas s O f bact e ria so m e tim e s f o und in milk a re

di se a se pr o duci ng and agai nst th e pr e s e nc e Of th e s e in mil k

-

th e c o nsum e r sh o uld hav e ab so lut e p r o te cti o n .

Th e B a cte ri a in th e Ud d e r I t was fo rm e r ly supp o s e d that

.

milk in th e udde r o f th e co w was ste ril e but we no w kn o w ,

that it is practically imp o ssibl e e ve n und e r as e ptic co nditio ns

, ,

to se cure s t e ril e milk Th e numb e r o f ba ct e r ia in th e m ilk

.

tak e n unde r th e b e s t co nditio ns vari e s fro m two hundre d to a

tho usand ba cte ria p e r cc .

Th e N um b e r f B acte ri a in M k t M il k
o Th e numb e r O f
ar e .

bact e ria fo und in ma rk e t mi lk va rie s gre atly f ro m a fe wtho u

sand p e r cc to many millio n p e r cc I n what may b e co n
. .

305
3 0 6 DI S T RI BU TI ON OF B ACTERI A
si d e re high e st grad e Of milk o n th e ma r k e t name ly ce r
d th e , ,

ti fi e d milk th e numb e r is usually limit e d i n th e c o ntract to

,

te n th o u sa nd bact e ria p e co and practically a c o n sid e rabl e

r , , ,

amo unt o f thi s milk is de live re d to th e h o m e o f th e co nsum e r

with o nly a small fracti o n o f thi s numb e r pre s e nt Othe r .

milk co ll e cte d unde r l e ss ide al c o nditio ns but c o nstituting ,

what wo ul d b e co nside r e d a go o d milk h as t e ns o r hundr e ds o f ,

tho usands o f bact e ria p e r cc A numb e r o f citie s have limit e d

.

th e numb e r o f bact e ria p e rmissibl e in g o o d milk to two

hundre d and fifty tho usand and o th e rs to fi v e hundr e d ,

tho usand .

A g o o d ma ny milks pro duc e d u nd e r p o o r c o nditi o n s o r ,

ke pt aft e r b e ing co ll e cte d fo r l o ng time s unde r unfavo r abl e

, ,

c o nditi o ns co ntain e n o rmo us numb e rs o f bact e ria Fo r in

,
.

stance P ark fo und in N e w Y o rk City an ave rage in twe nty

,

sampl e s bacte ria p e r cc a s th e milk was re ce ive d in

.

th e city .

F ro m t e n e m e nts midwint e r , ,

F r o m w e ll to d o di s tricts midwi nt e r
- -

, ,

F r o m t e ne m e nts S e p t , ,

F r o m w e ll to d o districts
- -

Th e S o urce o f B a cte ri a in M il k — Att e nti o n h a s alre ady.

b e e n cal l e d to th e initial bacte rial co nte nt o f mi lk a nd it ,

p e rhap s o ught to b e add e d that th e atte mpt is so m e time s made

to r e duc e thi s by di s ca r di ng th e f o r e milk o r th e fir s t fe w cubic ,

ce ntim e t e rs d r awn fro m e ach t e at but whi l e the re are m o re ,

bact e ria in th e fo r e milk than in o th e r parts o f th e milking ,

 B ACTERI A OF MI LK AN D I TS PR O DUCT S 3 07

th e diffe re nce is no t so g r e at that th e di scardi ng o f th e f o re m ilk

make s a ny gr e at imp ro ve me nt in th e ge ne r al characte r o f th e
who le mi lki ng S O that whe the r th e fo re milk is r e tain d o r
.
e

di s card e d is a m att e r o f l ittl e imp o rtanc e in th e t o t al numb e r

o f bact e ria .

C o ntam ina ti o n fr o m th e Ani m al Th e c o at o f th e animal

.

is a frui tful s o urc e Of bact e ria i n milk Dirty animal s intro

.

duce an e n o rmo us numb e r o f bacte ria Wh e n particl e s o f .

manur e a re di sl o dge d and fall into th e milk duri ng th e pro ce ss

O f milking th e y carry with th e m gr e a
,
t numb e r s o f bact e ria and ,

th e s e are o f an e sp e cial ly unde si r abl e ki nd Wh e n th e animals .

a re no t k e pt cl e a n th e numb e r o f bact e ria whi ch fall i nt o th e

,

pail is gre at Eve ry particl e Of dust h as a numb e r o f bacte ria

.

ridi ng o n it and th e same thi ng is tru e o f th e hai r s o f th e ani

,

mal This so urce O f co ntaminatio n can b e pr e ve nt e d to a co n

.

sid e rabl e e xt e nt by in th e first plac e k e e pi ng th e animal s cl e a n

, , ,

and whe re p o ssibl e currying th e m fr e que ntly at s o me o the r

, ,

tim e than just b e fo r e milking k e e ping th e hair o f th e hi nd

,

quart e rs clipp e d sho rt and furth e rmo re by mo iste ning th e

,

udde r and flanks o f th e co ws with a damp cl o th just b e fo re mil k

in g . I t is als o d e sirabl e to u s e a small t o pp e d mi lk
-
pail and ,

many o f the m are no wso co n st r ucte d that th e y p re ve nt larg ly ,

e ,

th e e ntranc e o f bact e ria and dirt into th e pail during th e

pro ce ss Of milking Th e use o f th e se pails also p re ve nts th e

.

e ntranc e o f bact e ria fr o m th e n e xt so urc e to b e co n s id e r e d .

Th e Entrance o f B acte ri a i nto th e M il k fro m th e Air If .

milk is to b e pro duce d unde r as go o d co ndi tio ns as p o ssibl e ,

3 08 DI S TRI BUTI O N OF B ACT E RI A
it is de sirabl e that th e air o f th e stabl e wh e r e th e milki ng is ,

do ne sh o ul d b e as fr e e fr o m bact e ria as p o s sible This m e ans

, .

that th e co ws sho uld no t b e fe d with d ry f o o d just b e fo r e o r dur

ing th e milki ng tim e si nc e th e shaking up o f th e hay and o th e r
,

dry f o dde r is th e m e ans o f g e tting into th e air a g r e at amo unt

o f du s t and i ts c o nc o mitant bact e r ia Thi s ca n b e pre ve nt e d
.

by avo idi ng th e use o f dry f o o ds and by using th e small to p -

milk pail Th e cl e anlin e ss o f th e m ilki ng ute nsils is an o th e r

.

p o int to b e co nside r e d .

C l e anl i ne s s o f M il king Ute n s il s Th e milking ut e nsils n e e d

.

to b e S O c o nstruct e d that th e s e ams a re all r o undi ng so that it ,

is imp o ssibl e fo r milk o r milk and wat e r to r e main in th e

, ,

se ams by capillary attracti o n C ar e is usually e x e rci se d no w

.

by th e manufacture rs to se e that all o f th e se ams are w e ll

ro unde d o ut with s o lde r Ut e nsils o ught to b e no t o nly
.

care ful ly cl e an e d but th e y must b e all o we d to dry and the y

, ,

o ught to dry c o mpl e t e ly v e ry S h o rtly aft e r th e y hav e b e e n

wa sh e d Othe rwis e th e wat e r which is l e ft particul a rly if

.
,

the re is quite a littl e mi lk in it affo rd s a cul ture m e dium fo r

,

th e gr o wth o f bact e ria and it may w e ll happ e n that b e tw e e n

,

milkings th e wat e r l e ft in a milking ute n sil may lite rally te e m

with bact e ria and wh e n th e se sam e ute nsils are use d in milki ng
, ,

th e y add v e ry co nsid e rably to th e ge rm co nte nt O f th e milk .

Th e M il ke r Th e milk e r a ff e ct s th e bact e rial c o nt e nt o f

.

milk in mo r e ways than o ne I n th e fir st place th e dirty suit

.
,

o f cl o th e s o r a suit us e d in th e s tabl e o r in th e gard e n may

, ,

intro duce a go o d many bacte ria and fo r this r e aso n in th e ,

 B ACTERI A OF MI LK AN D I T S PR ODU CT S 3 9
0

pro ducti on of a high grad e m ilk th e milke r sho ul d have a cl e an

-

suit o f cl o the s I n many dai ri e s th e milke r w e ars a whi t e

.

duck suit cl e an at e ach milki ng o r at l e ast o nce a day I n

, , .

o th e r ca se s sp e cial s uits a re w o rn but a re no t was e d s o fre

h ,

qu e ntly Th e hands o f th e mi lke r may e ffe ct th e bacte rial

.

co nt e nt O f th e mi lk Th e r e is a habit am o ng th e mi lkme n o f
.

mil ki ng a fe w st re ams o n to the ir hands and th e n milking we t ,

as it is call e d Thi s is a di sgu sting practice and sho uld b e

.

disco ntinue d Th e milkman o ught to wash his hands b e fo r e

.

i
b e ginnin g to mi lk and in so m e dair e s it is do ne b e fo re milking
,

e a ch co w . F o th o s e wh o Obj e ct to milki ng dry th e hands

r

may b e mo ist e n e d with vase lin e I t app e ars al so that s o me

.

milke rs ca n ge t a much b e tt e r mi lk fro m th e same co w und e r

th e sam e co nditi o n s tha n o th e r s F o r e xampl e it has b e e n
.
,

S h o wn that dairy s tud e nts milk e d m o r e e ffi ci e ntly than th e

o rdi nary f o r c e th e di ff e r e nc e b e i ng d ue to th e fact no d o ubt

, , ,

that th e s tude nts unde rstand and pay h e e d to th e rati o nale o f

th e pr o c e ss .

Th e C are o f th e M il k —
C are in c o ll e cting mil k is n e c e ssary
.

in o rde r to k e e p th e initial ge rm co nte nt l o w but th e sub se ,

qu e nt care o f th e mi lk is qui te as imp o rtant M ilk is an ide al .

f o o d fo r micro b e s and the y gr o w with gre at rapidity if the y

,

a re giv e n c o n di ti o ns fav o rabl e to d e v e l o pm e nt Wh e n th e .

bact e ria are in th e milk ab o ut th e o nly thing that th e y n e e d in

,

o rd e r to gr o w r apidly is th e p r o p e r t e mp e ratur e Th e milk .

fu rni she s e ve rything e lse I t is ab so lute ly n e ce ssary th e n in

.
, ,

o rd e r to k e e p th e g e rm c o nt e nt l o w to pre ve nt th e bact e ri a ,
3 10 DI S T RI BUTI ON OF B ACT ERI A
fr o m multiplying rapidly in th e milk Th e e ff e ct Of te mp e ra
.

ture up o n th e rate o f gro wth in bact e ria is in di cate d by th e

fact that if milk is k e pt at 5 F fo r twe nty fo ur h o ur s a
0
°
.
-

singl e g e rm may incr e ase fiv e fo ld I f th e se sam e bacte ria

.

are k e pt in milk at a t e mp e ratur e o f th e incr e as e is

7 2 0 f o.ld I n th e pr o ducti o n O f th e milk th e n,

it i,
s d e s irabl e

that th e milk sh o uld b e co o l e d imme di at e ly and th e co o ling

o ught to b e suffi ci e nt to tak e th e milk b e l o w th e t e mp e ra

ture at which bact e ria r e adily gro w Whe r e it is p o ssibl e

.
,

a co o ling apparatus sh o uld b e in s tall e d and th e milk

co o l e d to n e a rly th e f re e zi ng p o int as quickly as p o ssibl e at ,

l e as t withi n a fra ctio n o f an h o u r fr o m th e time o f milki ng .

And th e milk sho uld b e k e pt c o o l until it is r e ady to b e u se d .

I t S h o uld b e ic e d during d e live ry and b e k e pt c o o l a fte r i t is

d e live re d to th e h o me This is e sp e cially true o f mi lk to b e

.

use d fo r infant fe e ding .

 C HAP TER XXI X
B ACT ER I A OF M I LK AN D I T S PR ODUCTS (C o nti nue d)
Th e r Oi th e bact e ria whi ch
S p e ci e s o f B a cte i a in M il k — .

ge t int o milk th e r e are a g o o d many di ff e r e nt sp e ci e s The y .

may b e gro up e d in th e pr e s e nt co nside ratio n into tho se bac

t e ria whi ch pr o duce littl e o r no change in th e milk e ve n tho ugh ,

the y may multiply to a co n sid e rabl e e xte nt a nd thi s wo uld ,

i nclud e th e bacte ria usually f o und in th e udde r mo st Of th e ,

bacte ria fro m th e air e tc Ano th e r gro up o f bacte ria p o

,
. r

duce c e rtain fe rme ntati o ns in th e mi lk cau sing th e p re cip i ,

ta ti o n o f th e ca se in d ue to th e f o rmati o n o f acid s and e nzym e s ,

th e pr o du cti o n o f gas whi ch int e rf e r e s with th e u se o f mil k

fo r th manufactur e o f ch e e se o r th e pr o du cti o n o f di sag r e e

e ,

abl e O d rs and ta ste s and unnatural co l o rs and e v e n slimin e ss

o ,
.

An o th e r gr o up o f bact e ria that s o m e tim e s ge t into milk are

di se as e pro ducing
-
.

La cti c Aci d B a cte ri a — L actic acid bact e ria always find

.

th e ir way into mi lk Th e y ge t into th e milk usually fro m so me

.

co ntaminating influ e nce and th e numb e r that ge t in may b e

,

e ry sm all but th e lactic a cid bact e ria find milk th e cultur e

'
v ,

m e di um par e xce ll e nce a nd altho ugh th e y are fe w in numb e r

,

at first th e y so o n o utgro w th e o the r fo r ms and mil k l e ft to it ,

se lf almo st invariably unde rgo e s lactic acid f e rm e ntatio n .

3 11
3 12 DI S T RI B UTI O N OF B ACTERI A
Thi s fe rme ntatio n is und e sir abl e in that it pro duce s a taste
which is Obj e ctio nable to m o st p e o pl e a nd pro duce s a phy sical
chang e in th e milk whi ch is o rdi narily o b j e ctio nabl e M e tho ds .

o f pr e s e rvati o n a re aim e d alm o st e ntir e ly again s t this fe rm e n

tatio n .Thi s do e s no t m e an o f co u r se that th e fe rm e ntatio n

o f milk by th e lactic acid bact e ria pr o duc e s a ha r mful cha ng e

in milk M e tchniko ff h as call e d att e ntio n to th e fact that th e

.

B ulgarians mak e use o f so ur milk a s a b e ve r ag e and h e sug ,

g e sts that lactic acid o rganisms may s e rve a ve ry us e ful pur

p o s e in th e i nt e stin e s o f man since th e y antago nize th e putr e
factive bacte ria whi ch have a v e ry und e sirabl e i nflue nce o n
h e alth by fo rming m e tab o lic pro duct s that a e de t rim e ntal r .

I n th e pr o ducti o n o f butt e r at l e a s t in thi s cou ntry we are

, ,

d e p e nd e nt up o n th e f e rm e ntati o n s caus e d by th e se bact e ria .

One o f th e gr e at pr o bl e ms th e n o f th e dairyman is th e p ro
, ,

ducti o n O f a milk in which th e lactic acid f e rm e ntati o n is h e ld

in ch e ck fo r a r e a so nabl e l e ngth O f tim e ; and it is p o ssibl e ,

with th e gre at e st car e to pro duce a milk whi ch will r e main

,

sw e e t fo r many days .

