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The Biology of Death
The Biology of Death
T H E B I O L O G Y O F
D E A T H
BY
R A Y M O N D P E A R L
TMR JOHNS tmvKtm vmvumtrt
CHAPTBB PAQB
I. THE PBOBLEM 17
II. CONDITIONS OP CELLULAR IMMORTALITY 51
III. THE CHANCES OP DEATH 79
IV. THE CAUSES OP DEATH 102
V. EMBRYOLOGY AND HUMAN MORTALITY 138
VI. THE INHERITANCE OP DURATION OP LIPE IN MAN 150
VII. EXPERIMENTAL STUDIES ON THE DURATION OP LIPE 186
VIII. NATURAL DEATH, PUBLIC HEALTH, AND THE POPULATION PROBLEM 223
BIBLIOGRAPHY 259
INDEX 269
ILLUSTRATIONS
FIG. PAGE
1. Photograph of John Shell, claimed to be 131 years old, but actually
about 100, with his wife and putative son (From Nascher) 26
2. Showing the changes in nerve cells due to age (From Donaldson
after Hodge) 29
3. Paramecium, viewed from the oral surface (From Jennings) 31
4. Diagram showing the process of reproduction by fission in the uni-
cellular organism Paramecium 32
5. Conjugation in Paramecium 32
6. Planaria dorotocephala (From Child) 34
7. Beginning of process of agamic reproduction by fission in Planaria
(From Child) 35
8. Progress of agamic reproduction in Stenostomum (From Child) 36
9. Section across the posterior part of an embryo dog-fish (Acanthias)
of 3.5 mm. (From Minot after Woods) 38
10. First and second division in egg of Cyclops (From Child) 39
11. Diagram to show mode of descent (Modified from Jennings) 41
12. Artificially parthenogenetic frogs (Loeb) 52
13. Piece of tissue from frog embryo cultivated in lymph (From Harrison) 58
14. Group of nerve fibers which have grown from an isolated piece of
neural tube of a chick embryo (From Harrison after Burrows) 59
15. Human connective tissue cells fixed and stained with Giemsa stain
(After Losee and Ebeling) 60
16. Pennaria (From Wilson) 62
17. Culture of old strain of connective tissue (Ebeling) 63
18. Life table diagram 81
19. Comparing the expectation of life in the 17th century with that of
the present time 84
20. Comparing the expectation of life in the 18th century with that of
the present time 86
21. Comparing the expectation of life of ancient Egyptians with that of
present day Americans 88
22. Comparing the expectation of life of ancient Romans with that of
present day Americans 90
13
14 ILLUSTEATIONS
FIG.
23. Comparing the expectation of life of the population of the &
provinces Hispania and Lusitania with that of preset
Americans * *
24. Comparing the expectation of life of the population of the P*
provinces in Africa with that of present day Americans •*
25. Showing Pearson's results in fitting the dx line of the life tab*1*
5 skew frequency curves •*
26. Showing the relative importance of the different organ syst1***
human mortality
27. Diagram showing the specific death rate at each age for death1*
all causes taken together •-
28. The specific death rate at each age from breakdown of the circ**^
system, blood and blood forming organs •*
29. The specific death rate at each age from breakdown of the
tory system
30. Specific death rates at each age from breakdown of
secondary sex organs * *
31. Specific death rates at each age from breakdown of the kidney**
related excretory organs *-
32. Specific death rates at each age from breakdown of the skeleton-1
muscular systems *
33. Specific rates of death at each age from breakdown of the alimt* *
tract and associated organs of metabolism
34. Specific death rates at each age from breakdown of the nervotiJ
tern and sense organs »
35. Specific death rates at each age chargeable against the skin., ,
36. Specific death rates at each age from breakdown of the endrm
system , ,
37. Specific death rates from all othsr causes of death not covered •
preceding categories , ,
38. Percentages of biologically classifiable human mortality resulting
breakdown of organs developing from the different germ lay**
39. Specific death rates in males according to the germ layer from m
the organs developed „
40. Specific death rates for females...V „,
41. Survival curves of members of the Hyde family (Plotted from ]
data) „
ILLUSTRATIONS 15
FIC. *AGE
42. Influence of father's age at death upon longevity of offspring (After
Bell) 156
43. Influence of mother's age at death upon longevity of offspring (After
Bell) 157
44. Influence of age at death of parents upon the percentage of offspring
dying under 5 years (After Ploetz) 178
45. Snow's results on selective death rate in man 182
46. Male and female fruit fly (Drosophila melanogaster) (From Morgan). 187
47. Life lines for Drosophila melanogaster 188
48. life lines for different inbred lines of descent in Drosophila 192
49. Life lines showing the result of Mendelian experiments on the dura-
tion of lif e in Drosophila 195
50. Distribution of poverty in Paris U911-13). (After Hersch) 202
51. Death rates in Paris (1911-13) from all causes. (After Hersch) 203
52. Trend of death rates for four causes of death against which public
health activities have been particularly directed 230
53. Trend of death rates from four causes of death upon which no direct
attempt at control has been made 232
54. Trend of combined death rate from the four causes shown in figure
52 as compared with the four causes shown in figure 53 233
55. Course of the weighted average death rate for the countries in the A
and B groups, from typhoid fever 236
56. Like figure 55, but for diphtheria and croup 237
57. Record of. malaria control by antimosquito measures, Crossett, Ark.,
1916-1918. (From Rose) 241
58. Disappearance of yellow fever from Guayaquil, Ecuador, as a result of
control measures. (By permission of International Health Board) 242
59. Showing the change in percentage which deaths were of births in
each of the years 1912 to 1919 246
60. Theoretical curve of population growth 249
61. Curve of growth of the population of the United States 250
62. Curve of growth of the population of France 251
63. Curve of growth of the population of Serbia 253
64. Growth of a Drosophila population kept under controlled experi-
mental conditions 254
T H E B I O L O G Y O F
D E A T H
CHAPTER I
THE PROBLEM
In p r e s u m p t i v e VUHVH of K^**nt l o n g e v i t y , w h i c h o n
o t h e r g r o u n d n a r e w o r t h y of Hericitm conHideriition, it in
u s u a l l y in r e n p e c t of i t e m c - t h e p r o o f of identity—»tlmt
t h e evidence in weaken*. Mvery ntiident of Kt*neii!ii)Lcictiil
d a t a known how cany it in for tin* following n o r t of t h i n g
t o h a p p e n . J o l u i B m i t h \vm b o r n in tlio h i t t e r Imtf of
t h e e i g h t e e n t h c e n t u r y . Ilin biiptinm wn» d u l y a n d p r o -
p e r l y registered* H e u n f o r t u n a t e l y dim! n t tln» u p * of
24 BIOLOGY OF D E A T H
say 15. B y an oversight his death w a s not registered.
In the same y e a r that he died another male child w a s
born to the same parents, and given the name of John
Smith, in commemoration perhaps of his deceased brother.
This second John Smith was never baptized. H e at-
tained the age of 85 years, and then because of the appear-
ance of extreme senility which he presented, his stated age
increased by leaps and bounds. A study of the baptiBmal
records of the town disclosed the apparent fact that he
w a s just 100 years old. The case goes out to th« public
as an unusually well authenticated case of contenarianinm v
when of course it is nothing of the s o r t
Young applies vigorously the criteria above enumer-
ated first, to the historically recorded cases of great long-
evity such as Thomas Parr, et id genus omne, and rajeets
them all; and second to the total mortality e^perknee
of all the Life Assurance and Annuity Societies of Oreat
Britain and the annuity experience of the National Debt
Office. The number of persons included in the experience
was close upon a million. H e found in thin material, and
from other outside evidence, exactly 30 persons who lived
100 or more years. I n Table 2 the detailed results of
his inquiry are shown in condensed form.
It will be noted from this table that the most extreme
case of longevity which Young was able to authenticate
was about a month and a half short of 111 y u a n . Of
the 30 centenarians recorded 21 war© women and 9 w a r t
men. The superiority of women in expectation of life is
strikingly apparent at the very high age of 100 y e a r n W e
shall later see that this is merely a particularly noteworthy
instance of a phenomenon which i s common to a g r e a t
portion of the life span.
T H E PROBLEM 25
The contrast between these proved findinffH of Young,
exceedingly m o d e s t both in r e a p e d of numbcrx, and ex-
tremity of longevity, and the Iooaedala. on c u n t r i w i
TABLK2
Authentic Imtantm of Crntrnnrinnutm (frum Ytmng)
9 M HO .Til
9 M IDH 111
9 M 105 H «. ,
9 8 104 II in
9 M 103 11 2H
9 ? 103
9 M .1 7
1 H
102 11 %
9 102 21H
9 8 uri 2 HI
Q 8 102 1 H
9 8 102 *l
9* 8 102 W
? 102 *ti
<?t 8 20!
io 4
9 8 101 K 2ft
of* ? 101 2t»
? 101 "4
9 8 101 1 ' in4
9 8 101 1
? 101 82
9 8 101 %
d1 t im *!» 4
9 8 100 7 H
9 8 IOCJ II ii
9 M 100
o* M 100 '2 21
9 8 100 1 10
cP ? 100
FIG. 2.—Showing the changes in nerve cells due to age. 1, spinal ganglion cells of a still-
t>orn male child; 2, spinal ganglion cells of a man dying at ninety-two years; N. nuclei.
I n the old man the cytoplasm IB pigmented, the nucleus is small, and the nucleolus much
edburunken or absent. Both sections taken from the first cervical ganglion, X 250
cliameters; 3, nerve cells from the antennary ganglion of a honey-bee, just emerged in the
p>erfect form; 4, cells from the same locality of an aged honey-bee. In 3, the large
xxuoleus (black) is surrounded by a thin layer of cytoplasm. In 4, the nucleus is stellate,
m d the oell substance contains large vacuoles with shreds of cytoplasm. (From Donaldson
sdfter Hodge).
T h e r e a r e o t h e r a n d p e r h a p s even m o r e g e n e r a l a n d
s t r i k i n g m o r p h o l o g i c a l changes i n senescence t h a n the
o l a a n g e d r e l a t i o n b e t w e e n c y t o p l a s m a n d nucleus.
Oonfclin says:
By all odds the most important structural peculiarity of senescence is
•fclxe increase of metaplasm or differentiation products at the expense of
-fclxe general protoplasm. This change of general protoplasm into products
o f differentiation and of metabolism is an essential feature of embryonic
differentiation and it continues in many types of cells until the entire
is almost filled with such products. Since nuclei depend upon the
30 BIOLOGY OF D E A T H
general protoplasm for their growth, they also become small in
cells. If this process of the transformation of protoplasm into differedlft>1"
tion products continues long enough it necessarily leads to the deatk^ °*
the cell, since the continued life of the cell depends upon the interact* 011
between the general protoplasm and the nucleus. In cells laden witfr ^li**
products of differentiation, the power of regulation is first lost, then tilt*
power of division, and finally the power of assimilation; and this *«
normally followed by the senescence and death of the cells.
FIG. 6.—Planaria dorotocephala: m, month: p^, pharnyx: al, alimentary tract; nt, nerroua
system. (From Child).
THE PROBLEM 35
exclusive m o d e of r e p r o d u c t i o n , i n s o m e o r all f o r m s of
t h e t h r o e lowest g r o u p s of m u l t i c e l l u l a r o r g a n i s m s , t h e
s p o n g e s , flatworms, a n d e o e l e n t e r a t e s . M o r e r a r e l y it
m a y occur in o t h e r of t h e l o w e r i n v e r t e b r a t e
g r o u p s . I t m a y o c c u r in t h e f o r m of b u d d i n g
o r of fission c o m p a r a b l e t o t h a t of t h e Proto-
zoa. T h e a g a m i c r e p r o d u c t i o n of one of t h e
Hut w o r m s , Planaria dorotocephala, studied
by the w r i t e r m a n y y e a r s a g o , a s shown in
F i g u r e 6, m a y s e r v e a s an i l l u s t r a t i o n .
T h i s s i m p l y o r g a n i z e d w o r m , which lives
u n d e r s t o n e s In s l u g g i s h s t r e a m s a n d p o n d s ,
a f t e r a t t a i n i n g a c e r t a i n size, will u n d e r t h e
a p p r o p r i a t e e n v i r o n m e n t a l c o n d i t i o n s exhibit
a c o n s t r i c t i o n t o w a r d s t h e p o s t e r i o r end of
t h e body, an shown in F i g u r e 7.
F o r a t i m e t h e a n i m a l m o v e s a b o u t as a
r a t h e r u n g a i n l y double i n d i v i d u a l . I t finally
s e p a r a t e s into two. T h e l a r g e r a n t e r i o r p a r t
f o r m s a n e w tail, a n d t h e s m a l l e r p o s t e r i o r
fission p r o d u c t f o r m s a new h e a d a n d r a p i d l y
g r o w s t o full size. T h e p r o c e s s i s , in p r i n c i -
ple, exactly the s a m e a s t h e m u l t i p l i c a t i o n of
F a r a m e e i u m by fission. I n a n o t h e r m e m b e r
of t h e s a m e g e n e r a l g r o u p of a n i m a l s a s
Planaria, n a m e d Stenostomum, s e v e r a l fis-
sion p l a n e s m a y f o r m a n d t h e p r o c e s s s t a r t
Fin.—7. flkwta-a n e w b e f o r e t h e p r o d u c t s d e l i m i t e d b y t h e
nir»jc«»f ptnmmm
ftinmtn ff*prt<rfirst p l a n e h a v e s e p a r a t e d . A s a r e s u l t , w e
<lu<U Win hyftw-
g e t f r e q u e n t l y in t h i s f o r m c h a i n s of individ-
m*m cum
u a t e a t t a c h e d in a l o n g s t r i n g to e a c h other,
a s shown in F i g u r e 8*
I t i® o b v i o u s t h a t s o l o n g a s r e p r o d u c t i o n g o e s on in
36 BIOLOGY OF DEATH
I, /./.I.
J.I.
jr.1.
1.2.
7.J.
CelU
FIG. 9.—Section across the posterior part of an embryo dog-fish (acanthias) of 3.5 mm., to
show the compact cluster of germ cells on one side. The germ cells in later stages migrate
from this primitive position, moving singly or in small groups. Ed, ectoderm: Md, medullary
canal or primitive spinal cord; Nch, notochord; Mes, mesoderm: Eni, entoderm: X, cellular
strand connecting the germ cell cluster with the yolk. (From Minot after Woods, with the
permission of the publishers, G. P. Putnam's Sons).
rial. After union has taken place the fertilized ovum
or zygote presently begins to divide, first into two cells,
these again to four and so on, until by a continuation of
this process of division with concomitant differentiation
the whole body is formed. As the animal develops by
repeated cell division and differentiation, it is frequently
found that at the very early stage the cells which are to
be the germ cells of the next generation are clearly re-
THE PROBLEM 39
cognizable b y t h e i r s t r u c t u r e , a n d often a r e set aside
in a definite location i n t h e d e v e l o p i n g e m b r y o . T h u s ,
to t a k e b u t a single e x a m p l e of a p h e n o m e n o n of wide
generality, a t a v e r y e a r l y s t a g e i n t h e d e v e l o p m e n t of
the dog-fish, "when t h e only bodily o r g a n s of which even
the r u d i m e n t s a r e recognizable a r e t h e b e g i n n i n g s of w h a t
will p r e s e n t l y b e c o m e t h e s p i n a l c o r d a n d t h e back-bone,
d i a g r a m s h o w s t h e m o d e of d e s c e n t i n f o r m r e p r o d u c i n g
from t w o p a r e n t s . T h e lines r e p r e s e n t t h e lives of indi-
v i d u a l s ( a s i n A d i a g r a m ) , or of g e r m cells (in t h e B
FIG. 11. Diagram to show mode of descent in (A) unicellular animals reproducing agamic-
ally, and in (B) multicellular animals reproducing by germ cells. For further explanation see
text. (Modified from Jennings).
diagram) beginning a t the left a n d p a s s i n g t o t h e right.
In the A diagram, which represents uniparenta! reproduc-
t i o n b y fission, t h e l i n e of a n c e s t r y t r a c e d b a c k f r o m a n y
individual at the right is always single, and there is no
corpse t o be found anywhere, each present body trans-
f o r m i n g d i r e c t l y i n t o t h e t w o b o d i e s of t h e n e x t generation.
42 BIOLOGY OF DEATH
I n the B diagram, where we have bi-parental reproduc-
tion b y t h e u n i o n of g e r m cells, as in m a n , the solid black
t r i a n g l e s r e p r e s e n t t h e bodies, o r somata, a n d t h e lines
t h e g e r m cells. A line of a n c e s t r y t r a c e d back f r o m a n y
individual t o w a r d s t h e r i g h t end of tKe d i a g r a m forks
a t each g e n e r a t i o n , a n d in c o m p a r a t i v e l y few g e n e r a t i o n s
one h a s a m u l t i t u d e of a n c e s t o r s . T h e bodies of one
g e n e r a t i o n h a v e no c o n t i n u i t y w i t h the bodies of t h e p r e -
vious o r t h e following g e n e r a t i o n . I n each g e n e r a t i o n the
soma dies, while n e w s o m a t a a r e r e p r o d u c e d by t h e union
of g e r m cells f r o m d i v e r s e lines.
E . Life itself is a continuum. A b r e a k or discontin-
u i t y i n its p r o g r e s s i o n h a s n e v e r occurred since i t s first
a p p e a r a n c e . D i s c o n t i n u i t y of existence a p p e r t a i n s n o t
to life, b u t only t o one p a r t of the m a k e u p of a p o r t i o n
of one l a r g e class of living t h i n g s . This is certain, from
the facts a l r e a d y p r e s e n t e d . N a t u r a l d e a t h is a n e w t h i n g
which h a s a p p e a r e d i n t h e c o u r s e of evolution, a n d its
a p p e a r a n c e is concomitant with, a n d evidently in a b r o a d
sense, caused b y t h a t r e l a t i v e l y early e v o l u t i o n a r y spe-
cialization which set a p a r t a n d differentiated c e r t a i n
cells of t h e o r g a n i s m f o r t h e exclusive business of car-
r y i n g on all functions of t h e b o d y o t h e r t h a n r e p r o d u c -
tion. W e a r e able to f r e e ourselves, once and f o r all, of
the n o t i o n t h a t d e a t h is a n e c e s s a r y a t t r i b u t e or inevitable
consequence of life. I t is n o t h i n g of the s o r t Life can
a n d does all t h e t i m e go on w i t h o u t death. T h e somatic
d e a t h of h i g h e r m u l t i c e l l u l a r o r g a n i s m s is s i m p l y t h e
p r i c e t h e y p a y f o r t h e p r i v i l e g e of enjoying t h o s e h i g h e r
specializations of s t r u c t u r e a n d function which h a v e been
a d d e d on a s a side l i n e t o t h e m a i n business of living
t h i n g s , which is t o p a s s on i n u n b r o k e n c o n t i n u i t y t h e
n e v e r - d i m m e d fire of life itself.
THE PROBLEM 43
THEORIES OF DEATH
O n t h e b a s i s of these five g e n e r a l c l a s s e s of f a c t s
which h a v e been briefly r e v i e w e d a whole s e r i e s of specu-
l a t i o n s a s to t h e m e a n i n g of d e a t h h a v e been r e a r e d . T h e
first a t t e m p t a t a biological e v a l u a t i o n * of t h e m e a n i n g
of d e a t h which a t t r a c t e d t h e s e r i o u s a t t e n t i o n of scientific
men w a s t h a t of W e i s m a u n . I n h i s f a m o u s a d d r e s s of 1881
on t h e d u r a t i o n of life, W e i s m a n n p r o p o u n d e d t h e t h e s i s
that death was an adaptation, advantageous to the race,
a n d h a d a r i s e n a n d w a s p r e s e r v e d by n a t u r a l selection.
P r o b a b l y no m o r e p e r v e r s e e x t e n s i o n of t h e t h e o r y of
n a t u r a l selection t h a n this w a s e v e r m a d e . I t a p p e a r e d ,
h o w e v e r , j u s t a t t h e t i m e w h e n t h e p o s t - D a r w i n i a n at-
t e m p t to s e t t l e t h e p r o b l e m s of evolution b y s h e e r dia-
lectic w a s a t t h e zenith of its p o p u l a r i t y . N o w a d a y s such
a d o c t r i n e an W e i K m a n n ' s would n o t receive so respectful
a hearing.
MclchnikofF, w h o s e views excited so m u c h p o p u l a r
i n t e r e s t s o m e y e a r s a g o , held t h a t d e a t h w a s t h e r e s u l t
of i n t o x i c a t i o n , a r i s i n g f r o m t h e a b s o r p t i o n of p u t r e f a c -
tive p r o d u c t s of the a c t i v i t y of i n t e s t i n a l b a c t e r i a . T h e
chief difficulty w i t h t h i s view is t h a t i t is d e m o n s t r a b l y
n o t t r u e ; e i t h e r p a r t i c u l a r l y in t h e c a s e of m a n , w h e r e
it can easily be s h o w n t h a t m a n y s t a t i s t i c a l l y i m p o r t a n t
c a u s e s of d e a t h c a n n o t p o s s i b l y b e a c c o u n t e d f o r u n d e r
it, o r g e n e r a l l y in t h e a n i m a l k i n g d o m ; b e c a u s e a n u m -
b e r of eascm an* now known w h e r e a m e t a z o a n f o r m can
be successfully m a d e to lead a c o m p l e t e l y a s e p t i c life,
a n d still d e a t h o c c u r s a t a b o u t t h e u s u a l t i m e . (Of. C h a p -
t e r V I I I ) . M o r e npeculntive d e v e l o p m e n t s of t h e name
of varlouH theories of dt»ath, which the
writ«*r» though iliflforitiK (mm wmw of the e<mdu«ionii, h&» found useful
in th«t preparation of thin iiectUm, ha» l&taly Imn given by Child in his
and
44 BIOLOGY OF DKATH
b a s i c i d e a h a v e been p r e s e n t e d by Jiekeli a n d Moritj|
e r y . B o t h held t h a t because of the* mechanical i u r o m p
n e s s of the processes of m e t a b o l i s m , i n j u r i o u s find 1
s u b s t a n c e s tend t o a c c u m u l a t e in the* w l l s of th<* b
a n d t h a t senescence a n d d e a t h an* flu* n*Hti]tji
such accumulations.
A much b r o a d e r , a n d in the* l i g h t of all facts sous
view, is t h a t t h e d e t e r m i n a t i o n of di»groc*H of long*.;
a n d of t h e f a c t of d e a t h itself, is i n h e r e n t in tin* ini
h e r e d i t a r i l y d e t e r m i n e d biological c o n s t i t u t i o n of tin
dividual a n d t h e epecies. Thin view wan <»xpr*'88e<
J o h a n n e s Miillor a q u a r t e r of a c e n t u r y a g o in his Pi
ologie, by Cohnheim f o r t y yearn Inter, a n d h a s had n
l a t e r a d h e r e n t s . I shall r e t u r n to a diHcuHHion of it h
T h e r e h a v e been a n u m b e r of t h e o r i e s of HIMIPHI?
a n d d e a t h , differing widely in d e t a i l s , b u t h a v i n g t h e
p o i n t in c o m m o n of a t t r i b u t i n g t h e s e phcmotmmi
o r d e r l y c h a n g e s with a d v a n c i n g a g e in the* r e l a t i v e
p o r t i o n of nucleus to p r o t o p l a s m in t h e cells of Hit* h
H e r e m a y be m e n t i o n e d , w i t h o u t p a u s i n g t o go intn
tailed c o n s i d e r a t i o n of t h e i r different vjewn t V«*niw
M u h l m a n n , R i c h a r d H e r t w i g , a n d Minot.
A n o t h e r g r o u p of h y p o t h e s e s , till a d v a n c e d in i
p a r a t i v e l y r e c e n t times a n d associated w i t h the n a n u
K a s s o w i t z , Conklin, a n d Child, art! develojiecl ubout
metabolic a s p e c t s of a g e changed. T h e n * in nlmervi
d e c r e a s e in a s s i m i l a t o r y c a p a c i t i e s of mlh w i t h iliflfc
t i a t i o n a n d age- T h e s e m e t a b o l i c c h a n g e s a r e regal
a s f u n d a m e n t a l l y c a s u a l of t h e p h e n o m e n a of nanotie
a n d d e a t h . I n t h i s g e n e r a l g r o u p of hypotht?nf!« w
belong t h e v i e w s of m y colleague, D r . W . T . I l o w a r
B e n e d i c t in a detailed invegtigation of nentlttj
plants reaches the conclusion:
THE PROBLEM 45
"that the duration of life is directly linked with the degree of permeability
in that part of the living cell which places it in contact with the universe
about it, and that as the activities of life proceed the cell is being gradually
entombed by an inevitable decrease in the permeability of its protoplasm.
While decreasing permeability furnishes a possible explanation of the
more obvious symptoms of aenility, it cannot be the only degeneration of
first rank. All protoplasmic functions must be involved. Underlying these
primary causes of senile degeneration there must be some general funda-
mental cause from which they spring. This fundamental cause may well
be the colloidal nature of protoplasm."
D e l a g e a n d J e n n i n g s h a v e c o n s i d e r e d t h a t d e a t h is
t h e r e s u l t of diflferentiatiort J e n n i n g s h a s p u t t h e m a t t e r
in t h i s w a y
"the continuity of life in the infusoria is in principle much like that in
ourHclv«»Hr though with differences in details. As individuals, the infusoria
do not die, save by accident. Those that we now see under our microscopes
have bftem living ever since the beginnings of life; they come from division
of previously existing individuals. But in just the same sense, it is true
for ourHftlvcw that everyone that in alive now has been alive since the
beginning of lif«>. This truth applies at least to our bodies that are alive
now; every cell of our IKKIIGH IH a piece of one or more cells that existed
earlier, And thun unr entire body van be traced in an unbroken chain as
fur bitek into time* an life goes. The difference is that in man and other
higher orgaitinmM there have been left all along the way great masses of
relit* that did not continue to live. Thme masses that wore out and died
,ar# what wt? eall th« bocltati of the persons of earlier generations j but
tmr own bodie* nr? not defended by cell division from these; they are the
continuation of <'«?llf» that have kept on living and multiplying from the
f»arli<tf§i limit*, jtmt an have the existing infusoria."
' viewa r e g a r d i n g senescence in t h e p r o t o z o a
will be d i s c u s s e d in t h e n e x t c h a p t e r .
U n i c e l l u l a r o r g a n i s m s , an we h a v e seen, do n o t nor-
mally experience natural death. In the higher organisms
t h e r e h a s been 11 p r o g r e s s i v e n e t t i n g a p a r t of cells a n d
tiflatlet* t o p e r f o r m particular vital functions w i t h a con-
s e q u e n t U>m of t h e ability t o p e r f o r m all v i t a l functions
i n d e p e n d e n t l y . A s noon an a n y o n e of t h e s e cells o r t i s s u e s
begin*, f o r a n y a c c i d e n t a l c a u s e w h a t e v e r , t o fail t o per-
46 BIOLOGY OF DEATH
f o r m its special function p r o p e r l y , i t upsets the d e l i c a t e
balance of t h e whole a s s o c i a t e d community of cells and t i s -
sues. B e c a u s e of t h e differentiation and specialization o f
function, t h e p a r t s a r e m u t u a l l y dependent u p o n each,
other t o i e e p t h e m s e l v e s a n d t h e whole going. C o n s e -
q u e n t l y a n y d i s t u r b a n c e i n t h e balance which i s n o t
p r o m p t l y r i g h t e d b y some r e g u l a t o r y process m u s t e v e n -
tually end in death.
Since t h e publication of this m a t e r i a l in serial form,
a n objection t o t h e f o r e g o i n g s t a t e m e n t has been s u g -
g e s t e d on t h e g r o u n d t h a t differentiation per se does n o t
a p p e a r t o t h e critic t o h a r e m u c h t o do with the question of
n a t u r a l d e a t h i n the Metazoa. T o q u o t e ; " r a t h e r it is t h e
f a i l u r e after differentiation t o k e e p u p indefinitely t h e
s t a t e r e a c h e d . If, f r o m a n y i n t e r n a l or external a c c i d e n t ,
the differentiated p a r t suffers i n j u r y , the injury c a n n o t
be m a d e good a n y m o r e , since i n certain organs t h i s p o w e r
h a s b e e n lost. H e n e e , i n t i m e , loss after loss occurs a n d
the m a c h i n e w e a r s out. T h e p r o t o z o a n is as highly differ-
e n t i a t e d a s a n y cell of a m e t a z o a n ( o r much m o r e s o ) ; b u t
since i t " m u l t i p l i e s b y d i v i d i n g , " i t has r e t a i n e d t h e
p o w e r to m a k e g o o d a n y loss. Therefore, it is not t h e
differentiation per se, but t h e l o s s of power t o r e p a i r
that produces senescence."
T h i s seems t o m e t o "be i n t h e m a i n only a s o m e w h a t
different f o r m of s t a t e m e n t of precisely the idea t h a t I
h a v e e n d e a v o r e d to e x p r e s s . W h e n I have used the t e r m
" d i f f e r e n t i a t i o n " in this connection, I have always h a d
i n m i n d , a s o n e of i t s m o s t i m p o r t a n t physiological con-
c o m i t a n t s , j u s t t h e t h i n g s p o k e n of above. F u r t h e r m o r e ,
w h e t h e r t h e p r o t o z o a n cell i s a s h i g h l y differentiated a s
a m e t a z o a n cell, i s n o t t o t h e p o i n t at all. F o r , t o h a v e
a n y p e r t i n e n c e so^-far as t h e p r e s e n t issue is concerned,
the c o m p a r i s o n m u s t be between t h e differentiated p r o t o -
THE PROBLEM 47
zoan cell, and the whole metazoan soma, n o t one of its
c o n s t i t u e n t cells. I n t h e p r o t o z o a n , all t h e differentia-
tions a r e i n a n d a p a r t of one single cell o p e r a t i n g as one
metabolic u n i t , of small a b s o l u t e size, a n d c o n s e q u e n t l y
easier a n d m o r e labile i n t e r n a l physico-chemical r e g u l a -
tion. I n t h e m e t a z o a n s o m a we h a v e o r g a n differentia-
tion, w i t h t h e c o n s t i t u e n t cells i n e a c h o r g a n h i g h l y
specialized functionally, a n d d e p e n d e n t u p o n t h e n o r -
m a l functional a c t i v i t y of wholly o t h e r o r g a n s i n o r d e r
t h a t t h e y m a y k e e p g o i n g a t all. Remove these
tissue cells f r o m t h e soma, a n d p r o v i d e t h e m w i t h a n
a b u n d a n c e of s u i t a b l e n o u r i s h m e n t a n d o x y g e n , a s in
t i s s u e c u l t u r e s , a n d , so f a r as t h e evidence n o w a v a i l a b l e
indicates, t h e y will live f o r e v e r (cf. C h a p t e r I I ) .
Consider f o r a m o m e n t t h e m o s t h i g h l y d i f f e r e n t i a t e d
p r o t o z o a n k n o w n , o n the one h a n d , a n d m a n , on t h e o t h e r
h a n d , p u r e l y a s physico-chemical m a c h i n e s , w h i c h only
keep going if t h e i n t e r n a l balances a n d a d j u s t m e n t s a r e ,
in each case, held w i t h i n a n a r r o w zone of n o r m a l i t y .
Quite aside f r o m a n y q u e s t i o n of t h e i r different m o d e s
of r e p r o d u c t i o n , t h e t w o m a c h i n e s a r e n o t equivalent,
as machines, because of : ( a ) u n i c e l l u l a r v e r s u s multicel-
l u l a r s t r u c t u r e , (b) g r e a t absolute difference i n size of
t h e whole m a c h i n e s , w i t h c o n s e q u e n t r e q u i r e m e n t of a n
enormously m o r e complex i n t e r n a l r e g u l a t o r y m e c h a n i s m
in the one case t h a n i n t h e other, w h a t e v e r t h e i n h e r e n t
n a t u r e of t h i s m e c h a n i s m m a y be.
E s s e n t i a l l y t h e s a m e view of t h e m a t t e r a s t h a t
held by t h e p r e s e n t w r i t e r h a s b e e n well s e t f o r t h b y L o e b
in his m o s t r e c e n t p a p e r on t h e subject. H e s a y s :
"All this points to the idea that death is not inherent in the individual
cell, but is only the fate of more complicated organisms in "which different
types of cells or tissues are dependent upon each other. In this case it
seems to happen that one or certain types of cells produce a substance or
substances which gradually become harmful to a vital organ like the res-
48 BIOLOGY OF DEATH
piratory center of the medulla, or that certain tiM»tu*x cutmtnw or <I«»wtr<v
substancea which are needed for the life of muw vital orjk'ftn. Th« iium-hit*
of death of complex: organitnim way then be trarni U> th« activity of j
black sheep in the Hociety of tinnueH and organ** whirl* rcm»tttute & vmu
plicated rnulticellular organiHin."