S li m y M il k . S limy milk may b e pro duc e d a s th e r e sult O f

bacte rial gr o wth S o me bacte ria gr o w in milk and s e cre te a

.
,

m ucilagin o us substa nc e which caus e s th e milk to as sum e a

slimy characte r S o m e time s thi s slim e caus e s l o ng s t rings o r
.

r o p e s to app e ar in th e milk This is a n unde sirabl e fe rm enta

.

tio n altho ugh no t dang e r o u s to he alth Ve ry fre que ntly it

,
.

may b e carri e d fro m o ne farm to an o th e r thro ugh th e cr e ame ry

o r ch e e se fact o ry pr o duct s ; that is thr o ugh th e skim mi lk o r
,
 B ACT ERI A OF MI LK AN D I T S PR ODUCT S 3 3
1

whe y I f o ne patro n bri ngs inf e ct e d mi lk to a cre am e ry and

.

the n ano th e r tak e s away th e milk o r whe y whi ch co ntains

th e e o rganisms in th e same can s in which th e y b ring the ir milk
s

th e n e xt m o rning inf e cti o n i s lik e ly to b e di s tribut e d and may

,

b e co me wide spre ad .

B l ue M il k E pid e mics o f blue milk s o m e tim e s o ccur

.
-
.

This is d ue to th e gro wth in th e m ilk o f a bact e rium that p r o

duce s a blue co l o ri ng substance An e pide m ic o f thi s kind .

wa s d e scrib e d and sh o wn to b e co mmunicat e d fr o m o ne l o t o f

'

milk to ano th e r as lo ng ago as 83 8:by S te inho ff 1 .

R e d M il k M ilk som e tim e s app e ars re d o r bl o o dy d ue to

.
,

th e gr o wth o f c e rtai n bact e r ia i n it B p ro d igio sus s o me . .

tim e s ge ts into th e m ilk fro m th e air and pro duce s the re its
characte ristic re d co l o r A y e ast is wide ly distribute d in th e
.

ai r which pro duce s

a r e ddish pink co l o r .

5
Th e se change s o f
‘Q
co urse are to b e
sharply di ff e re nti
ate d fro m th e pre s '
F 6 — B a t ria pr d
IG 1 ing m ilk fa lt ( ) B
c e o uc u s 1
e nc e Of bl o o d n th e
. . . .

( bl m il k) ( ) B l
l b f i acto ru e a c e ns
g cy a no e nus ue . 2 .

d m il k ) la ti vi i r py m il k)
milk I n th e latte r ( ( 3) C
re .
( o ccus c s s co s o .

Af t r L ipm a e n
.

case th e re d co l o r
app e a r s imme di ate ly up o n mi lking whi l e in th e fo rm e r it ,

o nly r e sults fr o m th e gr o wth o f bact e ria and h e nc e w o uld ,

appe ar o nly afte r a p e rio d o f incubatio n o f a g re ate r o r le ss

l e ngth ( F ig .
3 4
1 DI S TRI B UTI O N OF B ACT ERI A
Th e Di s e as e pro d uti ng B a cte ri a i n M il k
-
. A third gr o up o f
bact e ria whi ch unfo r tunat e ly are s o m e time s fo und in milk are
tho s e capabl e o f pro ducing human di se as e s S o m e Of th e s e .

bacte ria are d e rive d di r e ctly fr o m th e co w a s B act tub e rcu ,

.

l o sis fo r e xampl e
,
I n o th e r cas e s th e inf e cti o u s ag e nt e nt e r s
.

afte r th e milk is drawn F ro m th e s tandp o int o f public

.

h e alth thi s gro up is far m o re imp o rtant than any o th e r gro up

O f bact e r ia f o und in milk I n th e ca se o f th e bact e ria which
.

a re d e riv e d di r e ctly fr o m th e animal it w o uld s e e m p o ssibl e

to pr o t e ct th public wh e n th e y a re sufli cie ntl y ar o us e d to d e

e

mand p ro t e ctio n I n tho se ca se s wh e r e th e di se as e pr o ducing

.
-

bacte ria e nt e r aft e r th e milk is drawn th e pro bl e m is m o re,

d ifli cul t b e caus e it s e e ms qui t e imp o ssibl e to d e t e ct mild

,

cas e s o f such di se as e s as typh o id f e ve r diphthe ria and sca r

, ,

l e t f e v e r in th o s e that handl e mil k I t s e e ms lik e ly that

.

o cca si o nal e pid e mics may o ccur fo r a g o o d ma ny y e ars al ,

tho ugh rigid insp e cti o n co ul d quickly bring th e s e e pide mics

to a minimum .

Di s e a s e B acte ri a d e ri v e d d ir e ctl y fr o m th e Anim al .

Tub e rcul o sis is ve ry co mmo n am o ng dairy cattl e I t is al so .

th e m o s t c o mm o n dis e as e O f man and that it may b e trans

,

m itte d fr o m cattl e to man se e ms b e y o nd do ubt Wh e n .

milk fro m tub e rcul o u s co ws is in o culate d into gui n e a pigs th e ,

di se ase is pro duce d as a numb e r o f inve s tigato rs have f o und

,

lo ng ago R usse ll fo r e xampl e fo und that cc O f milk

.
, ,
1 .

kill e d a rabbit ; B act tub e r cul o sis is p r e se nt in mil k no t o nly

.

wh e n th e r e is an appar e nt dise ase o f th e udde r but al so wh e n ,

 B AC TERI A OF MI LK AN D I TS PR ODU CT S 3 5
1

ca ttl e app e ar in p e rfe ct he alth Wh e n cattl e are suff e ri ng fro m

.

tub e r cul o si s th e bacilli app e ar in th e f a ce s so that no t o nly ,

a e di se a se d cattl e lik e ly to hav e tub e r cl e bacil li in th e i r milk

r
,

but o th e r cattl e in th e same h e r d f ro m pa r ticl e s o f manure

,

which are ve ry like ly to ge t in du ri ng milki ng pro ce ss .

A numb e r o f bact e ri l o gi st s h e ade d by K o ch have d e ni e d

o , ,

th e inte rtra nsrni ssib ili ty o f thi s di se a se T h e w o r k s o f r e c e nt

.

y e ars e sp e cially that o f R ave ne l and m o r e r e ce ntly by th e

, ,

English and G e rma n C o mmi s si o n s a nd by P a r k hav e sh o wn

, ,

that a c o nside rabl e p o rti o n o f tubbrcul si in childre n is d ue o s

to b o vin e i nf e cti o n And th e qu e s ti o n at th e pr e s e nt tim e is

.

no t wh e th e r th e di se a se i s i nt e r t r a nsmi s sibl e o r no t but ,

what pro p o rtio n o f human tub e rcul o sis co me s f ro m b o vi ne

so u r c e s . I t i s c e rtainly incumb e nt o n th o se r e sp o n sibl e fo r

th e r e ari ng o f chi ld r e n to u se milk fr o m cattl e f r e e fr o m

tub e rcul o si s wh e n thi s is p o s sible I f no t it wo uld s e e m b e tte r

.
,

to use milk f r o m a h e r d r ath e r tha n f r o m a si ngl e u nt e st e d

co w fo B o lli ng e r and G e bh a r dt sh o w e d milk whi ch p o

“
,
r r

d uce d dis e as e in gui ne a pig s was i nn o cu o u s wh e n dilut e d with

h e althy milk fifty to o ne hund re d tim e s its vo lum e Th e re .

f o re the r e is l e s dange r in mi xe d h rd mi lk than that o f a singl e

,
s e

co w u nl e ss it i s p o sitiv e ly k no w n that sh e i s u na ff e ct e d with

,

th e di se a se
”
.

Th e d e t e cti o n o f tub e rcul o si s is so e a sily and a ccurat ly d o ne e

by th e use f tub e r culin that the re is no xcuse fo havi ng

o e r

milk co ntami nate d with th e tub r cl e bacilli Al l that is e .

n e e de d is an a ro us e d public se ntime nt .
316 DI S T RI B UTI ON OF B ACT ERIA
t
P a h o ge ni c B acte i ar wh i ch e nte r M il k afte r i t i s Drawn .

Typ ho i d F e ve r. F r e qu e nt e pid e mics o f typh o id f e v e r have

b e e n trace d to milk supplie s F ro m 85 7 89 9 a hundre d

.
— 1 1

and nin e ty fi v e such e pide mics have b e e n r e co rde d Th e

-
.

g e r m g e ts into th e milk in o ne o f s e ve ral ways F o r instance .

,

th e milk e r may b e su ff e ring fr o m typ h o id f e v e r e ith e r i n a mild

f o rm o r h e may b e a g e rm ca r ri e r I n e ith e r cas e h e co n

’
.
,

tam ina te s th e milk dir e ctly I n th e s e c o nd place th e c o ntami

.

natio n may o ccur in di r e ctly by th e p e rso n milking O r handl ing

th e milk al so acting as nurs e o r th e trans mi s si o n may b e m o r e
,

indi re ct as fo r instance by th e use o f p o llut e d wat e r in th e

, , ,

cl e aning Of milk ve ss e ls As a n e xampl e o f th e fir s t m e tho d o f

.

infe cti o n indicat e d ab o ve th e e pid e mic at S o m e rvill e M ass in

, ,
.
,

189 2 ,
may b e cit e d Thirty fi v e cas e s o f typh o id fe ve r o ccurr e d
.
-

in this place ; thi rty o f th e se up o n inve stigati o n w e r e f o und to

b e th e c o nsum e rs o f milk fr o m a pa r ticula r d airy .At this
dairy th e so n o f th e milkman han dl e d and de live re d milk
whil e suff e ri ng fro m an indisp o sitio n whi ch was no t diagn o se d
as typho id fe ve r until aft e r th e inve s tigati o n .

At th e Unive rsity o f Virginia in 1893 th e re o ccurr e d fo urt e e n

cas e s o f typh o id f e ve r Al l Of th e patie nts live d at a particul ar
.

ho te l and u se d th e same milk Up o n inv e s tigatio n it was

.

f o und that th e milk cam e f ro m a dairy situate d o n a cr e e k .

This cre e k r e ce ive d s e wage fro m th e mai n unive rsity s e w e r .

I t was al so fo u nd that a n e g r o ha ing typh o id f e ve r had thro wn

v

th e d e j e cta o n th e bank o f th e cr e e k Th e milkman us e d th e

.

wat e r o f this cr e e k to wash th e udd e rs o f th e co ws during

 B ACTERI A OF MI LK AN D I TS PR ODU CT S 3 7
1

milk ing I n 89 5 an e pide mic o f typho id fe ve r o ccurr e d in

. 1

S tamf o r d C o nn ,
Alt o g e th e r th e r e w e r e th r e e hundr e d and
.

s e ve n case s This e pide mic was th o ro ughly inve stigate d I t

.
.

wa s f o u nd that th e g r e at e r numb e r Of pati e nts w e r e in th e

habit o f d rinki ng milk that p r actically all o f th e m to o k milk
,

f ro m a S ingl e dai ryman and that whil e th e pr e mi se s w e re in, ,

go o d co nditio n and no typho id fe ve r co uld b e de te cte d o n the

place th e cans aft e r th e y w e r e wa sh e d in th e h o us e
, , ,

w e r e tak e n to a w e ll to b e ri ns e d Thi s w e ll was situat e d .

in a pl o w e d fi e ld and was c o ve r e d with l o o se b o ards Th e

, .

fi e ld was manure d with night s o il a nd it is b e lie ve d that ,

th e B typh o su s pr e se nt in th e so il was carri e d o n th e f e e t o f

.

th e m e n and go t i nt o th e wat e r thr o ugh th e cracks in th e b o ard

,

co ve r.

Cho l e ra Asi a ti ca C h o l e r a can und o ubt e dl y b e carri e d

.

thr o ugh a mi lk supply alth o ugh th e fact that th e M sp co mma

,
.

is v e ry susc e ptibl e to acid s u nd o ubt e dly mak e s it quit e im p o s

sibl e fo r thi s ge rm to g ro w in milk to any e xte nt o r e ve n to live ,

fo a ny l e ngth O f tim e
r P rac tically o nly o ne e pid e mi c h as b e e n
.

t r a ce d to milk a nd thi s is th e w e ll kn o wn Simp so n ca se

,
-
.

C h o l e r a sudd e nly app e a r e d o n shipb o ard i n th ha rb o r o f C al e

cu tta . I t wa s f o und that te n m e n btai ne d milk fro m a native O .

Of th e se fi v e w e r e sick with ch o l e ra but r e c o v e r e d f o ur w e r e

, ,

sick a nd di e d whil e o ne wh o d r ank o nly a littl e milk e scap e d

,
.

Di p hthe i a r A c o n id e rabl e numb e r o f e pid e m ics has

. s

— 8 8 thi rty six

b e e n t r a ce d to milk suppli e s F ro m 87 7 9 . 1 1 -

e pid e mics o f di phth e ria caus e d by infe ct e d milk are d e

, ,
3 18 DI S TRI BUT I O N OF B ACTERI A
scrib e d Kl e in and s o me o th e rs b e li e ve that co ws may suff e r
.
.

fro m this dise as e and that th e g e rms may b e thus t r ans

,

m i tte d fr o m th e a nimal to m an M o st inve s tigato rs have .

b e e n unable to o btain e vide nce which wo uld co nfirm thi s b e li e f ,

and it is g e ne r ally r e co gni ze d that th e B act di phth e ria mu st .

ge t int o milk aft e r th e milk l e ave s th e b o dy o f th e animal in

o ne o f th e fi r st two ways i ndicat e d und e r typh o id f e v e r An .

e pid e mic whi ch is charact e ristic o f t hi s dis e a se o ccurr e d in

Hight s to w n N J in 89 2
,
. Th e re w e r e tw e nty e ight case s
.
,
1 .
-

a nd f o ur d e aths Al l th e s e cas e s o ccurr e d withi n o ne w e e k

.
,

a nd all o f th m u se d milk fr o m o ne particular dairy

e Up o n .

inve stigatio n it was fo und that a G e rman b o y wh o as si ste d in

milking was suff e ri ng fro m diphth e ria at th e time and was ,

th e re f o r e th e m e ans o f di stributi ng th e dis e as e

,
.

S ca l e t F e ve
r A co n sid e rabl e numb e r o f e pid e mics Of
r .

scarl e t f e ve r have b e e n t r ac e d to milk suppli e s F r o m 867 . 1

1 899 ni ne ty ni ne e pide mic s o f thi s kind w e r e r e p o rte d

-
As a .

typical e xampl e o f such e pide mi cs th e o ne wh ich o ccurr e d in

B u ffal o N Y may b e cit e d
,
. .
,
Twe nty ca se s o ccur re d in th e
.

city and w e r e f o und to b e all in th e famili e s o f tho se taking

milk fro m a pa r ticula r fa r m Up o n inve stigatio n it was fo und .

that f o ur p e rso ns wh o live d at thi s farm had had s ca rl e t f e ve r ,

a nd that o ne a c o nval e sc e nt had h e lp e d in th e milki ng a nd

, ,

th e ha ndling o f th e c o ws This was sto pp e d and no m o r e

.
,

cas e s de v e l o p e d .
 CHAP T ER XXX
TH E B ACTER I A OF THE HU MAN B ODY

In tro d ucti o n Th e h e althy human b o dy at all t ime s is

.
—

inf e ste d o r inhabit e d by a gr e at vari e ty o f bact e ria I n th e .

maj o rity o f cas e s the se bacte ria a e harml e ss alth o ugh patho
r ,

ge nic bact e ria may o cca si o nally b e pr e s e nt Whe n pr e se nt

,
.
,

th e se path o ge ni c bacte ria d o no t alway s n e ce ssa rily e nte r th e

tissu e s o f th e b o dy and pro duce an infe ctio n B acte ria are .

pr e s e nt o n th e skin and muco us me mbran e s and in all th e cavi

tie s O i th e b o dy which are in co mmunicatio n with th e e xte rio r .