A t this p o i n t 1 shall n o t s t a y to dincuHH critically each
of the h y p o t h e s e s so s u m m a r i l y reviewed, liintcad, I
shall m a k e bold to s t a t e n o m e w h a t c a t e g o r i c a l l y my own
views on t h e o r i g i n a n d m e a n i n g of d e a t h a n d t h e d e t e r -
mination, of l o n g e v i t y ; a n d in w h a t follow**, shall e n d e a v o r
to s e t f o r t h in o r d e r l y a r r a y t h e evidence winch twemH to
m e t o s u p p o r t t h e s e views. I n t h i s p r o c e s s , t h e r e l a t i o n s
of w h a t I shall s u g g e s t to t h e conclusions of e a r l i e r inves-
t i g a t o r s will, I think, sufficiently a p p e a r .
L e t u s consider, then, t h e following p i c t u r e of life
and d e a t h :
1* L i f e itself is i n h e r e n t l y c o n t i n u o u s .
2. L i v i n g tilings, w h e t h e r single-celled o r many*
celled organism**, a r e e s s e n t i a l l y only phyBieoHthemieal
m a c h i n e s of e x t r a o r d i n a r y c o m p l e x i t y ; b u t rttgurdlttiw of
t h e i r d e g r e e of c o m p l e x i t y only a m e n a b l e t o , a n d
a c t i v a t e d i n a c c o r d a n c e with, p h y s i c a l a n d chemical lftw§
and principles.
3. T h e d i s c o n t i n u i t y of d e a t h is n o t a nocetmary o r
i n h e r e n t a d j u n c t o r consequence of life, b u t in a r e l a -
tively n e w p h e n o m e n o n , which a p p e a r e d only whott a n d
because differentiation of s t r u c t u r e a n d function a p p e a r e d
in t h e c o u r s e of evolution.
4* D e a t h n e c e s s a r i l y occurs only in aueh a o m a t a of
multicellular o r g a n i s m s a s hav© lost, t h r o u g h differentia*
tion a n d specialization of function, t h e p o w e r of r e p r o *
d u c i n g each p a r t if it, f o r a n y accidental rettnon b r e a k s
down o r is i n j u r e d ; o r still p o s s e s s i n g such p o w e r in t h e i r
cells, h a v e lost t h e n e c e s s a r y m e c h a n i s m for s e p a r a t i n g a
THE PROBLEM 49
p a r t of t h e s o m a f r o m t h e r e s t f o r p u r p o s e s of a g a m i c
reproduction.
5. S o m a t i c d e a t h r e s u l t s f r o m a n o r g a n i c d i s h a r m o n y
of t h e whole o r g a n i s m , i n i t i a t e d b y t h e f a i l u r e of some
o r g a n o r p a r t t o c o n t i n u e i n i t s n o r m a l h a r m o n i o u s func-
t i o n i n g i n t h e e n t i r e differentiated a n d m u t u a l l y d e p e n d -
e n t s y s t e m . T h i s f u n c t i o n a l b r e a k d o w n of a p a r t m a y
be c a u s e d i n a m u l t i t u d e of w a y s f r o m e x t e r n a l o r i n t e r n a l
s o u r c e s . I t m a y m a n i f e s t itself i n a g r e a t v a r i e t y of
w a y s b o t h s t r u c t u r a l l y a n d functionally. M a n y of t h e s e
m a n i f e s t a t i o n s w h i c h h a v e b e e n r e g a r d e d a s c a u s e s of
senescence, m a y m o r e t r u l y be c o n s i d e r e d c o n c o m i t a n t
a t t r i b u t e s of senescence.
6. A s a consequence of o u r second t h e s i s which p o s t u -
l a t e d life t o b e a m e c h a n i s m , d e a t h , w h e t h e r of a single
s o m a t i c cell o r of a whole soma, is a r e s u l t of p h y s i c o -
chemical c h a n g e s in t h e cell o r o r g a n i s m ; a n d t h e s e
c h a n g e s a r e in a c c o r d a n c e w i t h o r d i n a r y p h y s i c o -
chemical l a w s a n d p r i n c i p l e s .
7. T h e t i m e a t w h i c h n a t u r a l d e a t h of t h e s o m a occurs
is d e t e r m i n e d b y t h e c o m b i n e d a c t i o n of h e r e d i t y a n d
e n v i r o n m e n t . F o r e a c h o r g a n i s m t h e r e is a specific long-
evity d e t e r m i n e d b y i t s i n h e r i t e d p h y s i c o - c h e m i c a l con-
s t i t u t i o n . T h i s specific l o n g e v i t y is c a p a b l e of modifica-
tion, w i t h i n r e l a t i v e l y n a r r o w limits, as a r e s u l t of t h e
i m p a c t of e n v i r o n m e n t a l f o r c e s ; t h e chief m o d e of action
of t h e e n v i r o n m e n t b e i n g i n t h e d i r e c t i o n of d e t e r m i n i n g
t h e r a t e a t w h i c h t h e i n h e r i t e d e n d o w m e n t is u s e d u p .
F o r no one of t h e s e p a r a t e e l e m e n t s of t h i s p i c t u r e c a n
I c l a i m a n y p a r t i c u l a r o r i g i n a l i t y . M o s t of t h e m w o u l d
p r o b a b l y be a g r e e d t o a t once, a t l e a s t b y s o m e biologists.
T h e n e e d i s f o r a s y n t h e s i z i n g i n t o a c o n s i s t e n t w h o l e of
a w i d e range of data, w h i c h h a v e a c c u m u l a t e d in v a r i o u s
4
50 BIOLOGY OF DEATH
fields of biology, a b o u t d e a t h a m i t h e d u r a t i o n of life,
S u c h a s y n t h e s i s will be a t t e m p t e d in w h a t follows,
G e n e r a l l y , t h o s e who h a v e s p e c u l a t e d a b o u t t h e biology
of d e a t h h a v e d r a w n t h e i r evidence f r o m , o r a t l e a s t h a d
t h e i r t h i n k i n g l a r g e l y colored by t h e facts in a relatively
small p a r t of t h e whole field. I n p a r t i c u l a r , few biologists*
h a v e a n y d e t a i l e d knowledge* of t h e m o s t i m p r e s s i v e
m a s s of m a t e r i a l , both in r e s p e c t of q u a l i t y a n d q u a n t i t y ,
which e x i s t s r e g a r d i n g the d u r a t i o n of life of a n y o r g a n -
ism. I r e f e r , of course, t o the e n o r m o u s v o l u m e of
r a t h e r e x a c t d a t a r e g a r d i n g h u m a n m o r t a l i t y . Much of
this m a t e r i a l , to be mire, w a n t s p r o p e r a n a l y s i s , n o t
only m a t h e m a t i c a l but biological. B u t , t h a t it in a rich
m a t e r i a l a d m i t s of n o d o u b t .
CHAPTER II
AKTIFICIAIi PARTHENOGENESIS
T h e m o s t e s s e n t i a l condition of t h i s i m m o r t a l i t y of
g e r m cells w a s m e n t i o n e d , b u t n o t p a r t i c u l a r l y e m p h a -
sized. I t i s t h a t two g e r m cells, a n o v u m a n d a s p e r m a t o -
zoon unite, t h e p r o c e s s of u n i o n b e i n g called fertilization.
H a v i n g u n i t e d , if t h e y t h e n find t h e m s e l v e s in a p p r o -
p r i a t e e n v i r o n m e n t a l conditions, d e v e l o p m e n t goes o n ;
n e w g e r m cells a n d a s o m a a r e f o r m e d , a n d t h e s a m e
process k e e p s u p g e n e r a t i o n a f t e r g e n e r a t i o n . N o w , while
union of t h e g e r m cells is ; g e n e r a l l y ' a n d i n m o s t o r g a n i s m s
a n essential c o n d i t i o n of t h i s p r o c e s s , i t is also t r u e t h a t
in a few f o r m s of a n i m a l life, m o s t l y f o u n d a m o n g t h e
i n v e r t e b r a t e s , d e v e l o p m e n t of t h e o v u m c a n t a k e place
without a n y p r e c e d i n g f e r t i l i z a t i o n b y a s p e r m a t o z o o n .
T h e p r o c e s s of r e p r o d u c t i o n , i n t h i s case is called par-
thenogenesis. I n a n u m b e r of f o r m s i n w h i c h p a r t h e n o -
genesis n e v e r occurs n o r m a l l y , so f a r a s is k n o w n , i t can
be induced b y a p p r o p r i a t e e x t r a n e o u s p r o c e d u r e s . T h e
discovery of t h i s e x t r a o r d i n a r i l y i n t e r e s t i n g a n d i m p o r -
t a n t fact f o r a n u m b e r of o r g a n i s m s , a n d t h e c a r e f u l
51
52 BIOLOGY OF DEATH
w o r k i n g o u t of i t s physico-chemical basis, we owe t o D r .
J a c q u e s Loeb, of the Rockefeller I n s t i t u t e f o r Medical
R e s e a r c h . Artificial p a r t h e n o g e n e s i s m a y be induced,
as G u y e r , B a t a i l l o n a n d L o e b h a v e Hhown, even in HO
highly organized a c r e a t u r e a s t h e frog, a n d t h e a n i m a l
m a y g r o w to full size. T h e f r o g s s h o w n in F i g u r e 12, while
t h e y p r e s e n t a n a p p e a r a n c e m u c h t h e s a m e m t h a t of
a n y o t h e r f r o g of t h e s a m e species, differ in t h e r a t h e r
f u n d a m e n t a l l y i m p o r t a n t r e s p e c t t h a t t h e y h a d no f a t h e r .
T h e role of a f a t h e r w a s p l a y e d in t h e s e c a s e s by a n
o r d i n a r y d i s s e c t i n g needle. Unfertilized e g g s f r o m a
v i r g i n female w e r e g e n t l y p r i c k e d with a s h a r p l y p o i n t e d
needle. T h i s i n i t i a t i o n of the process* of d e v e l o p m e n t took
place M a r c h 16,3916, in one c a s e , a n d F e b r u a r y 2 7 , 1 9 1 7 ,
in the other. T h e d a t e of d e a t h w a s , an t h e first c a s e , M a y
22,1917, a n d in t h e o t h e r M a r c h 2 4 , 1 9 1 8 .
I n t h e c o u r s e of L o e b ' s s t u d i e s of p a r t h e n o g e n e s i s in
lower m a r i n e i n v e r t e b r a t e s , ha became* i n t e r e s t e d in tho
question of t h e d e a t h of t h e g e r m cells which h a d failed
t o unite, or, h a v i n g united, failed of appropriate* envi-
r o n m e n t a l conditions. H i s r e s e a r c h e s t h r o w l i g h t on som©
of t h e conditions of cellular d e a t h , a n d on t h a t a c c o u n t
t h e y m a y be r e v i e w e d briefly h e r e . H e found t h a t t h e
unfertilized m a t u r e e g g s of t h e s e a - u r c h i n d i e c o m p a r a -
tively soon w h e n d e p o s i t e d in s e a - w a t e r . T h e n a m e e g g s ,
however, live m u c h l o n g e r , a n d will, if a p p r o p r i a t e s u r -
r o u n d i n g conditions a r e p r o v i d e d , g o on a n d d e v e l o p a n
a d u l t o r g a n i s m , if t h e y a r e c a u s e d t o d e v e l o p artificially
by chemical m e a n s o r n a t u r a l l y by f e r t i l i z a t i o n . L o e b
concluded f r o m t h i s t h a t t h e r e a r e t w o p r o c e s s e s g o i n g
on in t h e egg. H e m a i n t a i n e d , on t h e on® h a n d , t h a t t h e r e
a r e specific p r o c e s s e s l e a d i n g to d e a t h a n d d i s i n t e g r a t i o n ;
a n d , on t h e o t h e r h a n d , p r o c e s s e s which l e a d t o eell divi-
C O N D I T I O N S O F C E L L U L A R I M M O R T A L I T Y 53
sion a n d f u r t h e r d e v e l o p m e n t . T h e l a t t e r p r o c e s s e s m a y
be r e g a r d e d a s i n h i b i t i n g o r m o d i f y i n g t h e m o r t a l p r o -
cess. L o e b a n d Lewis 1 u n d e r t o o k e x p e r i m e n t s , b a s e d
u p o n t h i s view, to see w h e t h e r it w o u l d be p o s s i b l e by
chemical t r e a t m e n t of t h e e g g t o p r o l o n g i t s life. Since
i n g e n e r a l specific life p h e n o m e n a a r e p e r h a p s , o n t h e
chemical side, chiefly c a t a l y t i c p h e n o m e n a , i t w a s h e l d
t o be r e a s o n a b l e t h a t if s o m e s u b s t a n c e could b e b r o u g h t
t o a c t on t h e egg, which would i n h i b i t s u c h p h e n o m e n a
w i t h o u t p e r m a n e n t l y a l t e r i n g t h e c o n s t i t u t i o n of t h e
l i v i n g m a t e r i a l , t h e life of t h e cell should be c o n s i d e r a b l y
p r o l o n g e d . T h e first a g e n t chosen f o r t r i a l w a s p o t a s s i u m
cyanide, K C N . I t was known that this substance weakened
o r i n h i b i t e d e n t i r e l y a n u m b e r of e n z y m a t i c p r o c e s s e s i n
living material, without materially or permanently alter-
i n g its s t r u c t u r e -
I t wan f o u n d t h a t , n o r m a l l y , t h e u n f e r t i l i z e d e g g of t h e
s e a - u r c h i n would live i n s e a - w a t e r a t r o o m t e m p e r a t u r e ,
a n d m a i n t a i n itself i n condition f o r successful f e r t i l i z a -
tion a n d d e v e l o p m e n t , u p t o a p e r i o d of a b o u t t w e n t y - t h r e e
h o u r s . A f t e r t h a t t i m e t h e e g g s b e g a n t o weaken.. E i t h e r
t h e y could n o t be successfully fertilized, o r if t h e y w e r e
fertilized, d e v e l o p m e n t only w e n t on f o r a s h o r t t i m e .
A f t e r 32 h o u r s , t h e e g g s could not, a s a r u l e , be fertilized
a t all. T h e e x p e r i m e n t w a s t h e n t r i e d of a d d i n g t o t h e
s e a - w a t e r , i n which t h e u n f e r t i l i z e d e g g s w e r e k e p t ,
gmall a m o u n t s of K C N in a g r a d e d s e r i e s , a n d t h e n e x a m -
i n i n g t h e r e s u l t s of f e r t i l i z a t i o n s u n d e r t a k e n a f t e r a s t a y
of t h e unfertilized e g g s of 75 h o u r s in t h e solution. I t
will b e n o t e d t h a t t h i s p e r i o d of 75 h o u r s is m o r e t h a n
t h r e e t i m e s t h e n o r m a l d u r a t i o n of life of t h e cell i n
n o r m a l sea-water* T h e r e s u l t s of t h i s e x p e r i m e n t a r e
shown i a s u m m a r y f o r m i n T a b l e 4
54 BIOLOGY O F D E A T H
TABLE 4
Experiments of Loch and Lewis on the Prolongation of Life, of the Sm-urch
fyff by KCN
Concentration of Kesuit offrrttlisntittftaft«*r a 7§ hour** iitay
KCN in thfl uAuiitm
Pure sea -water No ogg segments
n/64000 KCN No egg Hcgments
n/16000 KCN No egg m^gutents
n/8000 KCN Very few <*gga nhow a lH*^immig of w*g-
inontiition
n/4000 KCN Vory few <»gg8 «how a U'gitiiting of m»g-
im?ii tut i cm
n/2(K)0 KCN F<»w Kggj* go through th<» <*nrly HtugttH of
T h e r e a p p o a r « in this n h o r t selected t a b l e a d e f e c t
C O N D I T I O N S O F C E L L U L A R I M M O R T A L I T Y 63
w h i c h is even m o r e a p p a r e n t i n h i s l o n g o n e s , n a m e l y ,
t h a t the figures f o r d u r a t i o n of life a r e d i s t i n c t l y r o u n d
n u m b e r s . T h e r e is n o evidence, f o r e x a m p l e , t h a t the
n o r m a l life s p a n of t h e m o u s e is 6 y e a r s . A l l w h o h a v e
statistically studied t h e m a t t e r a g r e e u p o n a m u c h smal-
l e r figure t h a n t h i s . B u t , l e a v i n g t h i s p o i n t a s i d e , i t is
a p p a r e n t t h a t t h e r e is a p a r a l l e l i s m of s t r i k i n g s o r t be-
t w e e n t h e c e p h a l i s a t i o n factor a n d d u r a t i o n of life. I n
o t h e r w o r d s , i t a p p e a r s t h a t the m a n n e r in w h i c h h i g h e r
v e r t e b r a t e s , a t l e a s t , a r e p u t t o g e t h e r in r e s p e c t of the
p r o p o r t i o n a l i t y of b r a i n and body is m a r k e d l y a s s o c i a t e d
w i t h the d u r a t i o n of life. I t w o u l d b e a m a t t e r of g r e a t
i n t e r e s t t o see w h e t h e r t h i s c o r r e l a t i o n b e t w e e n r e l a t i v e
b r a i n - w e i g h t a n d t h e e x p e c t a t i o n of life h o l d s i n t r a -
r a c i a U y as well a s i t does i n t e r - r a c i a l l y . T h e b e a r i n g of
t h e s e r e s u l t s of F r i e d e n t h a l ' s u p o n o u r r e s u l t s as t o t h e
d i s t r i b u t i o n of m o r t a l i t y u p o n a g e r m - l a y e r b a s i s , t o be
d i s c u s s e d i n C h a p t e r V infra, is obvious.
A n o t h e r p o s s i b l e i l l u s t r a t i o n of t h e g e n e r a l p o i n t
n o w u n d e r d i s c u s s i o n m a y be f o u n d i n s o m e r e c e n t w o r k
of R o b e r t s o n a n d R a y . T h e s e a u t h o r s , i n a r e c e n t p a p e r ,
h a v e a n a l y z e d t h e g r o w t h c u r v e s of r e l a t i v e l y long-lived
m i c e as c o m p a r e d w i t h t h e c u r v e s s h o w n by r e l a t i v e l y
short-lived individuals. I n the experiment both groups
w e r e subjected t o t h e s a m e kind of e x p e r i m e n t a l t r e a t -
m e n t of v a r i o u s s o r t s , a n d t h e c a r e w i t h w h i c h t h e e x p e r i -
m e n t s w e r e conducted i n r e s p e c t of c o n t r o l of t h e
environmental factors renders the results highly inter-
e s t i n g a n d v a l u a b l e . T h e long-lived a n i m a l s f o r m a g r o u p
w h i c h g r o w s m o r e r a p i d l y in e a r l y life, a n d at t h e s a m e
t i m e is less v a r i a b l e t h a n t h e s h o r t lived g r o u p . T h e
s h o r t - l i v e d a n i m a l s often g r o w m u c h m o r e r a p i d l y in
l a t e r life t h a n t h e long-lived, b u t this a c c r e t i o n of t i s s u e
70 BIOLOGY OF D E A T H
w a s found t o be r e l a t i v e l y u n s t a b l e . T h e y f u r t h e r found
t h a t t h e long-lived a n i m a l s r e p r e s e n t a r e l a t i v e l y s t a b l e
g r o u p , highly r e s i s t a n t to e x t e r n a l d i s t u r b i n g f a c t o r s ,
a n d s h o w i n g a m o r e o r less m a r k e d b u t n o t i n v a r i a b l e
t e n d e n c y to e a r l y o v e r g r o w t h a n d r e l a t i v e p a u c i t y of
t i s s u e accretion in l a t e life. T h e s h o r t - l i v e d a n i m a l s a r e
on t h e c o n t r a r y r e l a t i v e l y u n s t a b l e , sensitive to e x t e r n a l
d i s t u r b i n g f a c t o r s , and, as a r u l e , b u t not i n v a r i a b l y , d i s -
p l a y r e l a t i v e l y deiicient e a r l y g r o w t h a n d a t e n d e n c y t o
r a p i d a c c r e t i o n of t i s s u e in l a t e r life.
I n i n t e r p r e t i n g these r e s u l t s , R o b e r t s o n a n d K a y be-
lieve t h a t t h e differences a r e based upon t h e fact t h a t
in e a r l y or e m b r y o n i c life the o u t s t a n d i n g c h a r a c t e r i s t i c
of t h e t i s s u e s is a high p r o p o r t i o n of c e l l u l a r e l e m e n t s ,
w h e r e a s i n old a g e t h e r e in a m a r k e d i n c r e a s e in connective
tissues. They further point out t h a t connective tissue
e l e m e n t s a r e u l t i m a t e l y d e p e n d e n t u p o n c e l l u l a r tisBUtts
for t h e i r s u p p o r t , a n d t h a t t h e c o n n e c t i v e t i s s u e s a r e
expensive to m a i n t a i n . T h e y believe t h a t t h e r e a s o n t h a t
the s u b s t a n c e t e t h e l i n ( r / . C h a p . V J I infra) p r o l o n g s life
is because i t a c c e l e r a t e s t h e m e t a b o l i s m of t h e c e l l u l a r
elements to the d e t r i m e n t of t h e c o n n e c t i v e t i s s u e ele-
m e n t s . L o n g e v i t y on thin view in d e t e r m i n e d n o t by t h e
absolute m a s s of living s u b s t a n c e , b u t by t h e r e l a t i v e
proportioiiH of parenchymatotiH to s c l e r o u s t i s s u e s .
HKNRSCRNCK
T h e f a c t s p r e s e n t e d in t h i s and t h e p r e c e d i n g c h a p t e r
clearly m a k e i t n e c e s s a r y t o review w i t h s o m e c a r e t h e
c u r r e n t conception of senescence. Senescence, o r g r o w -
ing old, is commonly c o n s i d e r e d to be t h e n e c e s s a r y p r e l -
u d e to " n a t u r a l / 1 a s d i s t i n g u i s h e d f r o m accidental d e a t h .
3 N D I T I 0 N S O F C E L L U L A R I M M O R T A L I T Y 71
s t h e evidence r e a l l y sound a n d complete t h a t such
> fact?
c a r e f u l a n d u n p r e j u d i c e d e x a m i n a t i o n will inevi-
s u g g e s t to the open mind, I think, t h a t m u c h of the
i n g l i t e r a t u r e on senescence is r e a l l y of n o f u n d a -
a l importance, because it has unwittingly reversed
r u e s e q u e n t i a l o r d e r of t h e c a u s a l n e x u s . If cells
a r l y e v e r y s o r t a r e capable, u n d e r a p p r o p r i a t e con-
.is, of l i v i n g indefinitely in u n d i m i n i s h e d v i g o r , a n d
ogical normality, there is little ground for posta-
ge t h a t t h e o b s e r v e d senescent c h a n g e s i n t h e s e cells
> i n t h e body, s u c h a s t h o s e d e s c r i b e d b y M i n o t a n d
•s, a r e e x p r e s s i v e of specific a n d i n h e r e n t m o r t a l
>sses g o i n g on i n t h e c e l l s ; o r t h a t t h e s e cellular p r o -
s a r e the c a u s e of senescence, as M i n o t h a s concluded.
h a t t h e r e is such a p h e n o m e n o n a s senescence i s , of
5e, c e r t a i n . Tt is o b s e r v a b l e b o t h i n Protozoa and
"etazoa. T h e r e a l question, h o w e v e r , i s a twofold
viz: ( a ) i8 senescence in e i t h e r P r o t o z o a o r M e t a z o a
',evitable consequence of the s t r a i n o r t h e i n d i v i d u a l
:xg l i v e d ; a n d ( b ) is senescence a necessary asso-
a n d f o r e r u n n e r of n a t u r a l d e a t h ?
e t u s briefly r e c o n s i d e r t h e f a c t s . I n P r o t o z o a a
i n g down of t h e division r a t e i n c u l t u r e h a s been
a e n t l y o b s e r v e d ; a n d i t h a s been held, first, t h a t
i s a p h e n o m e n o n e s s e n t i a l l y h o m o l o g o u s t o aenes-
> in the m e t a z o a n ; a n d second, t h a t if n u c l e a r
g a n i z a t i o n , b y t h e w a y e i t h e r of e n d o m i x i s o r of
a g a t i o n , d i d n o t occur t h a t t h e s t r a i n w o u l d die out.
e d f JVmningH, in d i s c u s s i n g t h e m a t t e r i n his l a s t
says:
Thus it appears that in theae organisms nature has employed the
d of keeping on hand a reserve utock of a material essential to
72 BIOLOGY O F D E A T H
life; by replacing at interval** the; worn out material with thin re*erve,
the animalH are kept in a state of perpetual vigor; not, a« individual*,
growing old or dying a natural death. Xevcrthelew*, a wear ing out pro-
CC8H, such HH might be called getting old, cloen occur in the titrttrtttrea
employed in the active funetioiiH of life, and thene munt W replaced after
a time of nerviee. Ho fur HH the cronditionM in thene orgariinmH are typical,
deterioration and death do appear to N* a conH«wjiiiMUfi? c»f full and active
life; life carries within itadf the wvd* of death. It in not muting with
another individual that avoidu thin end; hut replacement of the. worn
material by a renerve The grt»at mnm of eelln mihjrct to death in the
higher animalM dwindlen in the infuHorian to the maeronueletm; thin alone
reprenentw a eorpno. But the diMMohition of thin tutrpMt? omirM within
the living body. It much re«emblim wieh ft pror***« &*i the wanting liwity
and dcHtruetion of minute parts of our own Ijodiett, whii'h we know fa
taking place at all Union, and which dmm not interrupt the life of
the individual."
These r e s u l t s s u g g e s t t h a t t h e r e is p r o d u c e d in some
cases by t h e b o d y o r s o m e of i t s p a r t s , a s u b s t a n c e
which inhibits t h e p o w e r of cells t o m u l t i p l y o r t o r e m a i n
alive. H o w g e n e r a l such a p h e n o m e n o n is i n occurrence
does n o t y e t a p p e a r , but, a p p a r e n t l y , i t m u s t be a b s e n t
in the c a s e of clonal r e p r o d u c t i o n i n p l a n t s a l r e a d y dis-
cussed, a n d i n t h e a n a l o g o u s c a s e of a g a m i c r e p r o d u c t i o n
in lower Metazoa, (cf. p l a n a r i a n s ) . I t seems possible
t h a t the r e s u l t s of C a r r e l a n d E b e l i n g m i g h t be o p e n to
a slightly different i n t e r p r e t a t i o n t h a n t h a t which t h e y
give, which h y p o t h e c a t e s a specific i n h i b i t i n g substance
in t h e s e r u m , i n c r e a s i n g i n e i t h e r a m o u n t o r specific
potency w i t h age. I t seems t o m e t h a t all of t h e i r facts
could be i n t e r p r e t e d w i t h equal cogency on t h e supposi-
tion t h a t t h e s e r u m f r o m a n old a n i m a l i s itself senes-
78 BIOLOGY OF DEATH
cent a s a w h o l e ; t h a t is, h a s u n d e r g o n e a phynieo-chemi
cal a l t e r a t i o n ( a s c o m p a r e d with t h a t of a y o u n g ani
m a l ) , which m c o m p a r a b l e t o tho m o r p h o l o g i c a l am
physiological c h a n g e s which a r e o b s e r v a b l e in seneseen
cells. I t m a y f u r t h e r q u i t e r e a s o n a b l y he? s u p p o s e d thn
" s e n e s c e n t " s c r u m , because of t h e s e physieo»chemiea
a l t e r a t i o n s , does n o t f u r n i s h BO f a v o r a b l e a n u t r i e n t me
dium f o r in vitro c u l t u r e s a s d o e s " y o u n g " s e r u m . Sue]
a view a v o i d s t h e necessity of p o s t u l a t i n g a apecifi
" s e n e s c e n t " s u b s t a n c e , t h e existence of which would b
exceedingly difficult to p r o v e .
B u t in a n y case, w h a t e v e r e x p l a n a t i o n is HUggentei
for C a r r e l a n d KheHng'a b r i l l i a n t nmultft, it d o e s no
seem to m e t h a t t h e result** t h e m s e l v e s , which alone a r
the realities p e r t i n e n t in t h e p r e m i s e s , e i t h e r offer an;
obstacle to or, indeed, a l t e r t h e i n t e r p r e t a t i o n of mme$,
cence which I h a v e s u g g e s t e d above- F o r , w h a t the* re
s u i t s r e a l l y d e m o n s t r a t e is, essentially, that the* s e r u m o
old a n i m a l s is a less f a v o r a b l e c o m p o n e n t of tlm mitricni
m e d i u m of cells in vitro t h a n in t h e s e r u m of y o u n g utii
m a l s . T h i s fact in a c o n t r i b u t i o n t o o u r k n o w l e d g e a
t h e p h e n o m e n a a n d a t t r i b u t e s of HenftHccmee of ftrat-clafi
i m p o r t a n c e ; b u t it does n o t per mf an it uppcmrH t o mi
p e r m i t of a n y nmf g e n e r a l i z a t i o n a s t o t h e etiology a
senescence.
CHAPTER III
T H E CHANCES OF D E A T H
U P to t h i s p o i n t in o u r discussion of d e a t h a n d lon-
g e v i t y we h a v e , f o r t h e m o s t p a r t , d e a l t w i t h g e n e r a l a n d
qualitative matters, and have not made any particular
e x a m i n a t i o n a s t o t h e q u a n t i t a t i v e a s p e c t s of t h e p r o b -
l e m of longevity. T o this p h a s e a t t e n t i o n m a y n o w be
d i r e c t e d . F o r one o r g a n i s m , a n d one o r g a n i s m only, do
we k n o w m u c h a b o u t t h e q u a n t i t a t i v e a s p e c t s of l o n g e v i t y .
I r e f e r , of c o u r s e , to m a n , a n d the a b u n d a n t r e c o r d s which
exist a s t o t h e d u r a t i o n of his life u n d e r v a r i o u s condi-
tions a n d c i r c u m s t a n c e s . I n 1532 t h e r e b e g a n i n L o n d o n
t h e first definitely k n o w n compilation of weekly " Bills
of M o r t a l i t y . " S e v e n y e a r s l a t e r , t h e official r e g i s t r a -
tion of b a p t i s m s , m a r r i a g e s a n d d e a t h s w a s b e g u n in
F r a n c e , a n d s h o r t l y a f t e r t h e o p e n i n g of t h e s e v e n t e e n t h
century similar registration was begun in Sweden. In
1662 w a s p u b l i s h e d t h e first edition of a r e m a r k a b l e book,
a book which m a r k s t h e b e g i n n i n g of t h e s u b j e c t w h i c h w e
n o w k n o w a s l c vital s t a t i s t i c s . ' ' I r e f e r t o ' ' N a t u r a l a n d
P o l i t i c a l O b s e r v a t i o n s Mentioned in t h e F o l l o w i n g I n d e x ,
a n d m a d e u p o n t h e Bills of M o r t a l i t y " b y C a p t a i n J o h n
G r a u n t , Citizen of L o n d o n . F r o m t h a t d a y to t h i s , in
a n e v e r w i d e n i n g p o r t i o n of t h e i n h a b i t e d globe w e h a v e
h a d m o r e o r less c o n t i n u o u s p u b l i s h e d r e c o r d s a b o u t t h e
d u r a t i o n of life of m a n . T h e a m o u n t of s u c h m a t e r i a l
which h a s a c c u m u l a t e d is e n o r m o u s . W e a r e only a t t h e
79
80 BIOLOGY OF D E A T H
b e g i n n i n g , h o w e v e r , of its p r o p e r m a t h e m a t i c a l a n d bio-
logical a n a l y s i s . If biologists h a d been f u r n i s h e d with
d a t a of a n y t h i n g like t h e s a m e q u a n t i t y a n d q u a l i t y for
a n y o t h e r o r g a n i s m t h a n m a n it is probable* t h a t a vastly
g r e a t e r a m o u n t of a t t e n t i o n would have been devoted to
t h e m t h a n e v e r h a s been g i v e n t o vital s t a t i s t i c s , no-called,
and t h e r e would h a v e been a s a r e s u l t m a n y fundamental
a d v a n c e s in biological knowledge now lacking, because
m a t e r i a l of t h i s s o r t BO g e n e r a l l y Hc*c*ms t o tlm profes-
sional biologist to be s o m e t h i n g nlxmt which he in in nc
w a y concerned.