S o m e S p e ci e s are quit e co ns tant in th e ir pr e s e nc e whil e o th e rs,

a re f o u nd o nly o cca si o nally I t sh o uld b e r e m e mb e r e d that

.

th e co nditi o ns r e quisit e fo r bact e rial gr o wth a re o r ga nic f o o d ,

a ce rtai n am o unt Of m o i s tu r e and a te mp e ratu r e varyi ng fro m

,

1 5
°
C. to 4 0
°
C Th. e s e c o n di ti o n s a re all pr e s e nt in th e

human b o dy I t is p o ssibl e that th e no n patho ge nic bacte ria

.
-

may se rve in so me unkn o wn way to pro te ct th e b o dy against

invasio n by th e path o g e nic o rganisms Thi s p o int has no t .

b e e n tho ro ughly inve stigate d .

B acte ri a o f th e S ki n a nd E x po s e d M uco u s M e mb a ne s r .

A g r e at vari e ty o f sp e ci e s o f bact e r ia and la rg e numb e r s o f

th e m a e f o und o n th e e xp s e d su rfac e s Of th e b o dy The
r o .

se cr e tio n s o f th e sub ace o us glands o f th e skin and th e d e squa

3 19
3 2 0 R BUT I ON
DI S T I OF B ACTERI A
mate d c e lls o f th e e pid e rmis furnish th e n e ce ssary nutritive
substance s .

Th e numb e r o f bact e ria pr e s e nt o n th e skin is i nfl u e nc e d by

th e h e at and m o istur e Of th e b o dy th e cl o thing w o rn th e

, ,

o ccupati o n o f th e individual and th e d e gr e e o f cl e anli ne s s

,

O b se rv e d .

Disinf e cti o n and st e rilizati o n Of th e hands h as r e c e iv e d

ca re ful atte nti o n in r e ce nt y e ars by su rg e o ns and b a cte rio l o
gist s I t has b e e n fo und that th e hands n o twithstanding
.
,

th e fact that th e y a re e xp o se d to all s o r t s o f o ut sid e i nflu e nc e s

and that th e y are fr e que ntly cl e an e d and fr e e d o f d e squa

s

mate d e pithe li al ce lls are co ve r e d with as large a numb e r Of

,

bacte ria as th o se surfac e s o f th e b o dy whi ch a e no t e xp o s e dr

and no t cl e anse d so Ofte n F o ste r fo und that it was im p o s

.

sibl e to r e m o ve all th e bact e ria in a sh o rt tim e by th e use o f

s o ap wate r and brush I t is claime d at th e pr e se nt tim e
, ,
.

that co nstant washi ng o f th e hands with ste ril e di sinf e cting

s o ap wate r and brush fo r tw e nty minut e s is n e ce ssary to
, ,

r e mo ve all th e bacte ria B o ckhart and Ftirb ringe r have

.

fo u nd large numb e rs o f bacte ria und e r th e finge r nail s Th e .

fo re go ing state me nts s e rv e to illust r at e th e pre val e nce o f

th e bact e ria o n th e S kin and h o w diffi cult it is to r e m o ve

the m .

Th e bact e ria u sually f o und o n th e surfac e s O f th e b o dy

b e l o ng almo st e ntire ly to th e C o cca ce a and th e B acte ia ce a r .

Th e micr o c o cci are p e rhaps th e c o mm o n e s t inhabitants .

Th e pus c o cci a re quit e num e r o us at all tim e s .

 THE B ACTERI A OF THE H MA U N B ODY 3 2 1

HOh e in has i nve stigate d th e numb e r o f bacte ria in plate

culture s mad e fr o m unde rcl o thi ng o f vario us kinds and wo rn
fo r di ffe r e nt p e ri o ds o f tim e S t e ril iz e d mat e rials w e r e pla ce d
.

in di r e ct co ntact with th e b o dy su r face s On lin e n wo o l and .

, ,

co tt o n th e co l o ni e s whi ch de ve l o p e d vari e d in numb e r f r o m

2 8 to 600 in o ne day a nd fr o m 4 8 to 6 7 99 in two days
1 0 In .

fo ur days th e nu mb e r O f co l o nie s co uld no t b e co unte d .

M aggi o ra and B o rdo ni Uffre d ussi hav e i nv e stigat e d th e

-

S p e ci e s O f bact e ria f o und o n th e e pid e r mi s O f th e f e e t Th e .

fo rm e r has i o lat e d tw e nty two no n patho g e nic sp e ci e s fro m

s -
‘
-

th e f e e t . S o m e o f th e s e wh e n gro wn artificially pr o duc e d

disagr e e abl e o do rs .

M iqu e l r e p o rts that th e wa sh wat e r fro m th e laundri e s o n

th e S e in e R iv e r c o ntai ns m o r e bact e ria than th e wat e r o f th e

P ari s s e w e rs .

Th e c o njunctiva o f th e n o rmal e ye p o ss e s s e s a la r g e vari e ty

o f bact e ria I t is claim e d that th e se ba ct e ria c o m e prima rily

.

fro m th e air and th e S p e ci e s f o und in th e e ye have al so b e e n

,

r e p e at e dl y iso lat e d fr o m thi s so u r ce M icro c o cci path o ge nic

.
,

and no n patho ge nic co mp o s e th e maj o rity o f th e o rgani sms

-
,

pre se nt .

B acte ri a o f th e G e ni to u ri nary T a ct-

O th e r e xp o s e d
r .

m e mbran o us surface s o f th e b o dy also Sho w a larg e numb e r

o f bact e ria F o r e xampl e th e muc o u s s e cr e ti o ns fr o m th e
.
,

surface s o f th e vulva and vagina o f th e f e mal e and th e s e cre

tio n s fo und in th e m e atu s urinarius o f th e mal e and f e mal e
S h o w a gr e at many sp e ci e s O f ba ct e ria T h e ut e rus F all o
.
,

Y
3 2 2 DI S T RI BU TI ON OF B ACT ERI A
p an tub e s and bladde r in h e althy individuals co ntai n no
i ,

bact e ria .

Th e s e c r e ti o n s Of th e f e mal e g e nit o urinary tract at c e rtain

-

time s s e e m to b e slightly g e rm icidal F o r e xampl e in th e .

,

late r stag e s o f p r e gnancy and at parturiti o n in th e h e althy

p rs o n v e ry fe w bact e ria if any a re f o und o n th e surface s
e , ,

whi ch co m e in co ntact with th e se cr e tio ns Th e sp e ci e s .

f o und a re u sually no n patho ge nic Und e r ce rtain co nditi o ns

-
.

th e g e nit o urinary o r gan s o f th e mal e and f e mal e may b e

-

subj e ct e d to se ve r e i nfe ctio n s .

B acte i a f th e N o s e
r o Th e muc o us m e mbran e o f th e n o s e
.

usually co ntains a gr e at va ri e ty and a large numb e r o f bac

t e ria Th e y are d e rive d principally fro m th e in spire d air
. .

Th e hairs a nd muco u s s e cr e ti o ns o f th e n o s e and upp e r re s

p i ra t o ry pa s sag e s catch th e bact e ria and as a r e s ult practically

,

n o n e r e ach th e alve o li o f th e lung s imm e diat e ly Co nse .

qu e ntly th e e xpir e d air is usually st e ril e Th e nasal mucus .

e x e r ts a v e ry slight g e r m icidal acti o n o n s o m e bacte ria .

P ath o g e nic bact e ria such as B act di phth e ri a and M p ne u

,
. .

m o ni a a re fr e qu e ntly fo und o n so m e n o rmal nasal muco us

,

m e mbran e s Th e no s e and muco us s e cr e tio ns o f dis e as e d

.

indi viduals o fte n co ntain patho g e ni c bacte ria Thi s is s o m e .

time s th e case in i ndivi dual s suff e ring fr o m tub e rcul o sis and
o th e r inf e cti o us di s e a s e s .

B a cte ri a o f th e M uth o A larg e am o u nt o f w o rk has b e e n

.

do n e up o n th e bact e ria whi ch are u sual ly pr e se nt in th e

m o uth Th e mo uth is line d by a muco us m e mbran e co m
.
 THE BACTERI A OF THE H MAU N B ODY 3 2 3

p o s e d o f stratifi e d squamo us e pithe lial ce lls and is co nstantly

k e pt m o ist by th e saliva Th e saliva co ntains an amyl o lytic
.

f e rm e nt calle d ptyalin Thi s f e rm e nt co nve rts s tarch into

.

sugar Th e co nve rsio n o f th e s tarch mate rial s into sugar by

.

th e ptyalin i s und o ubt e dly favo rabl e to bact e r ial acti o n .

F urth e rm o re th e stratifi e d squam o us e pith e lium lining th e

,

m o uth is co ntinu ally b e ing thro wn o ff and thi s mate rial fur
,

nish e s e xce ll e nt nitro ge no us fo o d fo r th e bacte ria whi ch may

b e pr e s e nt.

Th e t e e th a re o f c o urs e b a thbd in s aliva all th e time

, , .

The re are always pr e se nt in th e m o uth n ume ro us irre gular

d e pr e ssio ns o f th e muco us m e mbran e and the s e to ge th e r with
,

th e space s b e tw e e n th e t e e th se rve as l o dging plac e s fo r

particle s o f fo o d which a re k e pt mo ist by th e sal iva Th e .

saliva is said by s o me w rite rs to b e slightly antis e ptic but this ,

actio n is o f no practical co nse que nce Th e t e mp e ratur e o f th e

.

m o uth is r e lative ly high ( 9 8 6 F o r 3 7 °

. and it is at o nce
°

e vid e nt t h at a b e tt e r plac e fo r th e incubati o n and gr o wth

o f bact e ria cann o t b e f o und Th e main pr e r e quisit e s fo r

.

bacte rial gro wth are at hand nam e ly mo i sture h e at and

, , , ,

o rganic f o o d .

Th e ave rag e m o uth c o nta ins a gr e at vari e ty o f sp e ci e s o f

bact e ria C e rtain o f th e sapro g e nic o r putr e factive bacte ria

. ,

th e chr o m o g e nic o r pigm e nte d bact e ria th e a éro ge ni c o r gas

,

pro ducing bacte ria and th e patho ge nic o r di se ase pr o ducing

,
-

bacte ria may b e pr e s e nt One w rite r de scribing th e bact e ria

.

O f th e m o uth says that alm o st e v e ry o rganism whi ch has b e e n

 2
3 4 DI S T RI BU TI O N OF B ACT ERI A
de scrib e d in any p o sitio n has b e e n fo und in th e human mo uth .

As a matt e r o f fact ab o ut fifty S p e ci e s o f bact e ria have b e e n

,

iso lat e d and cul tivate d fr o m th e m o uth Th e co nn e ctio n o f .

th e m o uth with th e e xt e ri o r acc o unts fo r th e larg e fl o ra p r e s

e nt . T h e bact e ria usually e nt e r by inh alati o n thr o ugh th e

m o uth by f o o d o r by drink Th e y may b e intro duce d in

,
.

dir e ctly by b e ing i nhal e d th ro ugh th e n o s e and also p o s t

nasal discharge s co ntaining bacte ria may find th e ir way into
th e m o uth .

Th e re are a large numb e r o f sp e ci e s o f bact e ria who se pr e s

e nc e in th e m o uth is accide ntal and th e re are also se ve ral ,

sp e ci e s whi ch are f o und rath e r co nstantly To th e latt e r .

class o f o rganisms th e nam e mo uth bacte ria has b e e n

give n by so m e inv e stigato rs As far as is kn o wn th e se bacte ria
.

pr o duce no de le te ri o us e ff e ct o n th e b o dy B acte ri o l o gists .

have b e e n unabl e to cultivate th e s e m o uth bact e ria and ,

”

c o ns e que ntly ve ry littl e is kno wn ab o ut th e ir bi o l o gy Th e .

o rigin o f th e s e bact e ria is unkn o wn .

A larg e am o unt o f r e s e arch w o rk o n th e bact e ria o f th e

m o uth wa s do ne by M ill e r in th e e arly p e rio ds o f bacte ri o
l o gical wo rk Th e fo ll o wing bact e ria w e r e iso lat e d and
.

name d by hi m : Le pto thrix inn o mina ta L e pto thrix buc 1

,

cali s maxima I o d o co ccus v agina tus Spirillum sp utige num

,
-

,
2
,

S p iro ch a ta d e ntium and B acillu s buccalis maxima

,
Le p .

to thrix buccalis maxima i s th e o rganism supp o se d to have

L pt thrix i a t rm whi h w appli d t filam t ba t ia

e o s e c as e o e n o us c er .

2
I d o i a
o co ccus f rm f ba t ri a
s co ccus ot lik th m i
O c e no un e e cro co cci.
 THE BAC TERI A OF THE H MAU N B ODY 3 2 5

b e e n s e e n by Anto n v o n L e e we nh o e k and de scrib e d by hi m

a s a n a nimalc ul e in a c o mmunicati o n to th e R o yal S o ci e ty .

o f L o ndo n i n 683 1 .

Th e ab o v e t e rms are no t us e d at th e pr e s e nt tim e Th e .

bacilli co cci and spirilli fo und in th e m o uth we re supp o se d

, ,

by M ille r to b e inte rme diate stage s in th e de ve l o pme nt o f

Le pt o thrix buccal is F isch e r say s th a t th e t e rm Le pto thrix
. .

sh o uld o nly b e appli e d co ll e ctive ly to th e m o uth bacte ria .

M ill e r and o th e r s hav e i so lat e d a larg e numb e r o f Sp e ci e s o f

bact e ria fr o m th e m o uth but th e id e ntity o f all the s e o r
,

g a n i s m s h a s no t b e e n c o mpl e t e ly e stablish e d .

Th e c o mm o n sap r o g e nic and zym o g e nic bact e ria a re n e arly

alway s pr e se nt in th e m o uth Th e fo llo wing patho ge ni c bac

.

t e ria are quit e O ft e n fo und in th e mo uth o f no r mal indi vidual s

M pn e um o ni a M py o g e ne s var aur e us and var albus
.
,
. . .
,

S t py o g e ne s M ; sputi se p ticus S a r te trage na

.
,
Th e path o
,
. .

ge nic bact e ria whi ch o ccasi o nally fi nd acce ss to th e mo uth

a e a s f o ll o w s : B act diphth e ri a
r B act mall e i B act tub e rcu
.
,
.
,
.

l o sis B t e tani P S p yo cya ne a and c e rtain o f th e path o g e ni c

,
.
,
.
,

fungi such as Actin o myce s var b o vis and var h o m inus . .