L e t u s e x a m i n e some of t h e g e n e r a l facts a b o u t thi
n o r m a l d u r a t i o n of life in m a n . W e m a y p u t t h e m a t t e i
in t h i s w a y : S u p p o s e we s t a r t e d o u t a t a given i n s t a n t oi
time w i t h a h u n d r e d t h o u s a n d i n f a n t s , e q u a l l y distributee
a s to sex, a n d all b o r n a t t h e s a m e i n s t a n t of time- Hov
m a n y of t h e s e i n d i v i d u a l s would die in e a c h Hiicceedin|
y e a r , a n d w h a t would be t h e g e n e r a l picture* of tlm change)
in t h i s c o h o r t with t h e p a s s a g e of thru* 1 T h e facts on thii
p o i n t for t h e R e g i s t r a t i o n A r e a of the U n i t e d S t a t e s ii
1910 a r e exhibited in F i g u r e 18, which is b a s e d oi
G l o v e r ' s U n i t e d S t a t e s Life T a b l e s .
I n this t a b l e a r e seen t w o c u r v e d linmf arm m a r k e d I
a n d the o t h e r ds. T h e lz line iitflieatos flu* n u m b e r o
i n d i v i d u a l s , o u t of t h e o r i g i n a l 100,000 Mturtitig togiHLhe:
a t b i r t h , w h o s u r v i v e d a t t h e b e g i n n i n g of <*ac?li y o u r o
t h e life s p a n , i n d i c a t e d a l o n g t h e b o t t o m of t h e diugrani
T h e dm line s h o w s t h e n u m b e r d y i n g within each y e a
of t h e life s p a n . I n o t h e r w o r d s , if we s u b t r a c t the* nuno
b e r d y i n g w i t h i n e a c h y e a r f r o m t h e n u m b o r mirvivini
a t t h e b e g i n n i n g of t h a t y e a r we shall gc?t the* s e r i e s c?
figures p l o t t e d a s t h e lx line. W e note* t h a t in ilia ver;
first y e a r of life t h e o r i g i n a l h u n d r e d t h o u s a n d lose ove
T H E CHANCES OF DEATH 81
one-tenth of t h e i r n u m b e r , t h e r e b e i n g only 88,538 s u r -
viving a t t h e b e g i n n i n g of t h e second y e a r of life. I n
the n e x t y e a r 2,446 d r o p out, a n d i n t h e y e a r following
t h a t 1,062. T h e n t h e line of s u r v i v o r s d r o p s off m o r e
slowly between t h e p e r i o d of y o u t h a n d e a r l y a d u l t life.
A t 40 y e a r s of age, a l m o s t e x a c t l y 30,000 of t h e o r i g i n a l
100,000 h a v e p a s s e d a w a y , a n d f r o m t h a t p o i n t on t h e I z
line descends w i t h e v e r i n c r e a s i n g r a p i d i t y , u n t i l a b o u t
JN/TCD STATCS
UFC TABU
- mo
——.
"^
^
\
\
\
\
• — mi 3—« — s r—;ar-m
FIG. 18.—Life table diagram. For explanation see text.
a g e 80, w h e n i t once m o r e b e g i n s t o d r o p m o r e slowly,
a n d the l a s t f e w s u r v i v o r s p a s s o u t g r a d u a l l y , a few each
y e a r u n t i l s o m e t h i n g over t h e c e n t u r y m a r k is r e a c h e d ,
when t h e l a s t o n e of t h e 100,000 w h o s t a r t e d a c r o s s t h e
b r i d g e of life t o g e t h e r will h a v e e n d e d h i s j o u r n e y .
This d i a g r a m is a g r a p h i c r e p r e s e n t a t i o n of t h a t im-
p o r t a n t t y p e of d o c u m e n t k n o w n a s a life o r m o r t a l i t y
table. I t p u t s t h e f a c t s of m o r t a l i t y a n d l o n g e v i t y in t h e i r
best f o r m f o r c o m p a r a t i v e p u r p o s e s . T h e first such
table a c t u a l l y t o be c o m p u t e d i n a n y t h i n g like t h e m o d e r n
fashion w a s m a d e b y t h e a s t r o n o m e r , D r . E . H a l l e y , a n d
6
82 BIOLOGY O F D E A T H
w a s p u b l i s h e d in 1693, a l t h o u g h t h i r t y y e a r n before t h a t
time P a s c a l a n d F o r m a t (<:/. L e v a s n e u r ) h a d laid down
c e r t a i n m a t h e m a t i c a l rules for t h e c a l c u l a t i o n of the
p r o b a b i l i t i e s of h u m a n life. S i n c e H a l l e y ' s t i m e a g r e a t
n u m b e r of such t a b l e s h a v e been c a l c u l a t e d . D a w s o n
fills a s t o u t o c t a v o v o l u m e w i t h a collection of t h e m o r e
i m p o r t a n t of s u c h t a b l e s , c o m p u t e d f o r different coun-
t r i e s a n d different g r o u p s of t h e p o p u l a t i o n . Now they
h a v e become such a c o m m o n p l a c e t h a t e l e m e n t a r y classes
i n vital s t a t i s t i c s a r e r e q u i r e d to c o m p u t e t h e m (see for
e x a m p l e D u b l i n ' s N e w H a v e n life t a b l e ) .
n, T a b l e fi in i n s e r t e d , Kivinn; t h e f*xpt*rtntit»n# of
life from wltleli e^rfnin of the diiifframMi h a v e hmn%
F i g u r e 19 given t h e y*Mi!l8 from H n l l ^ f 8 lnl>lf%
upon t h e m o r t a l i t y e x p e r i e n c a in the inty nf Ifr**dtitt» in
Silesia, d u r i n g t h e yearn 1687 to 1691. Thin tjivw tin
a r o u g h , b u t in Itn g e n e r a l iwc»r»p nttfitc«ti?iitly uemtrnU*
p i c t u r e of tho forces of m o r t a l i t y t o w a r d * thit mul nf th«*
Bovcnteonth c e n t u r y From thin diagriiin i t uppcitri* thitt
a t b i r t h t h e e x p e c t a t i o n of lift of nn i n d i v i d u a l hunt in
B r e s l a u in the s e v e n t e e n t h c e n t u r y wan m u c h l«w«*r t h a n
T H E CHANCES OF D E A T H 85
t h a t of a n i n d i v i d u a l b o r n i n t h e U n i t e d S t a t e s i n 1910.
T h e difference a m o u n t s to a p p r o x i m a t e l y 18 y e a r s !
P r o b a b l y t h e a c t u a l difference w a s n o t so g r e a t a s this,
a s these e a r l y life t a b l e s a r e k n o w n t o be i n a c c u r a t e a t
the ends of t h e lifespan, p a r t i c u l a r l y a t t h e b e g i n n i n g .
A t 10 y e a r s of age, t h e difference i n e x p e c t a t i o n of life
h a d been r e d u c e d t o j u s t o v e r 12 y e a r s ; a t a g e 20, t o a
little less t h a n 10 y e a r s ; a t a g e 30 t o 7 - % y e a r s ; a t a g e
50 t o j u s t over 4 y e a r s ; a t a g e 70 t o 1 - % y e a r s . A t
a g e 80 t h e lines h a v e c r o s s e d , b u t o w i n g t o t h e i n a d e -
quate methods; of g r a d u a t i o n u s e d b y t h i s p i o n e e r a c t u a r y ,
together with the paucity and p r o b a b l y somewhat inac-
c u r a t e c h a r a c t e r of h i s m a t e r i a l , no ; s t r e s s is t o be l a i d
u p o n the crossing of t h e lines, o r u p o n the* s u p e r i o r
expectation of life a t t h e high, a g e s i n t h e s e v e n t e e n t h
c e n t u r y m a t e r i a l . W h a t t h e d i a g r a m shows is t h a t t h e
expectation of life a t e a r l y a g e s w a s v a s t l y i n f e r i o r
in t h e seventeenth c e n t u r y t o w h a t i t is now, w h i l e a t
advanced ages t h e chances of l i v i n g w e r e s u b s t a n t i a l l y
t h e same. L e t u s d e f e r t h e f u r t h e r d i s c u s s i o n of the
m e a n i n g a n d e x p l a n a t i o n of this c u r i o u s f a c t u n t i l we
h a v e examined some f u r t h e r d a t a .
F i g u r e 20 c o m p a r e s the e x p e c t a t i o n of life i n E n g l a n d
a t t h e middle of t h e e i g h t e e n t h c e n t u r y , o r a b o u t a cen-
t u r y l a t e r t h a n t h e l a s t , w i t h p r e s e n t c o n d i t i o n s in t h e
U n i t e d S t a t e s . A g a i n w e see t h a t t h e e x p e c t a t i o n a t
birth was greatly inferior then to w h a t it is now, b u t the
difference is n o t so g r e a t a s i t w a s a c e n t u r y e a r l i e r ,
a m o u n t i n g t o b u t 1 2 - % y e a r s i n s t e a d of t h e 18 w e f o u n d
before. F u r t h e r i t i s seen t h a t , j u s t a s b e f o r e , t h e expec-
t a t i o n s come closer t o g e t h e r w i t h a d v a n c i n g a g e . B y
the time a g e 45—middle life—is r e a d i e d t h e e x p e c t a t i o n
of life w a s s u b s t a n t i a l l y t h e s a m e i n t h e e i g h t e e n t h cen-
86 BIOLOGY O F D E A T H
S5
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6 £0 s
/S
10 si
5
0 =
YEAR'S OF LIFE
Fia. 20—Comparing the expectation of life in the 18th century with that of the presen
I t should be noted t h a t all d a t a as t o m o r t a l i t y in
s e v e n t e e n t h a n d eighteenth centuries lack the degree
a c c u r a c y which one desires for p u r e l y scientific p u r p o
B y e r r i n g g e n e r a l l y on the safe side t h e s e old morta
t a b l e s did well enough f o r i n s u r a n c e p u r p o s e s . B u t q
different r e s u l t s a s t o t h e detailed values of life t
c o n s t a n t s i n t h e s e e a r l y p e r i o d s a r e to be found in
l i t e r a t u r e . F o r example, R i c h a r d s constructed s
life tables f r o m N e w E n g l a n d genealogical r e c o r d s ,
c o m p a r e d t h e m w i t h W i g g l e s w o r t h ' s table, a n d also ^
tho^e of m o d e r n t i m e s . H i s g e n e r a l conclusion, for
T H E CHANCES OF D E A T H 87
N e w E n g l a n d p o p u l a t i o n , i s : " t h a t d u r i n g t h e l a s t half-
c e n t u r y longevity* i n M a s s a c h u s e t t s , a n d p r o b a b l y i n
New E n g l a n d , h a s i n c r e a s e d , t h a t f r o m 1793 t o 1850 t h e
increase is less c e r t a i n a n d f r o m t h e s e v e n t e e n t h to t h e
eighteenth c e n t u r y w h a t d a t a we h a v e p o i n t r a t h e r to
a decrease t h a n to a n y t h i n g e l s e . " T h i s r e s u l t m a y
m e a n a n y one of a n u m b e r of t h i n g s . I t m a y m e a n m e r e l y
i n a d e q u a t e a n d i n a c c u r a t e d a t a on w h i c h the s e v e n t e e n t h
c e n t u r y tables w e r e calculated. I t m a y m e a n a r e s u l t of
less s t r i n g e n t selection i n t h e m a k e u p of t h e p o p u l a t i o n
with t h e p a s s a g e of t i m e . I n a n y case i t a p p l i e s only to
a small a n d r a t h e r h o m o g e n e o u s g r o u p of p e o p l e .
T h e c h a n g e s i n e x p e c t a t i o n of life f r o m t h e m i d d l e
of t h e s e v e n t e e n t h c e n t u r y t o t h e p r e s e n t t i m e w h e r e t h e
r e c o r d s a r e m o s t extensive a n d reliable a p p e a r t o fur-
nish a r e c o r d of a r e a l e v o l u t i o n a r y p r o g r e s s i o n . I n this
respect a t l e a s t m a n h a s definitely a n d distinctively
changed, a s a r a c e , i n a p e r i o d of t h r e e a n d a half cen-
t u r i e s . T h i s i s , of course, a m a t t e r of e x t r a o r d i n a r y
interest, a n d a t once s t i m u l a t e s t h e d e s i r e t o go still
f a r t h e r back i n h i s t o r y a n d see w h a t t h e e x p e c t a t i o n of
life t h e n w a s . F o r t u n a t e l y , t h r o u g h t h e l a b o r s of K a r l
P e a r s o n , a n d his associate, W . R. Macdonell, i t i s p o s -
sible t o d o t h i s , if n o t w i t h p r e c i s e a c c u r a c y , a t l e a s t
to a r o u g h first a p p r o x i m a t i o n . P e a r s o n h a s a n a l y z e d
t h e r e c o r d s a s to a g e a t d e a t h w h i c h w e r e f o u n d
u p o n m u m m y cases s t u d i e d b y P r o f e s s o r W . S p i e g e l b e r g .
T h e s e m u m m i e s belonged t o a p e r i o d b e t w e e n 1,900
a n d 2,000 y e a r s ago, w h e n E g y p t w a s u n d e r R o m a n
dominion. T h e d a t a w e r e e x t r e m e l y m e a g r e , b u t f r o m
P e a r s o n ' s ! a n a l y s i s of t h e m i t h a s b e e n possible t o
* Richards somewhat loosely uses this term when he means "expectation
of life."
88 BIOLOGY OF DEATH
construct the diagram which in nhown in Figure 21.
Each circle marks a point whore it wan ponnible definitely
to calculate an expectation of life. The curve* running
through the circles is a rough graphic smoothing of the
scattered obfierved data. Unfortunately, there were no
records of deaths in early infancy. Hither there were
no baby mummies, or if there were they have disappeared
p ; » ; « •• . » » « ,. , ..
I -• ' r . ;* * .- :» ., « . » i ;* » \ ......, • -1
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F i o . 2 3 — C o m p a r i n g t h e e x p e c t a t i o n of life of t h e p o p u l a t i o n of t h e R o m a s provinces
Hispania and L u s i t a n i a w i t h t h a t of present d a y Americana. P l o t t e d from M a c d o n e l l ' s a n d
Glover's data.
t h a n that of t h e m a l e s w h i c h e x a c t l y r e v e r s e s t h e m o d e r n
a d v e r s e l y a f f e c t i n g t h e h e a l t h of f e m a l e s i n t h e E o m a n
E m p i r e , w h i c h n o l o n g e r o p e r a t e i n t h e m o d e r n w o r l d . U p
t o s o m e t h i n g like a g e 25 t h e e x p e c t a t i o n of life of d w e l l e r s
i n t h e city of E o m e w a s e x t r e m e l y b a d , w o r s e t h a n i n t h e
E o m a n o - E g y p t i a n p o p u l a t i o n w h i c h P e a r s o n studied, o r
i n the p o p u l a t i o n s of o t h e r p a r t s of t h e E o m a n E m p i r e a s
w e shall see i n t h e f o l l o w i n g d i a g r a m . M a c d o n e l l t h i n k s
92 BIOLOGY O F D E A T H
t h a t t h i s difference is r e a l a n d d u e t o circumstances pecu-
liar to Eome.
T h e g e n e r a l f e a t u r e s of t h e d i a g r a m f o r t h e popu-
l a t i o n of H i s p a n i a a n d L u s i t a n i a ( F i g u r e 23) a r e similar
to t h o s e t h a t w e h a v e seen, w i t h t h e difference t h a t t h e
expectation of life u p to a g e 20 o r 25 is n o t a s b a d a s i n
t h e city of E o m e itself. A g a i n t h e females show a lower
expectation p r a c t i c a l l y t h r o u g h o u t life t h a n do t h e males.
55 f—%«
UNITED STATES
SO
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10
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0
0 5 10 15 20 25 30 35 4-0 45 50 6"S 60 65 70 75 BO tiii 90 95 10
YEARS OF AGE
F I G . 2 4 — C o m p a r i n g t h e e x p e c t a t i o n of life of t h e p o p u l a t i o n of t h e R o m a n p r o v i n c e s in
Africa w i t h t h a t of p r e s e n t d a y A m e r i c a n s . P l o t t e d f r o m M a c d o n e l l ' s a n d G l o v e r ' s d a t a .
T h e lines c r o s s t h e m o d e r n A m e r i c a n lines a t a b o u t a g e
60 a n d f r o m t h a t p o i n t o n t h e s e colonial E o m a n s a p p a r -
ently h a d a b e t t e r e x p e c t a t i o n o f life t h a n t h e m o d e r n
A m e r i c a n h a s .