,

OI di um albicans and vari o us vari e ti e s o f sacchar o myc e t e s

’ '

.
,

M any o f th e bact e ria fo und in th e m o uth have th e p o w e r

o f pr o ducing o rgani c acid s One o f th e principal acids p ro
.

d uce d is lactic acid O ne o f th e bact e ria whi ch i s pro min e nt

.

in th e pro ductio n o f thi s acid is B act acidi l a ctici Thi s is .

-
.

th e bact e rium which caus e s th e s o uring o f milk and i n all ,

pro bability is intro duce d into th e mo uth in milk .

3 2 6 DI S T RI BUTI O N OF B ACT ERI A
De ntal C ari e s . Am o ng bacte ri o l o gical pro ce sse s
th e

which take place r e p e at e dl y in th e m o uth Of all i ndi vi dual s is

cari e s o r th e de cay o f th e t e e th Th e subj e ct o f carie s h as
.

b e e n s tudi e d by physicians and d e nti sts fo r ce nturi e s Th e r e .

a re ma ny th e o ri e s a s to th e cau s e o f thi s c o ndi ti o n Th e .

f o r e mo st is th e ch e mical and para sitic th e o ry whi ch may b e ,

b ri e fly stat e d as f o ll o w s : th e the o ry h o lds that de cay o f th e

t e e th r e sults primarily fro m th e actio n o f th e o rganic acids
pro duce d by bacte ria first o n th e e nam e l and se co nd o n th e
, , , ,

d e ntin e and p ul p and that as s o o n a s th e de ntin e is s o fte n e d

,

th e v ri o us putr e factive ba ct e ria infe ct th e t o o th and cau se

a

rapid disint e gratio n and de co mp o sitio n .

I t will b e r e call e d that th e cr o wn o f th e t o o th is c o ve r e d

with e name l which is th e ha r de s t tissu e in th e b o dy Unde r

, .

n e ath th e e name l is th e de ntin e which is th e principal co n

,

s ti tue nt o f th e t o o th This substance is trave rse d by smal l

.

micro sco pic tub ul a whi ch run almo st pa r all e l to th e lo ng

,

axis o f th e to o th The se mi nut e tub e s a e fill e d with p ulp and

. r

e mpty int o a c e ntral p e rp e ndi cula r pulp cavity Th e acids .

pro duce d by th e bacte ria di ss o lve th e e nam e l and change it int o

a ch e e sy ma ss Ab o ut 9 p e r ce nt calcium salts and 4 p e r
. 2 2

ce nt o f o rganic mate rial are lo st o n acco unt o f th e acids se

cre te d by th e bact e ria ( F i s ch e r) .

Wh e n th e so ft mass pro duce d by th e actio n o f th e bacte ria

up o n th e e name l is wash e d O ff as it is by th e saliva th e re
, ,

i s u s ually a slight e x cavati o n pr o duce d in th e und e rlying

de ntine which s e rve s a s a l o dging plac e fo r th e sap r o g e nic

,
 THE B ACT ERI A OF THE U
H MA N B ODY 3 2 7

bacte ria Th e de ntine is n e xt d e cal cifi e d and so fte n e d I t

.
.

is an int e re s ting o b se rvatio n that it take s l o nge r fo r th e b ac

t e ria to pr o duce a co mpl e t e change in th e so f t de ntin e than in
'

th e ha r d e nam e l . Onc e th e d e ntin e i s s o ft e ne d the bact e ria

f o ll o w th e tubul e s and e nt e r th e pulp cavity de stro ying ti ssue

,

a s th e y go . I t is p o ssibl e aft e r t hi s s tag e is r e ach e d fo r va r i

o us p a th o ge m c I m cro o rga ni sm s such a s th e p us c o c ci a cti no

, ,

m yce s e tc to e ff e ct an e ntranc e th r o ugh th e cari o us p o r

,
.
,

tio ns o f th e te e th and pr o duc e a ge n e raliz e d inf e cti o n in th e

b o dy .

F i sch e r r e f e rs to th e fact that bact e ria no t unlike th o se o c

cu r ri ng in th e mo uths Of p e o pl e in thi s day we r e fo und in th e

ho ll o w te e th o f so m e O f th e Egyptian mummi e s and thi s g o e s ,

to S h o w that bact e ria hav e b e e n inhabitants o f this r e gi o n o f

th e huma n b o dy fo r c e nturi e s .

B a cte ri a o f th e S to m ach a nd I n te s ti n e s . S inc e th e m o uth

co ntains such a l arge vari e ty o f bacte ria it is to b e e xp e cte d
that s o me o f th e se o rganisms will co nstantly pass do wn into
th e st o mach wit h fo o d and d r ink .

Th e bact e ria find th e h e althy sto mach an unfavo r abl e plac e

fo r gr o wth . T h e ga s t ric juic e in th e n o rmal s t o mach co n

tain s a quantity o f f re e hydro chl o ric acid (HC l ) Th e e . x

p e r i m e nt s o f S trauss and Wu r tz and o th e rs d e m o n s trat e th e

fact that th e n o rmal gast ric jui ce is de cide dly ge rm i c d al I t I .

o w e s thi s pro p e rty to th e f r e e hydr o chl o ric acid it c o ntains ,

a nd no t to any f e r m e nt . I t h as b e e n r e p e at e dly d e m o nstrat e d

by di ffe re nt o b se rve r s that th e n o rmal ga stric juice is fr e e f ro m

3 2 8 DI S T RI BUTI O N OF B AC TERI A
bact e ria and in fact is capabl e in c e rtain instanc e s o f kil ling
such mic ro organi sms as tho se o f Asiatic ch o l e ra and typh o id
f e ve r in fro m o ne ho ur to two h o urs Th e ve ge tativ e f o rms
.

o f B act anthracis hav e b e e n kill e d in fr o m fift e e n to tw e nty

.

minute s and B act tub e rcul o sis in fro m thirty si to f o rty e ight
.
-
x -

ho ur s Th e gastri c juice h as practically no e ff e ct o n th e S p o r e s

.

O f B act anthracis and th e vari o us sp e ci e s o f p us co cci a re al so

’

.
,

ve ry re sistant to its actio n A la rge numb e r o f th e no n

.

path o ge ni c bacte ria a re also re sistant to th e acti o n o f gastric

juice and pass thro ugh th e sto mach u nharme d into th e
inte stin e B act e ria may pass thro ugh th e sto mach int o th e
.

int e stin e s in particl e s Of fo o d and thu s e s cap e th e g e rmi cidal

actio n o f th e gastric juice I t is pro babl e that B act tub e r
. .

cul o si s and th e e nd o s po r e s o f all sp o r e pr o ducing bact e ria

-

r e adily pass thr o ugh th e sto mach to th e inte stin e .

I n c e rtain c o nditi o ns h o we ve r th e r e a re a larg e numb e r

, ,

o f bact e ria d e ve l o p e d in th e st o mach This is e spe cially

.

true as a r e sult o f dis e as e s in whi ch th e functi o nal activity

i
o f th e st o mach is l o w e r e d and th e gastric j u c e is r e duc e d in

amo unt o r b e c o me s f e e bly acid o r ne utral ( S te no sis o f .

pyl o rus carcino ma ) Us ually fe rme ntati o n is se t up in th e

,
.

s to mach by s o me o f th e bacte ria pr e s e nt This is th e co n .

diti o n in s o m e f o rms o f dysp e psia .

Ab e l o us inve stigat e d th e p o w e r O f a larg e numb e r o f th e

bacte ria f o und in th e h e althy sto mach to di g e st fibrin and ,

th e ability o f o th e r s to c o nv e rt starch int o d e xtr o s e Th e .

e xp e rim e nts w e r e mad e i n vi tro Hi s c o nclusi o ns w e r e th at

.
,
 THE B ACT ERIA OF THE U
H MA N B ODY 3 2 9

alth o ugh di ge sti on o f fibrin and c o nve rsi o n o f starch into

d e xtr o s e did tak e plac e as a r e sult o f bacte rial acti o n th e ,

time r e quir e d fo r such r e acti o ns was so l o ng that th e r e

was no basis fo r ass uming that any o f th e bacte ria we r e
c o nce rne d in th e pr o ce ss o f dig e sti o n in th e no rmal sto mach .

I n th e inte s tin e a l arg e vari e ty and an imme ns e numb e r

o f bacte ria are c o nstantly pr e s e nt S o m e o f th e s e bact e ria

.

a re pr e s e nt in th e inte stine c o nstantly whil e o th e rs o ccur

,

o nly o ccasi o nally T h e inte stinal c o nt e nts b e ing alkaline in

.

r e acti o n and th e t e m p e ratur e c o nstant and r e lative ly high ,

mo st o f th e bacte ria p r e s e nt find o ptimum c o nditi o ns A .

gr e at many o f th e bacte ria f o und in th e inte stine a re th e

sapr o ge nic o r putr e factive o rganisms T h e sp e ci e s o f bac
.

t e ria e nco unte r e d in th e inte stine are fo r th e mo st part

anae ro b e s o facultative a nae ro b e s Th e st ri ct e ro bic
‘

r . a
‘

bacte ria a e no t capable O f multiplying to any g re at e xt e nt

r

in th e inte stine o n acco unt o f th e scarcity o f o xyge n On .

th e in sid e O f th e int e stin e in c o nta ct with muc o us m e mbra ne

, ,

a e ro bic change s may tak e place to a limite d e xte nt B co li . .

is o ne o f th e many micr o o rganisms which is facul tative ly

ana e ro bic and a natural inhabitant o f th e inte stinal tract .

Th e e nd pr o duct s o f th e zymo g e nic and sapr o g e nic pr o c e ss e s

-

whi ch take place in th e int e stin e vary with th e charact e r o f

th e f o o d in th e int e stin e o n whi ch th e bact e ri a a re acting If .

p ro te in mate rial S imilar to m e at is acte d up o n th e pr o ce ss is

,

e ss e ntially sapr o g e nic and th e e nd pr o ducts pr o duc e d includ e

,
-

amino aci ds aro matics and gase s such as le ucin tyro sin
, , , , ,
33 0 DI S T RI B UTI O N OF B ACTERI A
indo l skat o l hydro ge n sulphi d e and ammo ni a I f ve ge tabl e
, , ,
.

mate rial co ntai ni ng a la rg amo unt o f carb o hydrat e is acte d

e

up o n by th e bact e ria u sually th e zymo ge ni c o r f e rm e ntive

,

pro ce s se s a e p re do mi nant and am o ng o th e r substance s

r ,

o rganic acid s a e p r o duc e d

r .

Th e int e s ti ne s o f ca r nivo r o us and o mnivo r o us animals co n

tain a gr e at e r numb e r o f bact e ria tha n th e h e rbivo ro u s animals ,

a nd th e r e are c o nsid e rably m o r e in th e larg e than in th e small

inte stin e ( De Gia a) x .

I t is claim e d by c e rtai n i nve s tigat o r s am o ng th e m G il l e p sie

, ,

that altho ugh th e maj o rity o f ba cte ria a re o f no use in gastric

o r pancr e atic dig e s ti o n ye t s o m e a re o f gr e at use in c o ntr o lling
,

putr e factio n in th e l o we r pa r ts o f th e small int e stine S o me .

o f th e o rga ni sms whi ch a e abl e to pa ss thr o ugh th e st o mach

r ,

a s s tat e d ab o v e pr o duc e o rganic acid s duri ng th e ir m e tab o lic

,

activity Th e se acids it is claim e d se rve to r e duce th e alka

.
, ,

l inity o f th e c o nt e nt s o f th e s mall i nt e s tin e and c o n se qu e ntly ,

in a m e asu r e co ntr o l th e putre factive pro c e sse s d ue to o the r

,

bacte ria S ap r o ge nic bacte ria are m o r e active in an alkaline

.

than in an acid m e dium I n th e large i nt e stine th e alkali ni ty

.

is so mark e d that th e acid s f o r m e d by bact e ria a re o f no co n

s e que nce in pr e v e nting sapro ge nic p r o ce s se s I n all pro ba .

b il ity t hi s acti o n o n th e part o f acid fo rming bact e ria is o f

-

littl e c o ns e qu e nc e Th e y are ce rtainly no t n e ce ssary auxil

.

i arie s in th e p r o c e ss o f dig e sti o n if th e y d o p o ssibly aid indi

re ctl y in c e r tai n i nsta nc e s Th e pr o ducts o f m e tab o lism o f

.

th e int e stinal bact e ria fo r th e m o st part are no t in any way

 THE B ACT ERI A OF THE H MAU N B ODY 33 1

use ful to th e human b o dy F isch e r says that th e fr e sh human

.

fa ce s co ntain 7 5 p e r ce nt wate r and p e r ce nt bacte ria

1 .

Th e inte s ti nal tract o f th e infant at bi r th is s t e ril e but in a ,

ve ry sh o rt p e rio d o f time afte r birth ( to 18 ho u r s) b e f o re any

12 ,

nutrim e nt h asb e e n take n bact e ria app e ar i n th e m e co nium

, ,

and as so o n a s mi lk is i ng e st e d th e y incr e as e rapidly in numb e r s .

S e ve n di ff e re nt spe ci e s o f bact e ria have b e e n i so lat e d fro m

th e i nt e s ti ne o f th e infant b e f o r e any nutrim e nt h as b e e n

tak e n Th e s e bact e ria pro bably co m e f ro m th e air B

. . .

c o li and B act aci di l a ctici are so me o f th e firs t o rganisms to

.
-

b e n o t e d in th e int e s tin e and th e fo rm e r usually r e main s an

,

inh abitant o f th e int e stinal tract thro ugh o ut lif e .

I t has b e e n S h o wn by car e ful e xp e rim e nts aft e r r e m o ving

,

a guin e a pig f oe tus f r o m th e p re gnant mo the r by C a se ian r

s e cti o n unde r strict a se ptic pr e cautio n s f e e ding th e yo ung

,

animal with st e rilize d f o o d and u sing e ve ry m e ans to p re ve nt

,

c o ntami natio n that afte r e ight days whe n th e a nimal was

, , ,

kill e d th e i nt e stine was ste ril e Th e e xp e rim e nt se rve s to

,
.

illust r ate that bacte ria have littl e o r no thing to d o with in

te stinal di g e sti o n at thi s p e r i o d at l e a s t a nd that th e y are
,

no t pr e se n t in th e i nt e s tin e s at birth .

Th e s ap r o phytic bact e r ia in th e n o rmal int e sti nal tract a re

abs o lut e ly harml e ss as l o ng as th e muco u s me mb r an e which

line s th e canal is uninjure d I f thi s b e i nju r e d so m e Of th e
.
,

bacte ria may e nt e r into th e ti ssue s a nd pro duce se ri o us dis

e as e d c o nditi o ns. Th e y may acco mplish thi s alo n e o r in co n
ne cti o n with s o m e o f th e path o g e nic sp e ci e s which o ccasi o n
33 2 DI S T RI B UT I ON OF B ACT ERI A
ally invad e th e int e stine F o r e xample B c o li is a fr e qu e nt
.
,
.

cause o f py o g e nic o r pus infe ctio ns in th e inte stin e and in co n ,

ne ctio n with B typh o su s fo r e xampl e may d o much to i n

.
, ,

cre as e th e s e ve rity o f th e i nfe ctio n .

B e sid e s th e ma ny no n path o g e nic bact e ria which a re pr e s e nt

-

in th e int e stin e no r mally at ce rtain tim e s patho g e nic bacte ria

,

may invade th e se pa r t s o f th e b o dy gro w and pro duce m o st

, ,

virul e nt infe cti o ns B typh o sus B dyse nta ria M sp co mma

. .
,
.
, .
,

a re am o ng thi s class o f micr o o rga nisms .

 I N DEX

An tag i m b tw n p i f ba t ria
on s e ee s e c eso c e ,

bi g i
A o e ne s s 8—13
64
1

A thrax ba ill
.

iz f 9
ti a id ba t i
.
,
n c us , s e O 2
Ace c c c e r um 1 7 2
Antiba t rial b ta
.
,

6 — 6
ti a id r tati
.
,
c e su s nce s , 2 1 2 2
Ace c c fe m e n —
o n 2 0 8 2 09
.

A ti —
id a t ba t ria ym
.

8
,
n e nz 65 67 e s, 1 2 2 2
Ac f s 98
Anti pti a ti n f h m i al b
.
,
-
c e

id r tati atty
.
,
se c c o o c e c su
Ac s fe m e n o n o f th e f

pr d d by ba t ria 7 4 7 5
, ,

—1
2 11
s ta 46 E
nce s, 1

Anti pti h m i al whi h

.

2 12 o uce c e 1

whi h pr d d b y a ti
,

f
se 46
cs , c e c s c a re 1

a ti n f n f rm ntati
,
c a re o uce c on o

ba t rial zy m f apr g i b
c e e n es o s o en c ac
54 ;
1 c o o , o e e o n,

t ria
e 2 2 2
2 0 5
Anti r m
.

Ae r b la f ba t ria kn wn 3
.

66
,
se u s, 2

A tit xi
.

o e s, c ss o c e o a s, 1 1
— 6 — 66
o bligat d fa ltativ 3 e an cu e 1 2
n
Ar m ati
5 8
o 65ns ,
f rm tati n f th
2 2 1, 2 2 .

Ae r tr pi m 4
.
,
o cs, e en o o e, 2 11
o o s 1 1

Ag ar a lt r m di m 7 4 7 5
.
,
2 12
—
Arth pr 5
.

as cu u e e u
Agg l ti i
.
,
ro s o e s, 0
—2
Art ba t rial f rm ntati n i th
6 63
.

u n ns, 2 2 .

s, c e e e o s n e,
Aggl ti id u 63 no s, 2
—
Ai ba t ria f th
.

2 1 4 5 2 1
89 — 9
A iati h l ra 9 7 9 9 ; arri d by
.

r, c e O e 2 2 2
—2
Al h l ba t rial pi g m t l bl i
.
,
s c c o e 2 c e

m ilk 3 7
,
co o c e e n s so u e n

95 ; i l bl i 9 5 9 6
,
1
—1
A i m ilati
.
,
1 nso u e n, 1

Al h l pr d d by pr 65 66
.

ss o n, 1 1
f f
iati ba t ria aff t d by 64
.
,
co o s o uce o ce ss o er
A
m tati 6 ; f rm tati f th sso c o n, c e ec e 1

A t lav
.
,
e n o n, 2 0 e e n on o e

hi g h r
u oc th 8 83 e e 2 -

2 1 1—2 12
A t ly i f ba t ria 8
.
, ,
e
—1 2
Al g r lati hip b tw f g i d
.
,
u o s so 8 c e 1 1

A t lyti ba t rial zym 8 8

.
,
a e o ns e een un an

; r lati hip f ba t ria t th

, ,
u o c c e e n e s, 1 1- 1 2 .

12 1 e o ns o c e o e

bl gr ue -
3 ; r lati hip f
e e n, 1 2 2
-
12 e o ns o

ba t ria t th tr
c e 3 o e ue 12
B
Alkali pr d d by ba t ria 7 5
.
,

es o uce c e 1

Am b pt r i m m b dy 6 B ab Ern t g anul 5 8
.
,

o ce o or une o 2 2 es -
s r e s,

Am i a id f fatty a id ri B a illi d fin d 7
. .
, ,

no c s o c se e s, 2 2 2 c e e 1

Am phitri h la f fl g l l t b B a ill u d nt ifi n dim i

. .
, ,

c o us c ss o a e a e ac c s e 9 r ca s, e ns o ns o f, 2

t ria 3 B a ill p digi hi t ry f 9 6

.

e 1 c us ro o s u s, s o o 1

Anab li m atab li m 66 fi
.
, ,

o s vs c o s 1 1 97
A ae r b blig at d fa lta B a t ria di v ry d hi t ry f th
. . .
,

n 3o e s, 1 o e an cu c e s co e an s o o e

tiv 3 t dy f 7 m rph l gy f 4 ff ;
,
.

e 1 2 s u o 2 — o o o o 1

A a e r bi lt r m th d 3 3 3 r tr t r
, .
, ,

—
4 4 ; tw
n o c cu u e e o s, 1 2 —1 f mo d fan s uc u e o 1 1 o

A aly i f wat r f ba t ria 9 9 3 rd r d tw b rd r f 5 ff ;

.
,

n s s o e or c e 2
— 01 o e s an o su o e s o 1

Anaphylaxi t ba t rial pr t i l w r ba t ria 5 ; hi g h r

. .
, ,

—2 1
57 s o c e o e ns , 2 o e c e 1 e 2 1

A ili dy f tai i g 6 9 5 9 6 4 ; i v l ti f rm a m g
.
, ,

n n — n o u on on
—2
es or s n n 2 4 7 o s 2

A i m al ki g d m q ti f la i fy dim i av rag
, ,
.
,

n n on o f —
ns o ns
o ue s c ss e 7 9; o 2 2 e e

i g ba t ria i in v g tabl ki g w ight f 9 ; l m ti f 9—3 5 ;

, ,

n c e n, o r e e e n e o 2 o co o on o 2

f r la f fl g l l t ba t ria 3
, ,

d m 6 —1 1 ou c sse s o a e a e c e 1
o 5 11

A i m al li f po tan g n rati rapidity f m v m t f 3 5 ; p

, .
,

n e s n e o us e e on o o e en o ca

l rr ding 3 5 3 7 ; m a
,

th ry 8 3
,

—1 su o un
—
eo , . su e s , ss
3 34 I N DE X
gr uping o ( Z g l a P lli l Of C l oo oe e c e s, o B a t rial di a
c e f plant 74 8 se se s o s, 2 -
2 0
r pr d ti f by B a t rial h ath 3 7
, .

i ) 37 4
on es — 1 e o uc on o c e s e s,

fi i 4 4 6 ; by p r f rm ati B a t ri l gy d finitio n nd hi t o ry
, , .

ss o n, 2
— s o e o o n, c e o o e a s
— ; rat f m l tipli ati
, ,

6 f 43 u on o
—
4 5 0 e o c 1 7
4 6 ; f d f 44 4 5 67 ; hi t l gy f B a t ri p rp ri 87 88
.
,

oo o
— s o o o c e o u u n, 1 —1

ba t rial ll 5 6 m i t B a t ri m a thra i 8— 9
, , .

th e c e ce 2 — 0 ; nu e c e u n c s, 1 1

tr t r f th ba t rial ll 5 65 B a t r id fr m l g m ba t ria 85
, .

uc u e o — ~
s e c e ce 2 c e o s o e u e c e 1
q ti l i ll 5 9—6 ;
, ,

ue s f on o nuc e us n ce s, 0 86
1

h m i al tr t r ll 6 —65 gr p 3
.

c e c f s uc u e o ce s, 0
'
B ggi t
e o o a ce a ou 2

t dy f by ltivati B rk f ld y t m f filt r 7 8
.
,

s u o 66 if ; cu o n, e e e s s e o e s,

t rilizati d di i f ti B i ary divi i r pr d ti by 4

, .

s e o n anf 77 s n ec on o n s o n, e o uc on 2
m th d f i lati
, ,

84 ; e 86—9 o s o so o n, 2 4 3
mi r pi al xam i ati
c o sco f 94 c e n o n o B l d r m a l t r m di m 7
oo se u as cu u e e u 2
la ifi ati B l m ilk pid m i f 3
, , .

10 1 ; f c 4;ss c n o o 10 2 —1 1 ue e e cs o 13

tax m y f 6 ; r lati hip B ill

,

ba t ria 67 68—7
, , .

o no o 10 2 — 1 2 e o ns ou f d f o n as oo or c e 1
d fi iti
,

Br ad m aki g zym g i ba t rial

.
, ,

o f 5 6
11 ;
—1 2 f 6 — 7 e n o n o 11 11 ; e -
n o en c c e
p i t f r m bla t th r f rm
,

zy m f i 5
, ,

o n s o e se nce o o e o s en es o use n, 2 1
f li f ; phy i l gy f Br w ia m v m t f ba t ria 3 5
.

—
o 7 e 11 7 12 0 s o o o 12 o n n o e e n O c e

r lati b tw th ir vir
,

B tyri a id f rm tati
, , .

ff ; e o ns e een e en on u c 9
— ; c e e n o n, 2 0 2 10
m t en d 7 64 ; wat r
an 12
—1 e ne ce s i r
n so um ilk 3
—
4 2 1 2 1

ary t gr wth f 9— 3 ; y
, , .

s o o o 12 1 1 ox

i fl
,

en d an f — n ue nce
g 3 33 ; 1 1 1 o C
t m p rat
,

e e — ure l ifi o n, 1 0 c a ss
34 4 ; 1

cati a rdi g t ff t f h at
on cco n o e ec o e o n, C ap l rr
su e s ndi n g ba t ria
su —
35 37 ou c e

a ti h m i al C ara m lizati g ar m dia 83

.
,

135 36;
-
1 f c on o c e c s o n, f e o n o su e

5 4 ; r lati f li g ht t C arm i tai i g 9 5

.
,

— 1 e n o o f ne or s n n
14 0 55 o 1

ff t f l tri ity C atab li m

.
, ,

e n, 66 67 68 s —1
1 59 ; ec 5 o e ec c o 1 9 o 1 1

6 ; la k f ff t f R o t g ray C atala xam pl f i nfra ll lar

.
, ,

1 2 c o e ec o n en s se e e o -
ce u
ff t f m v m t nzy m
,

6 63 —1
o n, 1 2 e ec o o e e n o n, e 78 e 1

ff t f pr r C ll mi t tr t r f ba t rial
.
,

1 63 ; e ec 6 ; O e ssu e o n, 1 3 e s, nu e s uc u e o c e
ff t f a iati
,

64 ; m t b o n o n, 1 65 —
e ec o sso c e a f 5 2

itri fyi g 8 C ll l ll wall 63 64

.

li m f 65 i —
'

o s O 1 n n 1 3 e u o se n ce s,

pr t tr phi 83 89 ; i Ch zy m g i ba— t ria i 3

.
, ,

85 ; —1 n en c n, 2 1
1 o o o c, 1 e e se o c e

tr g fi i g l g m S 5 86 C h m i al hang pI Od U d by
.
,

o en n or e u e ~
I e c c e s, C6 eh
'
-
x I

lph r ba t ria 86— 88 78; r lti g fr m d

,

2 85 87 ; —2 su u c e 1 1 zy m 77 e s, 1
—1 e su n o e

ir ba t ria 88 8 ; hr m og ni m p iti all d p t f

,

—1 d on
on c e 1 9 c o e c, co o s an so c -
e u re a c

ph t i ti f th pr t i m l l
,

19
9 7
01 1; g 8 —
; _ y o o e n c, 19 2 o 1 z on O
—
7 e o e n o e cu e 2 2 1 2 2

apr g i d M tab li m
.
,

m ge i n c, —2 1 n c an S
o 5 ; 2 02 s o e ee e o s

path g i C h m i al m p iti f f d
.

S p p h il i
a ro 6 7 ; c, 2 1 -
2 2 o e n c, e c co
9 os on o oo 12

p ifi d p ifi C h m i al d f di i f ti 84
.
,

2 28— 7 ; 2 3 s ec c an no ns e c c e c s, u se or s n ec o n,

i f ti di a d t
n ec o us 5 if ; se se s ue o, 2 3 a ti n f ba t ria 4 — 5 4 ; d i
c o o o n c e 1 0 1 s

m d f a ti f path g i i f ti g q aliti f 4 6 ff
, ,

o e o 43 c on o o e n c, 2 n ec n u es o 1

di t ib ti C h m i al tr t r f ba t rial ll
.
,

f 8 — f u n o uc u e
2 45 ; s r33 ; o 2 2 2 o e c s o c e ce

il 8 87 ; p i f 8
, ,

th e so 2 2 —2 s eces o 2 3 6 —6 5 0

f th air f wat r C h mi al b ta pi
.
, ,

2 85 ; o 89— ; e 2 2 92 o e e c sub s nce s, o so no us su

wag 3 3 4 ; f mi lk d ta pr d d by ba t ria 4 8 fi
,

an d se e 2
— 0 o an s nce s o uce c e 2
9
i t pr d t C h m i al th ry f i m m ity 7 3
.

f h ma
, ,

s 5 3 8;
o uc s , 30 1 o u n e c eo o un 2
-

b dy 3 3 C h m taxi
.
,

— —
o 19 32 e 4 o 43 44 ff s, 1 1 1 1

M i g la
.
, ,

B i
a cte r ace a — 8 u Ch e m g tro pi sm , 14 1 143 , ff
7 10 10 144
-

B a t rial l l dim i f 7 — 9
.
, .

c e ce s, e ns o ns o , 2 2 . C hl am y d o b acte riace a , 2 2 .
 I N DE X 33 5

h rid li
C lo e of me as a d i ta t 4 7
i s nfe c n 1 Diphth e ria tra d t milk ppli ce o su e s, 3 17
h l ra
.
,

—
C o e , 9 7 2 9 9 , 3 17 I 8
2 3
h l ra r a ti ipl i
.

C o e re d e c o n, th e , 1 7 6 D o co cc 16

hr at ph r ba t ria i a inf ti ta gi
. .
,

C om o o o us c e 19 1 D se se s nd ec o us a co n o us,
hr g i ba t ria K h p t lat tab
.
, ,

C o mo e n c c e 1 9 0 — 19 7 23 39 ; 2 —2 oc
’
s os u es es
hr par ba t ria ti b tw mi O
.
,

C o mo o us c e 19 1 li hi g s n co nne c o n e een crO r

id r aid a ti a id ba t ria i f pla t

.
,

C e , of ce c c c e n g i m an s d 45 ; s
74 an 2 o n s, 2

aki g 8 ; di trib ti f by air— br

,

m n , 2 09 2 0 s u on o o ne
ilia ba t ria ba t ria 9 9 S M ilk d
.
,

C on c e 2 9 If c e 2 1— 2 2 ee an

iti ab da ba t ria air f Wat r

,
.
, .

C es un nce o f c e in o e

Di i f ta t d di i f ti g 4 6 — 5 4
, , .

92 1 s n ec n s an s n ec n 1 1

C lad thrix gr p 4 Di i f ti pr
. .
,

o ou 2 s n ec f 7 7 ff 84o n, o ce ss o
'

C lad thrix d ri f ra t t f by m a f l t r 5 4
.
, , .
,

o 89 O o e 1 es o e ns o cu u e s, 1

C la ifi ati f ba t ria Di trib ti f ba t ria

.
, ,

—
ss c 4 9 on o c e 10 2 11 11 s 8 33
u o n o c e 2 2 — 2

D bl tai
, , , .