T h e E o m a n o - A f r i c a n p o p u l a t i o n d i a g r a m a p p e a r s t o
start a t n e a r l y t h e s a m e p o i n t a t b i r t h a s d o e s t h e m o d e r n
A m e r i c a n , a n d i n g e n e r a l t h e differences u p t o a g e 35
a r e n o t substantially m o r e m a r k e d f r o m m o d e r n condi-
tions t h a n t h e y a r e i n t h e s e v e n t e e n t h c e n t u r y B r e s l a u
b a s e d o n t h e e x p e r i e n c e of 1 8 7 1 to 1 8 8 0 i n E n g l a n d . T h i s
line g i v e s t h e d e a t h s p e r a n n u m of o n e t h o u s a n d p e r s o n s
b o r n i n the s a m e y e a r . T h e first c o m p o n e n t w h i c h h e s e p a -
r a t e d w a s t h e old a g e m o r t a l i t y . T h i s is s h o w n b y t h e
d o t t e d c u r v e h a v i n g its m o d a l p o i n t b e t w e e n 7 0 a n d 75
y e a r s , a t the p o i n t l e t t e r e d Ox o n t h e b a s e of t h e d i a g r a m .
PLARSQN'S GRADUATION OF
... —
..... :
— —— — — —— - — -•
—
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i
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o
or we
Fio. 25.—Showing Pearson'a resultn in fitting the dx lin© of the life table with fi skew
freauoncy curves. Plotted from the data of Pearson's original memoir on "Skew Variation"
in Phil. Trans. Roy. Soo.
— (.09092 (s —22.5)]'
4- 2.6 e + 8.5 I x—2 J — .3271 (x — 3)
I. general diseases.
II. Diseases of the nervous system and of the organs of special sense.
III. Diseases of the circulatory system.
IV. Diseases of the respiratory system.
V. Diseases of the digestive system.
VI. Non-venereal diseases of the gen i to-urinary system and annexa.
VII. The puerperal state.
VIII. Diseases of the skin and of the cellular tissue.
IX. Diseases of the bones and organs of locomotion.
X. Malformation.
XI. Early infancy.
XII. Old age.
XIIL External causes.
XIV. Ill-defined diseases.
T h e u n d e r l y i n g i d e a of t h i s r e a r r a n g e m e n t of t h e
c a u s e s of d e a t h i s t o p u t all tho,se l e t h a l e n t i t i e s t o g e t h e r
which b r i n g a b o u t d e a t h because of t h e f u n c t i o n a l o r g a n i c
b r e a k d o w n of t h e s a m e g e n e r a l o r g a n s y s t e m . T h e cause
of t h i s f u n c t i o n a l b r e a k d o w n m a y b e a n y t h i n g w h a t e v e r
i n t h e r a n g e of p a t h o l o g y . I t m a y b e d u e to b a c t e r i a l
infection; it m a y be due tq trophic disturbances; it may
b e d u e to| m e c h a n i c a l d i s t u r b a n c e s w h i c h p r e v e n t t h e
c o n t i n u a t i o n of n o r m a l f u n c t i o n ; o r t o a n y c a u s e w h a t -
soever. I n o t h e r w o r d s t h e b a s i s of t h e classification is
not t h a t of p a t h o l o g i c a l c a u s a t i o n , b u t i t i s r a t h e r t h a t
of o r g a n o l o g i c a l b r e a k d o w n . W e a r e n o w looking a t
t h e q u e s t i o n of d e a t h f r o m t h e s t a n d p o i n t of t h e biologist,
w h o c o n c e r n s himself n o t w i t h w h a t c a u s e s a c e s s a t i o n of
function, b u t r a t h e r w i t h w h a t p a r t of t h e o r g a n i s m ceases
t o function, a n d t h e r e f o r e c a u s e s d e a t h .
I n a s e r i e s of p a p e r s a l r e a d y p u b l i s h e d I h a v e given
a d e t a i l e d a c c o u n t of this classification, a n d t h e r e a s o n i n g
on w h i c h p a r t i c u l a r c a u s e s of d e a t h a r e p l a c e d i n i t w h e r e
106 BIOLOGY OF D E A T H
they are. Space is lacking here to go into the details,
and I must consequently ask the reader either to take it
on faith for the time being that the classification is at
least a fairly reasonable one, or to take the trouble to
go over it in detail in the original publication.*
ALIMENTARY
TRACT AND
ASSOCIATED
ORGANS
CIRCULATORYX 203.3
SYSTEM. 208. C
BLOOD 2M&
NERVOUS
SYSTEM /fW
SENSE
ORGANS
KIDNEYS AND
RELATED
EXCRETORY
ORGANS
PRIMARY AND
SECONDARY
SEX ORGANS
SKELETAL
MUSCULAR
SYSTEM
SKIN
ENDOCRINAL
SYSTEM
VS. REG AREA IS06-/O ENGLAND A*O WALES J&& SAO PS4UL0 f&(7
FIG. 36.—Showing th« relative importance of the different orgns nytttma \n Utittmu tumtmUty,
T H E CAUSES OF DEATH 109
IU«i«tr«li.in Arm,
Pmth** V. H. A.
No. taf ft licit lmi
1017
...mow
In
on a n o r K a n o l o g i c a l htmh* A ckmth r o a l l y d u o t o o l d
a g e , i n t h e sumMO o f Mi*ichntkoflF y rf»prcmc*ntH, f r o m t h e
p o i n t of v i o w o f the? premfnt riimMiHHion, a brt*nkiiig d o w n
o r w e a r i n g o u t o f a l l t h e o r g a n m*Ktmm o f t h o b o d y c o n -
temporancotiMly. I n a s t r i c t ncnne thin p r o l m W y niivcir,
o r a t bo»t e x t r e m e l y r i i r e l y , hnpiKmn. B u t phyHtcians
a n d reKifltrarH o f m o r t a l i t y ntill r e t u r n a c e r t a i n n u m b e r
of d e a t h s an clue t o " f l e n i l i t y . 1 f U n d e r the* c i r e t i m i t a n c c i i
110 BIOLOGY OF D E A T H
it is not possible to go behind such returns biologically.
The second line of Table 8, "Ill-defined d i s e a s e s , "
furnishes a striking commentary on the relative efficiency
of the medical profession in the United States and Eng-
land in respect of the reporting of the causes of death.
Only about one-fourth as many deaths appear in the
English vital statistics as due to ill-defined and unknown
causes as in the United States figures.
Returning now to the consideration of the general
results set forth in Table 7 and F i g u r e 26, a number of
interesting points about human mortality are apparent.
I n the United States, during the decade covered, more
deaths resulted from the breakdown of the respiratory
system than from the failure of any other organ system
of the body. The same thing i s true of England and
Wales. I n Sao P a u l o the alimentary tract takes first
position, with the respiratory system a rather close
second. The tremendous death rate i n S a o P a u l o charge-
able to, the alimentary tract is chiefly due to the relatively
enormous number of deaths of infants under two from
diarrhoea and enteritis. Nothing approaching such a
rate for this category as S a o Paulo shows is known in
this country or England.
In all three localities studied the respiratory and the
alimentary tract together account for rather more than
half of all the deaths biologically classifiable. These are
the two organ systems which, while physically internal,
come i n contact directly at their surfaces with environ-
mental entities (water, food, air) with all their bacterial
contamination. The only other organ system directly
exposed to the environment i s the skin. The alimentary
canal and the lungs are, of course, i n effect invaginated
surfaces of the body. The mucous membranes which
line them are far less resistant t o environmental stresses,
T H E CAUSES OF D E A T H 111
both physical and chemical, than is the skin w i t h its pro-
tecting layers of stratified and cornified epithelium.
The organs concerned with the blood and its circula-
tion—the heart, arteries and veins, etc.—stand third in
importance in the mortality list. Biologically the blood,
through its immunological mechanism, constitutes the
second line of defense which the body has against noxious
invaders. The first line is the resistance of the outer
cells of the skin and the lining epithelium of alimentary
tract, lungs, and sexual and excretory organs. W h e n
invading organisms p a s s or break down these first t w o
lines of defense, the battle* is then with the home guard, the
cells of the organ system itself, which, like the industrial
workers of a commonwealth, keep the body g o i n g as a
whole functioning mechanism. Naturally it would be ex-
pected that the casualties would be far heavier in the first
two defense lines (respiratory and alimentary s y s t e m s
and the blood and circulation) than in the home guard.
Death rates, when biologically classified, bear out this
expectation.
In the United States the kidneys and related excre-
tory organs are responsible for more deaths than the sex
organs. This relation is reversed in England and W a l e s ,
and in Sao Paulo. This difference is mainly due i n both
countries to premature birth. The higher premature
birth rate for these two localities as compared w i t h the
United States might conceivably be explained in any one
of several ways. I t might mean better obstetrics here
than in the other localities, or i t might mean that the
women of this country, as a class, are somewhat superior
physiologically in the matter of reproduction, when they
do reproduce, or it might be in some manner connected
with differences i n birth rates.
112 BIOLOGY OF D E A T H
T h e l a s t t h r e e o r g a n s y s t e m s , skeletal a n d m u s c u l a r
system, skin a n d endocrinal o r g a n s , a r e responsible for
so few d e a t h s relatively as n o t to be of serious moment.
T h e r e is one g e n e r a l consequence of t h e s e r e s u l t s u p o n
which I should like t o dwell a m o m e n t longer. I n a b r o a d
sense t h e efforts of public h e a l t h a n d hygiene h a v e been
directed a g a i n s t the affections c o m p r i s e d i n t h e first two
i t e m s i n t h e c h a r t , t h o s e of t h e r e s p i r a t o r y s y s t e m and
t h e a l i m e n t a r y t r a c t . T h e figures for t h e two five-year
p e r i o d s i n t h e U n i t e d S t a t e s , 1901-05 a n d 1906-10, indi-
cate r o u g h l y t h e r a t e of p r o g r e s s such m e a s u r e s a r e
m a k i n g , lqoking a t t h e m a t t e r f r o m a b r o a d biological
s t a n d p o i n t . I n reference to t h e r e s p i r a t o r y s y s t e m t h e r e
w a s a decline of f o u r t e e n p e r cent, i n t h e d e a t h r a t e be-
tween the two p e r i o d s . T h i s is s u b s t a n t i a l . I t is p r a c -
tically all accounted f o r i n p h t h i s i s , l o b a r p n e u m o n i a and
bronchitis. F o r t h e a l i m e n t a r y t r a c t t h e case w a s n o t
so good—Indeed f a r w o r s e .
B e t w e e n t h e t w o p e r i o d s t h e d e a t h r a t e f r o m t h i s cause
g r o u p fell only 1.8 p e r cent. A l l t h e g a i n m a d e in t y p h o i d
fever w a s a g r e a t d e a l m o r e t h a n offset b y diarrhoea a n d
enteritis ( u n d e r t w o ) , congenital debility a n d cancer.
Child welfare, both p r e n a t a l a n d p o s t n a t a l , seems by long
odds the m o s t hopeful direction i n w h i c h public h e a l t h
activities can expect, a t t h e p r e s e n t time, substantially to
r e d u c e t h e g e n e r a l d e a t h r a t e . T h i s i s a m a t t e r funda-
m e n t a l l y of education.
TOTALS
JOO
I
O \5 10 i5 20 25 SO 35 4O 45 SO 35 6O 6& 7O 75 60 65 BO OS /OO
AGE
FIQ. 27.—Diagram showing the specific death rate at each age for deaths from all causes
taken together.
the female curve lies, b y greater or less amounts, below
aoi
AOL
FIQ. 28.—Diagram showing tho ipecific death rat« »t r«.rb u^ Umu brmkdumu i>i lh#
circulatory »ytt«m, blood and btood-lormintf prgan* iGtuup I).
curve from what we have seen for the ease of all causes
of death. In the first place, the specific force of mortal-
i t y of this g r o u p of c a u s e s is r e l a t i v e l y l o w in i n f a n c y and
T H E CAUSES OF D E A T H 119
childhood. O u t of ?*• thounanri infantH of each *u»x e x p o s e d
to rink, only 7 mitlfK a m i 5 femalen d i e f r o m b r e a k d o w n
of this j r r o u p of o r g a n s d u r i n g t h e firnt y e a r of life.
The t r o u g h of t h e e u r v e a s s o c i a t e d w i t h t h e m o r t a l i t y of
childhood a n d y o u t h is v e r y m u c h less p o i n t e d t h a n in
the cane of **nll c a u s e s . M It in a s m o o t h l y r o u n d e d ,
r a t h e r t h a n a s h a r p l y p o i n t e d d e p r e s s i o n . I t in a l s o
noteworthy t h a t b e t w e e n a p p r o x i m a t e l y flic* a # e s of 5
and 35 tin* specific forci» of m o r t a l i t y from d i s e a s e s of t h e
circulatory n y s t e m a n d r e l a t e d o r g a n s is h i g h e r f o r
females t h a n it in f o r m u l e s . Thin condition of affairH 5B
probably c o n n e c t e d with t h e g r a v e r p h y s i o l o g i c a l change*)
and r e a d j u s t m e n t s called f o r t h by p u b e r t y in t h e f e m a l e
than a c c o m p a n y t h e g a m e vital cri*tm in t h e m a l a From
early a d u l t life*, nay i w 25-30 on, t h e Bpeciftc d e a t h r a t e
from dim # fWB of t h i * c i r c u l a t o r y HVHternnnd r e l a t e d o r g a n s
increa^en at itn ahiuwt itliHr»lntely c o n n t a n t r a t e until a g e
85 IH reached, Aft«*r t h a t , t h e r a t e of ittcreane HIOWB
down Hom<*w)mt. Of thorn* n o t c h i n g t h e ageg 9 5 4 0 0 , be-
tween 70 iiiiil BO o u t of each t h o u s a n d l i v i n g d i e f r o m
breakdown of thin g r o u p of orgnnH,
T h e upeeifm m o r t a l i t y c u r v e for d e a t h s f r o m b r e a k -
down of the* reMjunitnry «yHtt*mf tin nhown in F i g u r e 29,
pnmcmtN II ntjifitier «»f p o i n t s of p e c u l i a r iiit4Br«!8t, I n
the fitnt pliiei* w e n o t e tltnt t)tin orgiiri gyntern in m u c h
m o r e lifihli* t o b r e a k d o w n t h a n in the c i r c u l a t o r y ^y«tc*m
d u r i n g all t h e e a r l i e r yt*nr» of life tip to a b o u t age* GO-65.
Thd decline in t h e c u r v e from the* h i g h p o i n t of i n f a n c y t o
thci low p o i n t of t h « p e r i o d nbotit p u b e r t y in rnorci s h a r p
and «uildt*rt t h a n t h a t of tint c i r c u l a t o r y nyHUtm curve*
Again f htiwever,jtiKt an in i\m f o r m e r ctme,we noto t h a t t h e
specific force «f mnrtiiHty f r o m b r e a k d o w n of thi» o r g a n
impingiii m o r e heavily u p o n females than upon
120 BIOLOGY OF D E A T H
males in t h e y e a r s from 5-20. Thin difference la p r o b -
ably connected, as before, w i t h t h e g r e a t e r phyniological
d i s t u r b a n c e of p u b e r t y in t h e female t h a n in the* male.
40 4$ m if io m mmm m"
ACC
<4
all, b u t s h o u l d b e c h a r g e d , o n s u c h a b a s i s , a g a i n s t the
mother. T o g o further into detail, it is apparent that w h e n
a premature birth occurs it is because the reproductive
T H E CAUSES O F D E A T H 123
system of the m o t h e r , for s o m e r e a s o n o r other, did n o t r i s e
to the d e m a n d s of t h e s i t u a t i o n of c a r r y i n g t h e foetus t o
term. P r e m a t u r e b i r t h , in s h o r t , r e s u l t s f r o m a fail-
u r e or b r e a k d o w n i n some p a r t i c u l a r of t h e maternal
reproductive system. This failure m a y be caused in
various w a y s , which do n o t h e r e c o n c e r n u s . T h e e s s e n t i a l
feature from o u r p r e s e n t v i e w p o i n t is t h a t t h e r e p r o d u c -
tive system of t h e m o t h e r d o e s b r e a k down, a n d b y so
doing causes the d e a t h of t h e i n f a n t , a n d t h a t d e a t h i s
recorded s t a t i s t i c a l l y u n d e r t h i s title " P r e m a t u r e b i r t h . "
The d e a t h o r g a n i c a l l y is c h a r g e a b l e t o t h e m o t h e r .