12 0 ou e s ns , 1 00
i d fi iti f 5 ; di m i D ha m l ti f t dy f b
. .

f
’
C o cc e n o n o 1 e ns o ns o un s so u on or s u o ac

9 ; dir ti f divi i t i 7
, , ,

—2 i n o s o n, n re
2 7 ec o e r a, 1

pr d ti by fi i 4 3
.

o uc o n s s o n,
E
C h F rdi a d 5 3
.

o n, e n n 0

C ld lig ht val f i d tr yi g E t pla m lay r f ba t rial l l 5 3 5 4

.
, ,

o s ue o n es o n c o s e o c e ce
—

ba t ria 7 8 7 9 S T m p rat r Egg a lt r m di m 7

, , , .

— u e cu u e
c e ee e e s as e u 2

d Th rm al d ath p i t El t i ity ff t f ba t ria 5 9

.
, , .

an e e o n ec r c e ec o on c e 1

C l i f ba t ria 3 9—4
.
, , ,

o on es O 6 c e 1 1 2

C l r i hr m g ni ba t ria 9 ff Ed pr S Sp r
, .

o o s n c o d o e c c e 1 0 n os o es ee un e r o es

a ba ill E d t xi 49 2 5 4 5 5
, . .

C o m m th — c us, e 1 2 0 n o o ns, 2 —2
9
C m pl m t b ta E t pla m th 5 4
all d th
.
, ,

o e e n su s nce c e e n o s e

E vir m t ff t f ba t ria
, , , .

6 2 2 n on e n e ec o on c e

C idia
.
, , ,

n — 6
o 7 2 2 4 12 1

C tag i di a E zy m ba t rial d th ir m ha
. .
,

on
— o us se se s, 2 2 2 n e s, c e an ec n
3 39 e

C v r g la pr parati 9 4 9 5 i a ti a t lyti
.
,

i m 6— 8 ns , — n, 1 u o
o e ss 7 e ; o s O c o 1 0 c
-

Cr thrix p ly p ra 88 89 ba t rial 8 8 ; d f rm t
.

e no o s o 1 —1 c e 1
—
1 1 2 an e e n s,

C lt r m dia 66—7 6 6 ; pr t lyti

.
, ,

—2 0 —2 2 0
u u e e 7 ; im 2 02 o eo c, 2 1

C lt r m th d 85 9 f a i m al with r lt
.
,

u u e m i ei o s, — 2 1 2
—1 un z a t o n o n s e su
3 33
C lt r t ti g di i f ti by 5 4
, .
,

u u e s, i g i ti es ym 65
n s n ec o n 1 n n an e nz e s, 2

( bl gr Epid m i 9 5 9 9 ; a d by m ilk
al g )
. .
,

—2
Cy phy a no ce a ue — een a re e cs , 2 c use

hi p f ba t ria t ppli
,

l ti a o ns — 6— 8 o c e o, 12 2 12 su e s,
3 31 31
E m ar h r ll th 89
. .

s c o e

E ba t ria b rd r
.
, ,

D 5 4 ; f 5 u c e ,
1 ,
10 su o e s o ,
10

111

D fi iti f ba illi f ba t ria E ryth rm i rg a i m 3 8

.

e n 7; o n, o c 1 O c e u e c o n s s, 1

Exha ti th ry f i m m ity 7
.
,

7 11 us o n eo o un 2 2

la f b Ext r al diti b ari g f

.
,

D itri fyi g rg a i m
.

en n o n s s, c ss o ac e n co n o ns, e n o on

t ria all d 6— 7 f d i th ba t ria 7 64

,

—1
e c e 2 2 2 2 o un n e c e 12

s il
.
, ,

88 o 2

D tal ari
.
,

en 6 —
3 3 7 c e s, F 2 2

Dia m i a id b ta f rm d fr m
.

no c s , su s nce s o e o

by ba t rial a ti Fa ltativ para it d apr phyt

,

c e c o n, 2 2 2 2 2 3 cu e s es an s o e s,

Dig ti g zy m
, .

es n 8 en e s, _ 1 —2 2 0
—2 2
2 7 2 2 9
Dil ti m th d i i lati F rm tati zy m pr
. .

u 87 88
o n e o s o so o n,
— e e n o n, a n e n e o ce s s , 2 0 2

i f ba t rial ll 7 — 9 rg a iz d d rg a iz d
.

Dim e ns o ns o 4 c e ce s, 2 2 . 2 0 ; o n e an uno n e
3 3 6 I NDE X
ag t f 4— 6 ; d fi iti
en s of G idia 2 0 5 87 2 0 e n on o on 2 2 1, 1 2 1, 1

pr la ti a id Gram tai 9 7
, , , .

f 6 —2 0 ’
s s n,
o ce ss 5 o 2 0 c c c

a ti a id
, , .

6— 8 ;
2 0 2 0 8 9 ce c c 2 0 -
2 0

b tyri a id 9 — ; m tha
,

u c d c 2 0
H 2 10 e ne a n
m ilagi high r
,

uc ; f th no u s, 2 11 o e e

al h l ar m ati
co d fatty a id
o s, H abitat di trib ti
o f ba t ria
cs , an c s, s u o n o c e ac
art i r tti g f rdi g t 8
,

—
; i
2 1 1 2 12 th n e s, n e n o co n o 2 2

flax pr parati f indig ri g f H a ging dr p pr parati 9 4

.
,

e on o o cu n O n o e o ns ,

hid d t ba br ad m aki g H p l b t i 5 ; f rm typ

, , .

es an o cco , f e -
n a o ac e r a, 1 o es o

; r pr d ti a m g 4 5
, ,

t —2 1 —2 1 uc n on 2 — 0
e 4 c 5 2 1 5 1 e o o

F rm t rg a iz d d rg a iz d H p l b t in 5 9
. .
, , .

—
e e n s, o n e an u no n e a o ac e r a 10 10

H at val f i d tr yi g ba t ria
, , .

2 0
4 6 -
2 0 e ue o n es o n c e

Filtrati t rilizati by 7 8 ; f 7 9 3 4— 3 7 S T m p rat

.
, , ,

o n, s e on o 1 1 ee e e ure
wat r 3 3 wag 3 H m at xyli f tai i g 9 5
. .
, ,

e f 01
— 02 o se e 02 e n n n
o or s

H m ly i ba t rial 5 7 5 8
, , , .

0 e o s ns, c e 2 —2
3 4
Fi h ph ph r t ph t Hi g h r ba t ria ( Tri h b t ria)
. , .

s f d os o e s ce nce o ue o o o e c e c o ac e 2 1

g i ba t ria 9 8— r pr d ti
, , ,

en c c e 4; 1 m g t h 5
—
5 2 00 2 e o uc o n a o n e 0 1

Fi i n r pr d ti by 4 43 Hi t l gy f th ba t rial ll 5 —6
. .
, ,

ss o e o uc on 2 — s o o o e c e ce 2 0

Flag lla ba t ria 9 ff ; d t i H m th rm i rg a i m 3 8

. .
, , ,

e on c e 2 o no h o o e c o n s s, 1

with pr f fi i a b dy ba t ria
. .
,

t f 6 H m f t h — ss o n,
— u n o
e r e re 45 4 o ce s s o 3 9 33 c e o e 1 2

h m i al tr t r f 65 ; p ial
.
, ,

c e c s uc u e o s e c

tain d vi d f 9 8
,

s e se o r,
I
F lam t rilizi g by a dir t 7 9
.

e s e n ec

F l id lt r 9 I ll i rtai hi t ri al d t b
, ,
.

u cu u e s, 2 us o ns , ce n s o c ue o ac
F d f ba t ria 4 4 4 5 67 8 ; t ria 9 6— 9 7
.
,

oo o c e
— 11 e 1 1

la ifi ati a rdi g t 8— 9 ; I mm ity t path g i ba t ria 7

, , , , .

c ss c o n cco n o 12 12 un o o en c c e 2 1

am t r q ir d 8— 9 ; f p h th ri
, , ,

o un e u f —
73 ; e 7 73 12 12 O o 2 eo es o 2 2 2

t g i ba t ria I dig ba t rial zy m f

.
, ,

o en c 99 c e 1 in -
2 00 n o c e e n es o use
F rm ald hyd a di i f ta t 4 8 pr pari g 4
.
, ,

o e e as s n ec n 1 e n 2 1

I d l pr d ti f by ba t ria
.
, ,

51 0 75 n o o uc on o c e 1

Fra ti al m th d f i lati 86
.
, , ,

c o n 76 e o s o so o n, 1

Fr zi g all ba t ria t kill d by 7 9 I f ti fa t r i fl ig d

. .

ee n c e no e n e c o ns, c o s n ue nc n an

T m p rat nd Th rm al d ath m di fying 67 7 ; fl t f pr p

, , .

S ee e e ure a e e o 2
—
2 0 e e c O o

pi t ty f vir l mb r
,

o n 6 7 — 68 ; er o u e nce 2 2 nu e

F ngi r lati n hi p f ba t ria t f ba t ria ary t pr d

.
,

u e o s o n c e o 12 0 o c e ne ce ss o o uce a

i f ti av
, ,

12 2 68 69 ; f in n e c o n, 2
—2 e nu e O

ptib ility f i
.

f ti 69 ; ec o n, 2 su sce O h

G di id l t 7 ; m th d f inf v ua s o 2 0 e o O ec

ti in a f pla t 7 8 8
,

n s, 2 —2
on c se o 0

G a pr d d fr m m tab li m f I f ti di a 3 — 3 9
.

se s , o uce o e o s o n e c o us se se s, 2 2 2

ba t ria 7 4 ; whi h m y b p I t rm itt t filtrati f wag 3 3

.

c e 1 c a e ro n e e n on o se e 0

I nt rm itt t m th d f t rili zati

, .

d by pr f f rm tati
,
'

d uce o ce ss o e en o n, e en e o o s e o n,

6
2 0 8 —8 0 1

G latin lt r m di m 7 3 7 4 t ti ba t ria
. .

cu— I u e f t h —u n es ne s , o e 2 2
e e 3 9 33 c e

G lati plat m th d K h 6 89 9 I v l ti r 7 ; pr d t n
, . ,

e n — e fe m 4
— o oc
’
s, 1 n o u o n o s, 2 2 o uc i o

G lati tab lt r 9 —9 f by fi i ll b f r r a hi g
.
, ,

e ns cu u ef 1 2 o ss o n o ce s e o e e c n
G it rinary tra t ba t ria f th ad lt tag 43
.
, ,

en o u -
c c e o e u s e

I r n ba t ria 88 89
.
, ,

2 1
— 2 2 o c e 1
—1
3 3
G rm i idal a ti n f m h mi al I lati m th d f 86—9 ; f p h
. .
,

e c c o o so e c e c so o n, e o s o 2 o o

b ta i ba t ria f t dying
,

su 4 6 ff
s nce s, 1 t g . o en c c e or s u ,
2 01 .
 I NDE X 337

K th d d i t dyi g ba t ria 66 Me o s u se n s u n c e ,

10 1

Kir h r arly ba t ri l gi t
c e e M i r bi l gy c e o o s 2 c o o o 1

K h R b rt 6—7 ; d v l pm t f
.
,

Mi r i 6
, , .

oc o e e e o en o c o co cc 1

lt r m th d by 85 ; r l f t
, ,

Mi r iz f
.
,

cu u e e p g di o s u es o o c o co ccus ro re e ns, s e o
pr v ti b tw a mi r
, ,
,

o n e co n e c
9 o n e ee n c o 2

rg a i m d a di a 4 5
.

o n s Mi r p ba t ri l gi al 6 ; hi t ry
an se se , 2 c o sco e , c e o o c , s o
f th 9 3 —9 4 o e

Mi r pi al xam inati f ba t ria

, .

L c o sco c e on o c e ,
— 10 1
93
L a ti a id ba t ria 7 i m ilk 3 M i r pira 9
.

c c c c e 1 2 n 11 c os 1

; pr d d by ba t ria f th M ig la la ifi ati n f ba t ria by

, , , .

3 12 o uce c e o e u c ss c o o c e

m th 3 5
, ,
—10
ou 3 2 4 ff 10

L a ti a id f rm tati M il k f d f ba t ria 7 ba t ria

. .
,

c c 6— 8 c e en o n, 2 0 2 0 as oo or c e 1 c e
L w h k A th y V
.
,

f — 2 12 —2 1 — 1
eeu en oe 4 4 3 5 3 8 n on a n, 2 o 0

L g m ba t ria 85 86 85 87 M ld r lati hip f ba t ria t

.
, , , .

e u e c e 1
—
1 2
—2 o s, e o ns o c e o 1 2 1,

L if pr ll g ral hara
.
, , ,

e f o ce sse s o ce s, e ne c c L2
—1 2
3 4
tri h la f fl g l l t b
.

t f 65 66
er o 1M -
1 o no c o us c ss o a e a e ac
Light ff t f ba t ia 5 5 5 9 t ria 3
.
,

e ec o on c er 1 1 e 1
-

L im a di i f t a t 4 7 M rph l g y f ba t ria 4 ff
, , , ,
.

e as s n ec n 1 o o o o c e 1

L ati r lati n f ba t ria t M th ba t ria f th 3 3 5

,
. .
,

oc o ns , e o o c e o ou c e o e 2 2 — 2

M v m t f ba t ria ff t
, , .

28 2 o e en —
9 35 o c e 2 e ec

L m ti f ba t ria 9 —3 5 ; ff t h m i al ff t
.
, ,

o co o on o f 4 4 ; c e 2 e ec O c e c s o n, 1 1— 1 2 e e c

f h m i al f m v m t ba t ria 63
,

o c e c s o n, 1
—
1 1 2 o o e e n o n
4 4 c e 1

alkali m thyl bl f M ilagi f rm tati

. .
,

L ffl
’
oe ers ne ; e e ne ue or uc no us e e n o n, 2 11

tai i g 9 7
s nn m ilk 4 2 1

L ffl r bl d r m 7
.
, ,
.

M ii l l —
’
oe e s O F 4 5
oo se u 2 e r,

L ph tri h la f fl g ll t b M ltipli ati f ba t rial ll rat f

.
, . .
, .

o o c o us c ss o a e a e ac u c on o c e ce s, e o

t ria 3 R pr d ti
,

S —
e 1 43 46 ee e o uc on

L w r ba t ria (H pl b t i ) 5 r lati hi p f b
. . .
,

o e My b t i
c e a o ac e r a 1
— 2 1 xo ac e r a ce a e o ns o ac

r pr d ti am g 4 5 t ria t 4
, ,

e o uc on o n 2
— 0 e o 12

L m i ity p w r f i rtain b
.
, .
,

u no s o e o n ce ac

t ria 9 8
, ,

e ,
1 .

M N tralizati f b ill in pr para eu o n o ou on e

ti n ba t rial f d 69 o as c e oo

Ma g ping f ba t rial l l 3 7 N itrifi ati pr k w

.
,

ss ro u o 6 c e ce s, c o n, o ce ss no n as, 2 2

4 1 7 2 2

M m bran ba t ria f 3 9 N itrifyi g ba t ria 83 85

. .

m e s, uco us, n c e 1
—1
e c e o 1

Nitr g fi i g ba t ria 85 86 85
.
, ,

—1
o en -
x n c e ,
1 ,
2

M r ri
e cuhl rid n anti pti
c 87c o e as a se c 2

Nit g b ta pr d d by
.

47
1 ro e no u s su s nce s o uce
ba t ria 7 5 7 6
.

M i m p di
er s 6 o e a, 1 c e 1
—1

phili ba t ria 3 5 3 6 Nitr ba t ria 83 84

.
,

M —1 c e
—1
e so c c e 1 o so -
1

pr d N l b
.

M tab li m f ba t ria 65 7 lt r dia 7 5 7 6

, .
,

e o m i m — —1 0 o na u no us cu u e e
s o c e 1 o .

N ba t ria f th 3
, ,

t
uc s of 7 8 1 1— 1 2 o se c e o e 2 2

N xi r t ti th ry i ity
.

M ta hr m ati gra l
, .
, ,

e c 55 57 5 o
9 c f m m nu e s , — o o us e e n on eo o un ,

M tatr ph i ba t ria 67
.
,

e o 7 c c e 1 2 2

M tha d m ilag i f rm ta N l i ba t rial l l q ti n f

. .
,

e ne an uc no us e e n u c e us n c e ce ,
ue s o o ,

ti o n, 2 1 1 59 6 .
-
0 .

Z
33 8 I NDE X
0 P o iko l o th e rm i c ba t ria c e 13 7 —1 8
3
P lar gra l 5 5 5 7 5 9
, .

o nu e s , —

p ni va i P r lai fil t r 7 8
, .

O so 63 64 ; n s, nd 2 -
2 cc ne s a o ce n e s,

ag t i fl i g th 67 P tat lt m di m 7 7
.

en s n u e nc n e 2 o o e s a s cu ure e u 1- 2

O m ti pr r i ba t rial ll 5 5 Pr ipiti 64— 65

, .
, .

s o c e s su e n c e ce s, ec ns , 2 2

5 6 ; pla m ly i by Pr r ff t f ba t ria 63
.

9 s o s s 12 e ss u e e ec o on c e 1

Oxidativ ha ng Pr t i m l l r lt f d m p
, .
, , , .

74 e c e s, 1 o e n o e cu e e su s o e co o
O xy g varyi g ff t f ba t ria all d p tr fa ti f
.
,

e n, n e ec s o on c e i i
s d
t on an SO c -
e u e c on o

ary f
, , ,
— ne ce s s —2 2
1
3 33 1- 1
4 4 ; 1 1 1 2 or 2 2 1 7
pr d ti n f l ig ht by ba t ria P t i a aphyla i t ba t rial 5 7
, .

o uc o o c e 2 00 ro e ns , n x s o c e 2

Pr t lyti zym f ba t ria 7

.
, , .

o eo c en es o c e ,
2 1

P 2 2 0

Pr t
.

o e o se s, 2 2 1

P ara hr m at ph r ba t ria 9 Pr ti ta gr p f m i r rg a i m 6
.

c o o o o us c e 1 1 o s ou O c oo ns s, 1 1 :
P ara it ba t ia all d 8 ; fa lta Pr t pla m f ba t rial ll 5 3 —5 7
.
, ,

s e s, c er c e 12 cu o o s o c e ce s,

tiv 8 tri t blig at 9 3 Pr t tr phi ba t ria 67 83 89

.
,

e 2 2 s c or o e 2 2 —2 0 o o o c c e 1 1
—1
;
P aratr phi ba t ria 67 Pr t z a r lati hi p f ba t ria t
, , , , .

o c c e 1 o o o e o ns o c e o,

Pa t L i 5 —
.
, ,

s e ur, o u s, 12 6 —1 2
13 5
12

P a t —C ham b rlai p r lai filt r P d m a i di g f ra iz f 8— 9

.
,
.

s e ur e n o ce n e se u o on s n o e , s e o , 2 2 .

d l —
78 P se u O o zo8 g c
e a, 3 39
P ath g i f ph t g i ba t ria P y hr phili ba t ria 5 3 6
. .

o e ne s s o o o en c c e s c o c c e 13 —1

P t m ai pr d ti d g pi g f
.
,

2 01 o ns , o uc o n an ro u n O

P ath g i ba t ria 8 7 ; m d
.
,

o en c c e
—2 o e 8 1, 2 — 5 2 2 1
2 2 3 3 2 4 49
f a ti ff t P r lt r 85 86
.
, ,

O f — f
on o ec s o on u e cu u e s, —
4 c 45 2 3 2 e

b dy 4 6— 48 i m m ity t P rifi ati f wat r 3 —3

.
, ,

th e o 2 2 un o u c o ; f
n, o e 01 02 o

tra f i t m ilk 6 wag 3 3 3

, , ,

— en nce o n o —
7
2 73 ;
1 2 3 1 se e 0 2 0

i iz f 9
, , , .

3 7 1 Bi s co cc s e o 2

P ll i l 3 8—3 9 P tr fa ti d t apr g i d
.
.
, ,

e c e s, u e c o n, ue o s o en c an

P pt ba t ria 7 ; m h
.

o ne s , 2 2 1 hil i 6— ec
e p p
sa ro c c e 2 1 2 1

P ritri h la f fl g l l t ba t ria ; r lt f f
.
,

e c o us c ss o a e a e c e i m
an s f — o 2 2 0 2 2 1 e su s o o

pr t i m l l
, , ,

—
n o e cu e
3 1 o e 7 2 2 1 2 2

P hag yti th ry f imm ity 7

.
,

oc c eo o un , 2 2

73
2
R
P h ph r a d by ba t ria
.

os o e sce nce c use c e

98
1 Rapidity f m v m t f m t il b o o e en o o e ac

Ph t g i ba t ria 9 8— t ria 5
.

o o e n c c e 1 2 01 e 3
Phy i l g i al m th d f i lati 86 R at f m ltipli ati f ba t rial ll
. .
, ,

s o o c e o s o so o n, e o u c on o c e ce s,
—
87 4 46 3
Phy i l gy f ba t ria 7 ff R di Fra i 9
.

s o o O c e 12 e nc sco ,
P hyt t xi 5 5 R d m ilk 3 3
. .
, ,

o o ns, 2 e 1

Pig m t ba t rial — 9 6 R d ti b ta
. .
,

e n s, c e 19 1 1 e uc 73o n su s nce s, 1
Pla t r lati hip f ba t ria t 9 R g rat r 5
. .
,

n s, e o ns o c e o 11 e e ne o s, 1 0

ba t rial di a R lati hi p f ba t ria 5 if ; t

.
,

f — 8 ns c e 11 o
12 0
; 74 c e se se s o 2 2 0 e o o

P la m ly i t dy f tr t r f ll pla t E my t
, .
,

s s, s u o uc u e o s — n s, t 1 19 12 0 o u ce e s,
s o s ce ;
by 9 12 orf gi —
; t Cy
un ph 12 0 12 2 o a no y ce a
Pla m pty i ph m bl g alg
,
°
, ,

f th —1 2
s o p s s, e no e no n o a or ue 3 ; t re e n a 12 2 o e
-

pli d t ba t rial ll 5 7 tr alg

, ,

e o c e ce s, E phy u 3 ;
ce a t or ue a 12 o

i M ar A t i M yx m y t li m m ld 3
.
, ,

Pl e nc z , 4 cu s n o n u s, o ce e s, o r s e o s, 12

y th ry f 7
.

Pl e o go n eo o 2 10 3 4;12 t My b t i 4; o xo a c e r ace a 12

P l m rphi m 4— 7 3 t pr t z a
.
, , , ,

6 —1 2
eo o s ,
2 2 ,
10 . o 5 o o o ,
12 .
 I NDE X 339

R pr d ti f ba t ria 4 5 ; by
e o uc o n o p rt 4 7 c e 2 -
1 S o s,
bi ary divi i fi i 4 43
,

tai i g m th d appli d t t dy f
.

n s on or ss o n, 2 -
S nn e o s, e o s u o
am g th Tri h ba t ria 5 —5
on gra l 5 7 5 9 ; adva tag f in
e c o c e 0 1 nu e s , — n es o
R m bla f ba t ria t th r f rm
,

xa m i ati f ba t ria 9 4 fl
.
,

e se nce o c e o o e o s e n on o c e
f li f S tai i g pri ipl i v lv d i 9 9 —
, .

—1 2 0 nn
o 7 e 11 nc es n o e n, 10 1

R p i ati S tai vari ti f 9 5 9 8

.
, , .

6 n 68— ns , —
es r 5 7 o 1 1 1 0 e e s o

Rig r tat S taphyl i 6

.
, , , .

o 3 s e 0 1 o co cc I

Ro t g ray la k f ba t ri idal pr p S t a m t rili i g by 8 —84

, , .

n e n s, c o c e c o e s e z n 0

_ St th rm i ba t ria 3 8
, , .

ti
cr f 6 63 es o 1 2 -
1 e no e c c e 1

R Al xa d r 9 S t ril izati pr
.
,
- .
,

OSS , n e e f 7 7 84 e o n, o ce ss o -

S t m a h ba t ria f th
.
, , .

— o c c e
3 7 3 9 o e 2 2

tr pt i
, , .

S S 5 e o co cc 1

S tr pt th ix a ti m y
.
,

e o r c no ce s , 2 2

S and fl di a d t ph t g i S t i t blig at para it 2 9 3

.

ea se se ue o o o en c r c or o e s e s, 2 -
2 0

ba t ria S tr t r f th ba t rial ll 5 —65

, , .

c e 2 01 uc u e o e c e ce 2

S apr g ni ba t ria S b lt r
.
, , .

o e 6— 7 c c e
9
—9 2 1 2 2 u cu u e s, 1 2

S p p hil i ba t ria S ubi f ti pr all d 3 3

.
, .

a ro 6 7 c c e 2 1
-
2 2 n e c o n, o ce ss c e 2

S apr phyt ba t ria all d 8 8 S g ar fr b ill 7 7

.
, , .

o e s, c e c e 12 2 2 u -
ee ou o n, 0
-
1

3 ; fa ltativ 9 ; f d am g S lph r a di i f ta t 4 7 48
, , .

2 2 cu e 2 2 o un on u u as s n ec n 1
—1

air b r ba t ria 9 — 9 S lph ba t ria 5 4

, , .

o ne c e 2 1 2 2 u ur c e 1 10 1 11
— 11
4
-
.

S ar i a 6
, , , , ,

86 88 —1
c n 1 1

S arl t f v r pid mi f tra d t S nl i ght di i f ti g d t rilizi g

. .
,

c e e e e e cs o ce o u s n ec n an s e n
milk ppli 3 8 pr p rti f 7 8 ; ff t f n b
, , ,

su e s, 1 o e es o e ec o o ac

S h lz Fra z t ria 5 5 ff
.
, ,

c u e n 10 e 1

S hwa Th d r S y m bi i f ba t rial rg ani m

.
, , .
,

c nn, eo o e 64 11 os s o c e o s s, 1

S a i fl p n i b rn
. .
,

e so n, f n ue nce o u o a r -
o e

ba t ria 9
,

c e 2 1
T
S pti ta k th 3 4
.
,

e c n e 0

wa ba t ria Tax m y f ba t ria

.
, ,

Se g f
e 3
—
3 4 c e o 6 02 0 o no o c e 10 2
—1 2

S w r ba t ria i i fr m T m p rat r i fl ba t ria

. .
, , ,

e e s, 9 c e f n a r o 2 0 e e u e n ue nce o on c e

S h ath 4 ; rat pr d ti f
, .
, , ,

e 37 s, 2 1 , 34 f f 1 -
1 0 e o or o uc on o

S ilv r alt a ti pti 4 7 lig ht by ba t ria C ld

.
,

e s s as n S se d cs, 1 c e 2 00 ee o an

S ki ba t ria f th 3 9 3 2 H at
. .
,

n, c e o e 1
— 1 e

S li m ba t rial m a mp i g T tra i 6
. .
,

e c e sse s co os n e co cc 1

Thall phyta la ifi ati n f ba t ria

.
, , ,

38 o c ss c o o c e

S li m y m ilk ba t ria in 3
.
,

3 3 9 c e 12
-
1 as, 1 1 -
12 0

S il ba t ria f th Th rm al d ath p int th 3 4 3 5 3 8

. .
, ,

—1
o 8 — 87
c e o e 2 2 2 e e o e, 1 ,
1

S r m ilk b tyri a id ba t ria i nf t

, ,
.
,

ou 4 u c c c e ec 1 0

Th rm phili ba t ria 3 6
.
,

i g n 2 10 e o c c e 1

Thi ba t ria ( lph r ba t ria) f m

.

S palla za i L azzar
,
. ,

n n o , 10 o c e su u c e a

ily f 5 4
,

i ba t ria
.
,

Sp eces o f 8 — 85 c e 4 86 88 2 2 o 1 10 111 11 1
-
1
3
-

Thi th rix gr p 3 4 ; xam pl f

.
, ,

gr p 4
, .
, ,

—2
S ph til
a ro us ou 2 o ou 2 an e e o

l rl gr p f lph r ba t ria
,

S pirilla d fi iti ; dir t i

, .

f — e n o n o 1 2 1 ec on co o e ss ou o su u c e
9 ,

f divi i in r pr d ti n by fi i n
, ,

86 87 —1
o s o n, e o uc o ss o 1

T xi ba t rial pr t i
.
,

6— e ns,
43 49 5 57 o c c e o 2 2 2 .

T xi 4 8 5 5 4
,

pirilla i la
.

S M g ce a8 —
9 u 10 10 o ns, 2 2 1
-
2

Tra m i i f ba t ria fr m
.
,
.
,

Spi h t
ro c a e s, 2 0 i ns ss o n o c e o o ne n

S p ta g rati n th y 8 3 divid al t a th—r 3 9 4

.

—1 no —2 2
o n ne o u s e ne o eor u o e 2

r pr d ti by 4 6 Tri h ba t ria — 4 ; r pr d ti
.
,

S p r f rm ati
.
,

o e o o n, e o uc on c o c e 2 1 2 e o uc on

a g
,

h i al tr t r
,

5 ; 0 m c e f 6 5 m c t h 5 5 s uc u e o ,
. on e ,
0 1 .
3 4 0 I N DE X
T ri ch o b a cte ri na lig ht 57 58; ( hi g h r ba t ria )
ff t f m ti n e c e s ub o n, 1
-
1 e ec o o o

rd r ba t ria 63 ; ba t rial pi g
,

o e s O f, 10 9 f 1 11 o o n c e 1 c e
-

Tr ph tr pi t l bl i 9 4 9 5 ; pig
.
, ,

o o o sm
4 m 1 1 en s so u e n, 1
-
1

T b r l i di trib ti f by b t i l bl i 9 5 9 6 ; p
.
,

u e cu o s s, m s u on o ac en s nso u e n, 1 -
1 re v a

t ria i th air 9 9 ; a m g l f ba t ria i rigi

,

e n e 93 2 1—2 2 on e nce o c e n, 2 o n
dairy attl 3 4 3 5 f ba t ria i p i f
,

c e 1
— 1 o n,
94 95 ; c e 2 2 s eces o
-

Ty dall izati 8 —8 ba t ria in 9 5 9 8

.
,
”
n o n, 0 1 c e 2
-
2

Typh id pid m i 9 6 9 8 ; d t Wat r a aly i ba t ri l gi al 9 9

. .
,

o e e cs , 2
-
2 ue o e n s s, c e o o c 2

m ilk 3 6 3 7
,

3 1 -
1 01

W ig ht f av rag ba t ri m 9
. .
,

e o e e c e u 2

Will— th wi p ph m
, .