A considerable n u m b e r of cases of p r e m a t u r e b i r t h
a r e unquestionably d u e t o p l a c e n t a l defects a n d t h e p l a -
centa is a s t r u c t u r e of foetal o r i g i n , so such d e a t h s could
n o t be p r o p e r l y c h a r g e d t o t h e m o t h e r . On t h e o t h e r
hand, however, t h e y w o u l d still s t a y i n t h e s a m e t a b l e be-
cause the p l a c e n t a m a y f a i r l y be r e g a r d e d a s a n o r g a n
intimately concerned in r e p r o d u c t i o n .
The same r e a s o n i n g which a p p l i e s t o p r e m a t u r e b i r t h s ,
mutatis mutandis, a p p l i e s to t h e i t e m " I n j u r i e s a t b i r t h . "
A n infant d e a t h r e c o r d e d u n d e r t h i s h e a d m e a n s t h a t
some p a r t of t h e r e p r o d u c t i v e m e c h a n i s m of t h e m o t h e r ,
either s t r u c t u r a l or functional, f a i l e d of n o r m a l p e r -
formance in t h e t i m e of s t r e s s . U s u a l l y " i n j u r y a t
b i r t h ' ' m e a n s a c o n t r a c t e d o r m a l f o r m e d p e l v i s of t h e
mother. B u t i n a n y c a s e t h e d e a t h is p u r e l y e x t e r n a l a n d
accidental f r o m t h e s t a n d p o i n t of t h e i n f a n t . I t is o r g a n -
ically c h a r g e a b l e t o a defect of t h e s e x o r g a n s of t h e
mother. T h e female pelvis, i n r e s p e c t of i t s c o n f o r m a -
tion, is a s e c o n d a r y s e x c h a r a c t e r .
T h e i m m e d i a t e r e a s o n f o r i n c l u d i n g syphilis a n d
gonococcus infection h e r e is obvious, but, p a r t i c u l a r l y i n
r e l a t i o n to syphilis, t h e p o i n t n e e d s f u r t h e r discussion.
124 BIOLOGY OF D E A T H
A s a -cause of a c t u a l d e a t h , s y p h i l i s f r e q u e n t l y a c t s
through the central nervous system, and the question may
f a i r l y be r a i s e d why, i n view of t h i s fact, s y p h i l i s in n o t
tabled t h e r e . T h e p o i n t well i l l u s t r a f m one of t h e fun-
d a m e n t a l difficulties in a n y o r g a n o l o g i c a l classification
of disease. I n t h e case of syphilis, h o w e v e r , the difficulty
in p r a c t i c e i s n o t n e a r l y so g r e a t an i t in in t h e o r y . A B
a m a t t e r of fact, m o s t of t h e d e a t h s from the effect of
syphilitic infection on t h e n e r v o u s s y s t e m arc* r e c o r d e d
i n v i t a l statistics b y r e p o r t i n g p h y s i c i a n s a n d vital s t a t i s -
ticians as d i s e a s e s of t h e n e r v o u s s y s t e m . F o r e x a m p l e ,
i t is p e r f e c t l y c e r t a i n t h a t m o s t of t h e dctuthH r e c o r d e d
a s due to " l o c o m o t o r a t a x i a " a r e f u n d a m e n t a l l y s y p h i l -
itic in origin. T h e r a t e of 5.4 for t h e R e g i s t r a t i o n A r e a
of t h e U n i t e d S t a t e s in 1906-10 f o r d e a t h s d u e t o s y p h i l i s
i s f a r lower, a s a n y clinician k n o w s , t h a n the* n u m b e r of
d e a t h s r e a l l y a t t r i b u t a b l e to syphilitic* infection. T h e s e
o t h e r d e a t h s , d u e to syphilis, a n d n o t r e p o r t e d u n d e r t h a t
title, a r e r e p o r t e d u n d e r t h e o r g a n which p r i m a r i l y
b r e a k s down a n d c a u s e s d e a t h , a s , for e x a m p l e , t h e b r a i n ,
a n d will i n t h e p r e s e n t s y s t e m of classification be included
u n d e r t h e n e r v o u s s y s t e m . A f t e r careful c o n s i d e r a t i o n ,
i t h a s seemed a s f a i r a s a n y t h i n g which could be d o n e t o
p u t t h e r e s i d u e of d e a t h s specifically r e p o r t e d a« d u e
t o syphilis u n d e r P r i m a r y a n d S e c o n d a r y S e x O r g a n s .
T h e r a t e , i n a n y event, i s s o s m a l l t h a t w h a t e v e r s h i f t w a a
m a d e could n o t sensibly affect t h e g e n e r a l r e s u l t s t o
which w e shall p r e s e n t l y come.
T u r n i n g n o w t o t h e c o n s i d e r a t i o n of F i g u r e 30, which
gives t h e c u r v e s of specific m o r t a l i t y f r o m b r e a k d o w n of
t h e r e p r o d u c t i v e o r g a n s , w e n o t e a t once the h i g h specific
d e a t h r a t e of i n f a n t s u n d e r one, r e c o r d e d by t h e f e m a l e
line. T h i s r a t e i s o v e r 40 p e r t h o u s a n d e x p o s e d t o r i s k .
T H E CAUSES OF D E A T H 125
I t includes, of c o u r s e , b o t h male* a n d f e m a l e i n f a n t s , dy-
ing from c o n g e n i t a l debility, p r e m a t u r e b i r t h a n d i n j u r i e s
at"birth,bwanK<su<rcordingt(; the r e a H o u i n g j u a i e x p l a i n e d ,
loorz
K> k B £J 3d JB Jb S 36 h 60 k
Of.
Fio. 32.—Diagram ahowing«p«cifio<Ifmth r&am »t#n@H *m (mm brmk*tvmn of thm
tt»d muaoular uyutvam (Group V),
too
ABL
WIQ. 33.—Diagram * how Jog th« »r*dfk rat<* of < frum breakdown
alimentary tract and aiwociated orgs
OOt _L
io is io JO 4$ so h do
ABC
Fio. 34.—Diagram showing the upcciflc death ttXm Rt «^rh *m from t»r#*lt«bwit o( th«
nervous tystom nod »en»c orgmwt (Group VIIh
a ot
Fio. 35.—Diagram showing the wslfi© d«*tk r*tm at €*<th tg« th*rgt*bte *«*Jt*t &t
skin (Group VIII). ^^
curve lies well below the male curve practically through-
out its course-
Deaths from failure to function properly of the organs
T H E CAUSES OF D E A T H 133
of the endocrinal system, including the thyroid gland,
suprarenal glands, etc., d o not become significant until
middle life in the case of the male, as shown i n F i g u r e 36,
ENDOCRINAL SYSTEM
- ^ \
\ /
0.01 \ 1 1
tO 15 20 ZS 3O 35 4O 45 SO 55 6O 65 7O 75 6O 65 &Q OS 100
A3E
a l t h o u g h i n t h e f e m a l e t h e c u r v e " b e g i n s t o r i s e f r o m p u -
b e r t y o n . T h e s p e c i f i c r a t e s a t a l l a g e s , o f c o u r a e , a x e
e x t r e m e l y s m a l l , p r a c t i c a l l y n e v e r r i s i n g t o m o r e t h a n
134 BIOLOGY OF D E A T H
1/10 of one person per thousand exposed to risk. The
well-known fact that these glandular organs, whose se-
cretions are so important for the normal conditions of
KOOOr—
O.I
iO E6 30 35 AQ 43 JO £5 60 65 7
FIG. 37—
. Daigramcosvheorew AOL from al other cause* of dea
dnigin ththee pspreecceifdicingdecaathtegorarietess(Group X).
life, are much more unstable and liable to breakdown
in the female than in the male, is strikingly shown by
this diagram.
T H E CAUSES OF D E A T H 135
Finally, we have the diagram for our omnium gatherum
group, the " A l l other causes of death," i n Figure 37.
H e r e we see that, because of accidental and violent deaths,
the male specific mortality curve lies far above the
female, from youth until old age has set in, about age 75.
F r o m that point on to the end of the span of life both
curves ascend rapidly together, as a result of the deaths
recorded as resulting from senility. Eventually it is
t o be expected that no deaths will be registered as result-
i n g from senility. W e shall have them all put more nearly
where they belong.
These diagrams of specific forces of mortality give
altogether a remarkably clear and definite picture of how
death occurs among men. W e see that failure of certain
organ systems, such as the lungs, the heart, the kidneys,
to maintain their structural and functional integrity, has
an overwhelmingly great effect in determining the total
r a t e of mortality as compared with some of the other
organ systems. One cannot but be impressed, too, with
the essential orderliness of the phenomena we have ex-
amined. The probability of any particular organ system
breaking down and causing death is mathematically def-
inite at each age, and changes in a strikingly orderly
manner as age changes, as is shown in Table 11. Thus
w e find that in the first year of lif e it is the alimentary
tract and its associated organs which most frequently
break down and cause death. F r o m age 1 to age 60
t h e specific force of mortality from breakdown of the
respiratory system is higher (with a few insignificant
exceptions in the females) usually by a considerable
amount, than that associated with any other organ system
of the body. F r o m 60 to 90 years of age the circulatory
136 BIOLOGY OF D E A T H
s y s t e m t a k e s t h e f r o n t r a n k , w i t h a h i g h e r specific mor-
tality r a t e t h a n a n y o t h e r o r g a n s y s t e m .
TABLE 11
The most fatal organ systems at different ages
MALES FEMALES
Per cent, of all Per cent, of all
biologically Organ system Age Organ system biologically
classifiable Group classifiable
concerned in largest deaths
deaths due to concerned in largest
proportion proportion due to
breakdown of of fatalities of fatalities breakdown of
specified organ specified organ
system system
68.8 Alimentary tract 0— 1 Alimentary tract 40.6
50.1 Respiratory 1— 4 Respiratory 51.3
41.2 Respiratory 5— 9 Respiratory 42.5
27.1 Respiratory 10—14 Respiratory 33.3
43.6 Respiratory 15—19 Respiratory 43.8
52.6 Respiratory 20—24 Respiratory 46.0
49.7 Respiratory 25—29 Respiratory 44.2
45.6 Respiratory 30—34 Respiratory 39.5
39.9 Respiratory 35—39 Respiratory 33.2
33.3 Respiratory- 40—44 Respiratory 27.5
28.0 Respiratory 45—49 Respiratory 22.1
23.6 Respiratory 50—54 Alimentary tract 21.6
25.0 Circulatory 55—59 Alimentary tract 22.6
28.4 Circulatory 60—64 Circulatory 24.4
30.9 Circulatory 65—69 Circulatory 25.6
32.5 Circulatory 70—74 Circulatory 28.0
32.9 Circulatory 75—79 Circulatory 28.4
33.3 Circulatory 80—84 Circulatory 30.4
85—89 Circulatory 30.8
TABLE 13
Showing the relative influence of the primary germ layers in human mortality
(Items 64 and 65 charged to mesoderm)
Death rate per 100,000 due to functional breakdown
of organs embryologically developing from
Locality
Ecto- Per Meso- Per Endo- Per
derm cent. derm cent. derm cent.
United States Registration
Area, 1906-10 116.9 8.7 499.4 37.4 719.6 53.9
United States Registration
Area, 1901-05 137.3 9.8 480.4 34.2 786.2 56.0
England and Wales, 1914... 107.9 6.7 443.2 36.0 681.5 55.3
Sao Paulo, 1917 101.3 6.3 501.6 31.1 1009.9 62.6
c a s e s still unsettled, q u e s t i o n s of embryology. F u r t h e r -
m o r e , t h e original s t a t i s t i c a l r u b r i c s u n d e r which t h e d a t a
a r e compiled b y r e g i s t r a r s of v i t a l statistics were n e v e r
p l a n n e d w i t h such a n object a s t h i s i n mind. Still t h e t h i n g
s e e m e d w o r t h t r y i n g because of t h e biological i n t e r e s t
w h i c h would a t t a c h t o t h e r e s u l t , even t h o u g h i t w e r e some-
140 BIOLOGY OF D E A T H
w h a t c r u d e and, i n r e s p e c t of m i n o r a n d insignificant
details, open t o criticism. I t is n o t possible h e r e to go into
details as to how t h e c a u s e s of d e a t h w e r e combined in
m m
m
M M . i s ?
1 i
ENDODERM MESODERM ECTODERM
FIG. 38.—Diagram showing the percentages of biologically classifiable human mortality
resulting from breakdown of organs developing from the different germ layers. Upper bar
of pair gives upper limit of mortality chargeable to ectoderm: lower bar gives lower limit of
mortality chargeable to ectoderm.
making up the final tables. F o r these details one must
refer to the original papers.
Tables 12 and 13, and F i g u r e 38, g i v e the results for
t h e crude m o r t a l i t y of t h e U . S. R e g i s t r a t i o n A r e a , E n g -
land and Wales, and S a o Paulo, Brazil.
E M B R Y O L O G Y A N D H U M A N M O R T A L I T Y 141
T h e figures s h o w t h a t in m a n , the highent p r o d u c t of
organic evolution, a b o u t 57 pc*r cent, of all t h o biologically
classifiable d e a t h n m m It f r o m a b r e a k d o w n a n d f a i l u r e
f u r t h e r to function of orgmiH a r i H I n g f r o m t h e cmdodorm
in t h e i r e m b r y o l o g i c a l d e v e l o p m e n t , w h i l e b u t f r o m 8
p e r cent, to 13 p e r cent, can be r e g a r d e d an a r e s u l t of
b r e a k d o w n of o r g a n HyntemH n r i m n g f r o m t h e e c t o d e r m .
T h e r e m a i n i n g 30 to 35 p e r cent, of t h e m o r t a l i t y reHults
from f a i l u r e of m e n o d e r m i c o r g a n s T h e t w o v a l u e s
stated f o r e c t o d e r m a n d meworform, s h o w n b y t h e t w o
b a r s in t h e d i a g r a m , differ by v i r t u e of t h e f a c t t h a t t w o
i m p o r t a n t c a u s e s of d e a t h , c e r e b r a l h e m o r r h a g e a n d
apoplexy, arid s o f t e n i n g of t h e b r a i n , a r e p u t in t h e
one cane with t h e e c t o d e r m a n d in t h e o t h e r c a s e w i t h
the m e s o d e r m . T h e p a t h o l o g i c a l a r g u m e n t s f o r tho one
disposition a s agairiHf t)m o t h e r of thene t w o d i s e a s p s a r e
i n t e r e s t i n g b u t btck of s p a c e p r e v e n t s t h e i r e x p o s i t i o n
here. I h a v e choson r a t h e r t o prc*si»nt tho facts in
both w a y s .
T a k i n g a g e n e r a l v i e w of c o m p a r a t i v e a n a t o m y a n d
e m b r y o l o g y i t in e v i d e n t t h a t in tho e v o l u t i o n a r y h i s t o r y
t h r o u g h which m a n a n d t h e h i g h e r v e r t e b r a t e s h a v n p a s s e d
it is t h e e c t o d e r m which him been m o s t w i d e l y differ-
e n t i a t e d f r o m i t s primitive* condition! t o t h e v a l i d i t y of
which s t a t e m e n t t h e c e n t r a l n e r v o u s s y s t e m f u r n i s h e s tho
m o s t p o t e n t evidonco. T h e e n d o d e r m h a s been l e a s t p r o -
g r e s s i v e l y c h a n g e d s t r u c t u r a l l y a n d f u n c t i o n a l l y in t h e
p r o c e s s of e v o l u t i o n , while tho m r a o d e r m occupies, on tho
whole, a n i n t e r m e d i a t e p o s i t i o n in t h i s r e s p e c t .