8— O
’
e s e no e no n, 1 1
9 99
- -

Wi aid f a ti a id ba t ria in m k
.

ne , o ce c c c e a
Va i 67
cc ne s , 2 ing 9 2 0
V g tabl ki g d m la ifi ati n f
.
.
,

e e e n o c ss c o o

ba t ria i 5 6
,

—
c e n, Y
11 11

V g tabl lt r m di a 7 7
.

e e e s a s cu u e e 1
-
2

V g tat v f fn f b t 4 6
.
,

e e i
Y a t r lati n hip f t ba t ria
e or s O ac e n a . e s s, e o s o o c e
V g ar a t
i ne d f rm tat n ce
; n r
ic
d in f rm ntiv
a cI e en
.

io 111
12 1 12 2 co ce ne
,
,

e e
,

e
m a fa t r
, -

fnu 9 prc u e o 6 2 0
o ce s se s , 2 0
Vir l pr p rty f 67 68
.
, .

u e nce , o e o 2 —2 .
,

Z
W
Z it q t d ff t f l tri ity e uo e on e ec o e ec c on

Wall f ba t rial ll 5 —5 ; h m i ba t ria 6 — 6

,

s o c e ce s, 2 3 c e c e 1 0 1 2

m p iti Zi hl arb l f h i f tai i g 9 7 :

.
,
’
l ca f 63 64
co os on o -
e s c o uc s n or s nn
Wat r r q i it f gr wt h f ba t ria Z o gl a f rm ati f 3 5 3 6 3 7 ff
.
, ,

e e u s e or o o c e o oe o ono
—

3 ; g rm i idal a ti Zy m g i ba t ria
.
, , , , ,

— f c on o s un n c
9 12 1 1 e 5 c o e c e 2 02 2 1
-
.
,
 Labo ra t o r
y G u id e i n El e m e n ta r
y
B a c te ri o l o gy
By W I LL I AM D O D G E F R O S T Ph D , . .

Ass i s ta nt Pro fe s so r o f B a ct e ri o l o gy U niv e rs it y o f Wi sco nsi n , .

Third Re vi d se E diti on . In t rl av d
e e e . I ll trat d us e . Ne w
Y rk o ,
1 9 04

Cl o th , 3 05 p age s, 8710, ne t

Th e o bj t
thi b k i t giv ad q at dir ti n f th p
ec of s oo s o e e u e ec o s or e er

f m n
or f
a rtai n f ndam ntal x r i in ba t ri l gy Th
ce o ce u e e e c se s c e o o . e

g n ral dir ti n ntai n th

e e ecnti l p art
o f th x r i whi h
s co e e s se a o e e e c se c

d n t p rm it f ny n id rabl variati n whil th p ial

oes o e O a co s e e o ,
e e s ec

d ir ti n
ec m bra
o h f at ur
s e m t ubj t t m d ifi
ce s uc e es as are os s ec o o

ca ti no f i n ta n th p rti lar rga ni m t b u e d th

,
as, or s ce , e a cu o s o e s , e

kind f m di um th i n ubati n t m p rat ur t

o e ,
e c o , e e e , e c .

M a nu a l o f De te rm i na tiv e
B a c te rio l o gy
By F R E D E R I C K D . C HE ST E R
B a t ri l gi t f th D lawar C ll g Agri l t ral E p i
c e o o s o e e e o e e , cu u x er

m nt S tati n
e d Dir t r f th L ab rat ry f th S tat
o ,
an ec o o e o o o e e

B ard f H alth f D lawar

o o e o e e

Cl o th , 4 0 1 p age s, 872 0, ne t

Th e p re se nt tabl e s se rv e o nl y fo r p urp o se s o f id e ntifi catio n and ,

no t ne ce s s aril y fo r th o se o f cla ss i fi cati o n To th e s t ud e nt w o rk .

i ng i n th e lab o rat o ry th e d e t e rm i nati o n o f unkno w n ba ct e ri a ha s .

b e e n al m o s t i m po s sibl e W ith th e use Of th e pre se nt m a nual it is

.

b e li e v e d that th e t e a ch e r ca n p la ce a give n cult ure in th e hand s o f

h i s p up il and e x p e ct h im to d e t e rmi ne it as is d o ne with o th e r ,

o rga ni c fo rm s T h e w o rk d o e s no t clai m to b e a t e xt b o o k o n
.
-

b a ct e ri o l o gy b ut aim s o nl y to s upp l e m e nt th e latt e r

,
.

UBL I S H E D BY
P

TH E M A C M I L L A N C O M PA NY
6 4 6 6 F i fth A n
-
N wY k ve ue ,
e or
The P ra c ti c a l M e th o d s Of —
O rga ni c
C h e m i stry
By L UD W I G G AT T ER M AN Ph D ,
. .

Pr f r in th Univ r it y f Fr ib rg Tra n lat d b y

o e sso e e s o e e s e
W illiam B S h b r P h D A i t nt Pr f r f Orga ni
.

c oe e . ss s a o e ss o o c
C h m i try i n L high U niv r it y N w Y rk S nd
. .
, ,

e s e e s e o e co
E diti n
.
,

o n t e

M e th o d s o f O rga ni c A n a l ysi s
By HE N RY C . S HE R M AN , Ph . D .

Adj n t Pr u c o fe s so r o f An al y ti al c C h m i try
e s in C o l um bia
Univ r ity e s . N e wYo rk ,
1 90 5
Cl o th , 2 43 p g
a e s, ne t

The o re ti c a l
O rga ni c C h e m istry
By J UL I US B C O HE N .

L nd n 9 7 o Cl th 5 0 p g
o ,
1 n t
0 o , 0 a e s , 12 772 0, e

Alth gh thi ou writt n fr m th Engli h ta nd p i nt y t it h

s Is e o e s s o e as
r iv d a nti nuall y n r a i ng i n it fi r t i ntr d u ti n t
,

e ce e co I c e s use s ce s s o c o o
Am ri a n p r f
e c r f Orga ni C h m i try Th b k m t b
o e sso s o c e s e oo se e s o e
p arti ularl y w ll ada p t d t Am ri a n n d nd f ab ut th
.

c e e o e c ee s, a o o e
right i z nd grad f di ffi ult y I t p r babl y m t n arl y
S e a e o c o os e
a ppr a h th p rf t t xt b k f u with a ll g la in
.

o c es e e ec e oo or se co e e c ss

thi bj t f ny n w p ubli h d
s su ec o a o s e .

P ra c ti c a l O rga ni c C h e m is try fo r

A d v a n c e d S tu d e n ts
By J UL I US B C O HEN .

L nd n 9 7 o Cl th 7 p g
o ,
1 n t 0 o ,
2 2 a e s, 12 7710, e

T hi b k alth gh n t ai m i ng t b a
s oo m p l t lab rat ry g id
ou o o e co e e o o u e

i i nt nd d t p r vid a y t m ati r f p ra ti al i n tr ti n
, ,

s e e o o e s s e c co u se O c c s uc o

whi h hall rv t a m p a ny th a th r T h r ti al O rga ni

c s se e o cco e u o
’
s eo e c c

C h m i try m bi nati n f th tw b k W ill b f nd

”
e Th s e co o o e se o oo s e ou

m t ati fa t ry
.

os s s c o .

PU B L I S H E D BY
THE M A C M I L LA N C O M PA N Y
64 6 6 F i fth A n
-
N wY k ve ue , e or
A n I ntro d u c ti o n to Zo o l o gy
By R O BE R T W . HE G N E R , Ph D . .

I ns tru t r in Z l gy in th U niv r it y
c o oo o e e s of M i higa n
c

On y l a f w a ni m al b l ngi ng t th m r i m p rta nt p hy la
e s e o o e o e o ,
as

vi w d fr m n v l uti nary t nd p i nt
e e o n id r da Th y e o o s a o ,
a re co s e e . e a re ,

h w v r i nt n iv l y t di d in n nd av r t t a h th funda
o e e ,
e s e s u e a e e o o e c e

m nt l p ri n i p l
e f Z l gy in a wa y that i n t p
a c ibl wh n a es o oo o s o o ss e e

up rfi ial
s xa m i nati n f t yp fr m all th p hyla i m ad
e c e o o es o e s e .

F u th m r _m p h l gy i n t p iall y m ph a iz d b ut i
er o re ,
or o o s o s ec e s e , s

co o rdi nat d with p h y i l gy l gy nd b havi r nd rv t

e s o o ,
e co o ,
a e o ,
a se es o

ill u trat b y a m p arativ t ud y th p r babl ur f v l ti n

s e co e s e o e co se o e o u o .

Th a ni m l e n t tr at d i n rt bj t f di ti n b ut
a s a re o e e as e O ec s or sse c o , as

livi ng rga ni m wh a tiviti a f fundam nt l im p rta n

o s s o se c es re o e a o ce .

N arg um nt
O n ary t j u ti fy th t yp ur
e s d a re e ce ss o
'

s e
“
e co se ,
e

ve ol p d wi th th p r bl m f rga ni v l uti n in m i nd nd
e e o e s o o c e o o ,
a

d ali ng with d ynam i

e w ll tati p h n m na c as e as s c e o e .

hav r d y ur h a p t r (Th Crayfi h nd Arth p d in

I e ea o c e e s a ro o s

G n ral ) nd
e e n xp r m y ati fa ti n with r f r n t th
a ca e e ss s s c o e e e ce o e

g n ral arra ng m nt f th m att r

e e w ll with r f r n t e e o e e ,
as e as e e e ce o

th d ta il
e T h wh l tr a tm nt i up t dat
e . tak i ng a nt f
e o e e e s o e ,
cco u o

th m d rn advan m nt in u kn wl dg
e o e f th yfi h nd ce e o r o e e o e cra s e s, a ,

chi f f all th m r i m p rta nt f at ur in th nat ur l hi t ry

e o ,
e o e o e es e a s o

o f th a nim al v ry p r p rl y p a at d fr m th ni m p rtant
e se s a re e o e se r e o e u o

on I thi nk t h i
es . ha p t r giv th n f what w k n w s c e es e e sse ce o e o

ab ut yfi h
o nd ny t ud nt m ight
cra ths b k d n
e s, a a s e u se e oo a va

t g a ly I n fa t I k n w n
e o us th r t xt b k whi h giv u h a
. c ,
o o o e e -
oo c es s c

w al th f i nf rm ati n up n f w p ag — P OF S S O A E
e o o o o so e es .
”
R E R . .

O HM AN N C rn gi M u um
RT ,
a e e se .

Th e p la n
v ry ati fa t ry nd th b k w ill b v ry in is e s s c o ,
a e oo e e

stru tiv f la u I m v ry glad that y u hav h n th

c e or c ss se . a e o e c o se e

b y
e e as r
u i n t t yp ( C ha p tor X II ) — D E E P HILLIP Sse c e . e . R . . .
,

D p ar tm nt f Agri lt ur W a hi ngt n DC
e e o cu e ,
s o ,
.

P U B L I S H E D BY
THE M A C M I L L AN C OMP A N Y
64 6 6 F i fth A nu
-
N wY k ve e ,
e or
B a cte ri a i n R e l a ti o n to C o untry Life
By JAC O B LI PM AN , M P , Ph D G . . . . . ne t

di i r bl h alth
A scuss o n o f th e p o e m o f e a nd co m fo rt in th ntry e co u
as aff t d b y th m i nut rga ni m whi h fl at in th air w
ec e e se e o s s c o e e
br ath nd in th w t r w dri nk nd p rf rm n i m p rta n t w rk
e e a e a e e a e o a o o
in th il fr m whi h
e so f d i xtra t d — R i w f R i w
o c o ur oo s e c e .
”
ev e o ev e s .

M u h pa alm t a third f th b k i q it p r p rl y giv n

“
c s ce os o e oo s u e o e e
t th r lati n f ba t ria t th il nd t th gr wth f pla nt
, ,

o e e o o c e o e so a o e o o s
T hi p art f th ubj t i v ry i nt r ti ng r adi ng nd h w
.

s o e s ec s e e es e a s o s
l arl y h w m p l x th p r bl m f i nti fi farm i ng hav b
c e o co e e o e s o sc e c e e
co m nd h w w rth y f x p n iv
e a x p ri m ntati n S ar l y l
o o o e e s e e e e o c ce e ss
i nt r ti ng alth gh x di ngl y bri f i th di u i n f th
.

e es ou e ce e e s e sc ss o o e
wag pr bl m — N w Y k E ni ng P t
, ,
”
se e o e . e or ve os .

Di se ase s f E co no m i c P l ants
o

By F L S T E V EN S Ph D . .
, . .

Pro fe sso r o f B o ta ny a nd V e ge tabl e Path o l o gy o f th e N rth o

C ar li n o a Co e e o f A ll g gri lt r nd M ha i cu u e a ec n c A s, rt a nd

B i l gi t o o s o f th e A cu gri lt ural Ex p rim nt S tati e e o n, a nd

J . G . HALL ,
M A . .

i ta nt in V g tabl Path l gy N rth ar li na

/
Ass s e e e o o i n th e o C o
A gri ult ural Ex p ri m nt S tati n
c e e o

Cl o t z, 5 13 page s, 1 2 7m , ne t

T hi i d ign d a t xt b k t m th n
s s es f that la
e f as e -
oo o ee t d e ee s o c ss o

st ud nt f e Agri lt ral C ll g
s o nd Agri ult u ral H igh
o ur cu u o e es a c

S h l wh wi h t l ar n t r
c oo s gni z nd tr at p la nt di a
o s o e o e co e a e se se s
with ut th b rd n f l ng t ud y t th ir a
o e I t i ndi at
u e o o s as o e c use s c es

hi f h ara t ri ti f th m t d tr u tiv p la nt di a
.

th e c e c c e f s cs o e os es c e se se s o

th U nit d S tat
e a u d b y ryp t g m i p ara it f ngi ba t ria
e e s, c se c o a c s e s, u c e
li m m ld nd giv i nf rm ati n r gardi ng th b t
, ,

and s e ou s, a es o o e e es

m th d
e f p r v nti n
o ur
s o f th di W hil in the e o or c e o e se s e a se s e e

m ai n n n p ara iti di a n t di d a f w f th m t
.
,

o -
u s c se s e s a re o sc s s e e o e os

n pi f thi la bri fl y m nti n d al d i a

, ,

co u s cuo s o s c ss a re e e o e a s a re so se se s
a u d b y th m t mm n p ra iti fl w ri ng p lant
,

c se e os co o a s c o e s .

T e x t bo o k
-
o f P l a nt Di se ase s
C a use d b y C ryp t o gam i c P aras it e s
By G EO R G E M ASS EE
L nd n T hird E diti n 9
o o ,
o ,
1 07 /
Cl o t i , 45 8p age s, 1 2 172 0, ne t

P U B L I S H ED B Y
THE M A C M I L L AN C O M P AN Y
64—6 6 F ifth A nu N wY k ve e ,
e or