D e g r e e of d i f f e r e n t i a t i o n of o r g a n s in e v o l u t i o n im-
plies d e g r e e of a d a p t a t i o n t o e n v i r o n m e n t . F r o m t h e p r e -
s e n t p o i n t of v i e w we sea t h a t t h e g u m layar f t h e e n d o -
d e r m , which h a s evolved o r bueome d i f f e r e n t i a t e d l e a s t in
142 BIOLOGY OF D E A T H
the p r o c e s s of evolution in l e a s t able t o m e e t successfully
t h e vicissitudes of the e n v i r o n m e n t . T h e e c t o d e r m has
changed m o s t in the c o u r s e of evolution* Of thin t h e cen-
t r a l n e r v o u s s y s t e m of m a n is t h e boat proof. T h e r e
h a v e also been formed in t h e proctma of differentiation,
protecti\ r e m e c h a n i s m s , t h e skull a n d v e r t e b r a ! column,
which v e r y well keep t h e delicate a n d h i g h l y organized
central n e r v o u s s y s t e m a w a y f r o m d i r e c t c o n t a c t with
t h e e n v i r o n m e n t . T h e skin a l s o exhibits m a n y differen-
tiations of a highly a d a p t i v e n a t u r e t o r e s i s t environmen-
t a l difficulties. Tt is then n o t B u r p m i n g t h a t t h e o r g a n
s y s t e m s developed from the e c t o d e r m b r e a k down and
lead to d e a t h lens f r e q u e n t l y t h a n a n y o t h e r . T h e fig-
u r e s m a k e i t c l e a r t h a t raan'n g r e a t e s t e n e m y ia h i s own
endoderm. E v o l u t i o n a l l y Bpeaking, it in a v e r y old-
fashioned a n d out-of-date a n c e s t r a l relic, whieh c a u s e s him
a n infinity of t r o u b l e . P r a c t i c a l l y all public lmalth ac-
tivities a r e directed t o w a r d s o v e r c o m i n g t h e difficulties
which a r i s e because m a n c a r r i e s a b o u t thin a n t e d i l u v i a n
s o r t of e n d o d e r m . W e e n d e a v o r t o modify t h e environ-
ment, a n d soften its aaperiticm d o w n to t h e p o i n t w h e r e
o u r own inefficient e n d o d e r m a l m e c h a n i s m c a n cope with
them, by such m e t h o d s a s p r e v e n t i n g b a c t e r i a l contam-
i n a t i o n of w a t e r , food a n d t h e like, w a r m i n g the a i r we
b r e a t h e , etc. B u t o u r e c t o d e r m r e q u i r e s no imch exten-
sive a m e l i o r a t i o n of the e n v i r o n m e n t T h e r e a r e a t m o s t
only a v e r y few, if a n y , g e r m s whieh c a n g a i n e n t r a n c e to
t h e body t h r o u g h t h e n o r m a l , h e a l t h y u n b r o k e n ikin*
W e do, t o be s u r e , w e a r clothes* B u t i t m a t leant a d e b a t -
able question w h e t h e r , u p o n m a n y p a r t * of t h e e a r t h ' s
surface, w e should n o t b e b e t t e r off w i t h o u t t h e m from
t h e p o i n t of view of h e a l t h .
T h e s e d a t a indicate f u r t h e r in a n o t h e r m a n n e r how
E M B R Y O L O G Y A N D H U M A N M O R T A L I T Y 143
i m p o r t a n t a r e the f u n d a m e n t a l e m b r y o l o g i c a l f a c t o r s
i n d e t e r m i n i n g t h e m o r t a l i t y of m a n . Of t h e t h r e e local-
ities compared, E n g l a n d a n d t h e U n i t e d S t a t e s m a y be
f a i r l y r e g a r d e d as m u c h m o r e a d v a n c e d in m a t t e r s of
public h e a l t h a n d s a n i t a t i o n t h a n S a o P a u l o . T h i s f a c t
is reflected w i t h p e r f e c t p r e c i s i o n a n d j u s t i c e i n t h e r e -
lative p r o p o r t i o n of t h e d e a t h r a t e s f r o m e n d o d e r m a n d
ectoderm. I n t h e U n i t e d S t a t e s a n d E n g l a n d a b o u t 55
p e r cent, of t h e classifiable d e a t h s a r e c h a r g e a b l e t o e n d o -
d e r m a n d a b o u t 9 t o 14.5 p e r cent, to e c t o d e r m . I n S a o
P a u l o 62.6 p e r cent, fall w i t h t h e e n d o d e r m , a n d b u t 6.3
to 8.4 p e r cent, w i t h t h e ectoderm. Since p u b l i c h e a l t h
m e a s u r e s can a n d do affect p r a c t i c a l l y only t h e d e a t h
r a t e chargeable t o e n d o d e r m , t h i s result, which is a c t u a l l y
obtained, is precisely t h a t which would be expected.
A question which n a t u r a l l y occurs is as t o w h a t t h e
a g e d i s t r i b u t i o n of b r e a k d o w n of ectodermic, m e s o d e r -
mic, o r e n d o d e r m i c o r g a n s m a y be. A r e t h e e n d o d e r m i c
o r g a n s , for example, r e l a t i v e l y m o r e liable to b r e a k d o w n
i n e a r l y life, a n d less so later, a s g e n e r a l o b s e r v a t i o n
w o u l d lead one t o conclude?
T o a n s w e r this a n d s i m i l a r q u e s t i o n s which come t o
m i n d i t is n e c e s s a r y t o d i s t r i b u t e t h e specific r a t e s of
T a b l e 9 u p o n a n embryological basis.
I n F i g u r e 39 t h e r e s u l t of d o i n g t h i s is s h o w n f o r
m a l e s . W e note t h a t p r i o r to a g e 60 t h e c u r v e f o r t h e
b r e a k d o w n of o r g a n s of e n d o d e r m i c o r i g i n lies a t t h e
t o p of t h e d i a g r a m ; n e x t below i t comes t h e c u r v e f o r
thej b r e a k d o w n of o r g a n s of m e s o d e r m i c / o r i g i n ; a n d
finally a t the b o t t o m t h e c u r v e for the b r e a k d o w n of or-
g a n s of ectodermic origin. All t h r e e of t h e c u r v e s h a v e
i n g e n e r a l t h e f o r m of a specific d e a t h r a t e c u r v e . T h e
r a t e s f o r all t h r e e g e r m l a y e r s a r e r e l a t i v e l y h i g h i n in-
144 BIOLOGY OF D E A T H
fancy a n d d r o p a t a p r a c t i c a l l y c o n s t a n t r a t e t o a low
p o i n t in e a r l y y o u t h . I n infancy t h e h e a v i e s t m o r t a l i t y
i n m a l e s is due t o t h e b r e a k d o w n of o r g a n s of e n d o d e r m i c
(,000 rz
100
, ._
0.5 JO i5Z0Z5dO3540453OS560657O7560Q59OS5
AGE.
FIG. 40.—Showing specific death rates for females, classified in the same manner as in Fig. 3 9 .
outlines one p a r t of t h e p i c t u r e of t h e g e n e r a l biology o f
death. I t h a s b e e n s h o w n i n w h a t h a s p r e c e d e d t h a t n a t -
u r a l death is n o t a n e c e s s a r y o r i n h e r e n t a t t r i b u t e o r
E M B R Y O L O G Y A N D H U M A N M O R T A L I T Y 147
consequence of life. M a n y cells a r e p o t e n t i a l l y i m m o r -
t a l a n d t h e p o t e n t i a l i t y is a c t u a l l y r e a l i z e d if a p p r o p r i a t e
conditions a r e p r o v i d e d . P r o t o z o a a r e i m m o r t a l . G e r m
cells a r e i m m o r t a l . V a r i o u s s o m a t i c cells, a n d e v e n t i s -
sues h a v e been p r o v e d to be p o t e n t i a l l y i m m o r t a l b y
d e m o n s t r a t i n g in a v a r i e t y of w a y s t h a t u n d e r a p p r o -
p r i a t e c o n d i t i o n s t h e y c o n t i n u e to live indefinitely. This
is t h e lesson t a u g h t XXH on t h e one h a n d b y successive
t r a n s p l a n t a t i o n s of t u m o r cells, which a r e only modified
s o m a t i c cells, a n d on t h e o t h e r h a n d by successful cul-
t u r e of m a n y s o r t s of s o m a t i c cells in vitro.
A n a l y t i c a l c o n s i d e r a t i o n of t h e m a t t e r s h o w s v e r y
c l e a r l y t h a t t h e s o m a t a of m u l t i c e l l u l a r o r g a n i s m s
die b e c a u s e of t h e d i f f e r e n t i a t i o n s a n d s p e c i a l i z a t i o n s
of s t r u c t u r e a n d function which t h e y e x h i b i t in t h e i r
m a k e - u p . C e r t a i n cells a r e d i f f e r e n t i a t e d t o c a r r y on
c e r t a i n specialized f u n c t i o n s . I n t h i s s p e c i a l i z a t i o n t h e y
f o r e g o t h e i r p o w e r of i n d e p e n d e n t a n d indefinitely con-
t i n u e d existence. T h e cells l i n i n g t h e l u n g s , for e x a m p l e ,
m u s t d e p e n d in t h e b o d y u p o n t h e u n f a i l i n g n o r m a l ac-
t i v i t y of t h e cells of t h e a l i m e n t a r y t r a c t a n d t h e blood in
o r d e r t h a t t h e y , t h e e p i t h e l i a l cells of t h e l u n g s , m a y g e t
p r o p e r n u t r i t i o n . I f in s u c h a n i n t e r l o c k i n g a n d m u -
tually dependent system any one p a r t through accident
o r in a n y w a y w h a t e v e r g e t s d e v i a t e d f r o m i t s n o r m a l
functioning, t h e b a l a n c e of t h e whole s y s t e m is u p s e t If
t h e d e p a r t u r e of a n y p a r t f r o m its n o r m a l f u n c t i o n a l
c o u r s e is g r e a t e n o u g h t o be beyond c o r r e c t i o n p r o m p t l y
t h r o u g h t h e n o r m a l r e g u l a t o r y p o w e r s of t h e o r g a n i s m ,
d e a t h of t h e w h o l e will s u r e l y e n s u e .
W h a t I h a v e t r i e d to s h o w in t h i s a n d t h e p r e c e d i n g
c h a p t e r i s a q u a n t i t a t i v e p i c t u r e of h o w t h e different
o r g a n s y s t e m s g e t o u t of b a l a n c e , a n d w r e c k t h e w h o l e
148 BIOLOGY OF D E A T H
machine. T h e b r o a d o r d e r l i n e s s a n d lawfulness of the
whole business of h u m a n m o r t a l i t y is i m p r e s s i v e . W e
h a v e seen t h a t different o r g a n s y s t e m s h a v e well-defined
times of b r e a k d o w n . Or, p u t in a n o t h e r w a y , we see t h a t
in the h u m a n o r g a n i s m , j u s t as i n t h e automobile, the
serviceability of the different p a r t s v a r i e s g r e a t l y . The
h e a r t o u t w e a r s t h e l u n g s , t h e b r a i n o u t w e a r s both. B u t
we h a v e f u r t h e r , I believe, got a n i n k l i n g of t h e funda-
m e n t a l r e a s o n w h y t h e s e t h i n g s a r e so. I t is b r o a d l y
speaking, b e c a u s e evolution is a p u r e l y m e c h a n i s t i c p r o -
cess i n s t e a d of b e i n g a n intelligent one. A l l t h e p a r t s a r e
not perfected b y evolution t o even a n a p p r o x i m a t e l y equal
degree. I t is conceivable t h a t a n o m n i p o t e n t p e r s o n
could h a v e m a d e a m u c h b e t t e r m a c h i n e , a s a whole, t h a n
t h e h u m a n b o d y which evolution h a s p r o d u c e d , assuming,
of course, t h a t h e h a d first l e a r n e d t h e t r i c k of m a k i n g
self-regulating a n d s e l f - r e p r o d u c i n g m a c h i n e s , such as
living m a c h i n e s a r e . H e would p r e s u m a b l y h a v e m a d e an
e n d o d e r m w i t h as good r e s i s t i n g a n d w e a r i n g qualities
a s the m e s o d e r m or ectoderm. E v o l u t i o n by the h a p -
h a z a r d p r o c e s s of t r i a l a n d e r r o r w h i c h w e call n a t u r a l
selection, m a k e s each p a r t only j u s t good enough to get
by. I n t h e v e r y n a t u r e of t h e p r o c e s s itself i t cannot
possibly do a n y t h i n g a n y m o r e c o n s t r u c t i v e t h a n this.
T h e w o r k m a n s h i p of evolution, f r o m a mechanical
p o i n t of vi£w, is e x t r a o r d i n a r i l y like t h a t of t h e a v e r a g e
automobile r e p a i r m a n . If evolution h a p p e n s to be fur-
nished b y v a r i a t i o n w i t h fine m a t e r i a l s , a s in the case
of t h e n e r v o u s system, i t h a s n o objection to u s i n g them,
b u t i t is equally r e a d y t o u s e t h e s h o d d i e s t of e n d o d e r m
p r o v i d e d i t will hold t o g e t h e r j u s t l o n g enough t o get
the machine by t h e reproductive period.
I t f u r t h e r m o r e seems t o m e t h a t t h e r e s u l t s p r e s e n t e d
E M B R Y O L O G Y A N D H U M A N M O R T A L I T Y 149
in t h i s c h a p t e r a d d one m o r e link to t h e a l r e a d y Htrong
chain of evidence which i n d i c a t e s the h i g h l y i m p o r t a n t
p a r t p l a y e d by i n n a t e c o n s t i t u t i o n a l biological f a c t o r s
as c o n t r a s t e d w i t h e n v i r o n m e n t a l f a c t o r s in t h e d e t e r -
m i n a t i o n of t h e o b s e r v e d r a t e s of h u m a n m o r t a l i t y . Here
we h a v e g r o u p e d h u m a n m o r t a l i t y i n t o b r o a d classes
which r e s t u p o n a s t r i c t l y biological b a s i s . W h e n t h i s
is d o n e i t is f o u n d t h a t t h e p r o p o r t i o n a t e s u b d i v i s i o n of
the m o r t a l i t y a m o n g t h e s e v e r a l c a u s e s — i n s h o r t t h e
d e a t h ratios in t h e s e n s e of Fisher—-is s t r i k i n g l y s i m i l a r
in s u c h widely d i s s i m i l a r e n v i r o n m e n t s a s t h e U n i t e d
S t a t e s , E n g l a n d a n d S o u t h e r n Brazil.
CHAPTER VI
T H E I N H E E I T A N C E O F D U R A T I O N OF
LIFE IN MAN
10 15 ZO 25 30 35 40 45 SO 55 6O 65 70 75 6O 65 30 35
A6L
FIG. 41.—Showing survival curves of members of the Hyde family (Plotted from Bell's data).
b e g i n n i n g «fc a g e 15 a n d c o n t i n u i n g t o a g e 90, t h e f e m a l e
c u r v e lies below t h a t f o r t h e m a l e s , w h e r e a s n o r m a l l y f o r
t h e g e n e r a l p o p u l a t i o n i t lies above it. T h i s d e n o t e s a
s h o r t e r a v e r a g e d u r a t i o n of life i n t h e f e m a l e s t h a n i n
t h e m a l e s , t h e a c t u a l figures b e i n g 35.8 y e a r s f o r t h e m a l e s
a n d 33.4 y e a r s f o r t h e f e m a l e s . Bell a t t r i b u t e s t h e dif-
f e r e n c e t o t h e s t r a i n of c h i l d - b e a r i n g b y t h e f e m a l e s i n
154 BIOLOGY OF D E A T H
t h i s r a t h e r highly fertile g r o u p of p e o p l e , belonging in
t h e m a i n t o a p e r i o d w h e n r e s t r i c t i o n s u p o n size of family
w e r e less common a n d less e x t e n s i v e t h a n now. I n the
second place, t h e f e m a l e lx c u r v e is a c t u a l l y convex to
t h e base t h r o u g h o u t a c o n s i d e r a b l e p o r t i o n of middle
life w h e r e a s , n o r m a l l y , t h i s p o r t i o n of t h e c u r v e p r e s e n t s
a concave face to t h e b a s e .
A p a r t f r o m t h e s e d e v i a t i o n s , which a r e of no partic-
u l a r significance f o r t h e u s e which Bell m a k e s of the
d a t a , t h e H y d e m a t e r i a l is essentially n o r m a l a n d simi-
l a r to w h a t one would expect t o find in a r a n d o m sample
of the g e n e r a l p o p u l a t i o n . In t h i s m a t e r i a l t h e r e were
2,287 cases in which t h e a g e s a t d e a t h of t h e p e r s o n s and
t h e ages a t d e a t h of t h e i r f a t h e r s w e r e known. I t occurred
to Bell t o a r r a n g e t h i s m a t e r i a l in Huch a w a y aa to
show w h a t , if any, r e l a t i o n exiated between a g e a t d e a t h
of the p a r e n t a n d t h a t of the offspring. H e a r r a n g e d
t h e p a r e n t s into f o u r g r o u p s , a c c o r d i n g t o t h e a g e a t which
t h e y died, a n d t h e offspring i n t o five g r o u p s u p o n the
s a m e b a s i s . I n t h e c a s e of the p a r e n t s t h e g r o u p s w e r e :
F i r s t , those d y i n g u n d e r 4 0 ; second, between 40 a n d 6 0 ;
t h i r d , between 60 a n d 8 0 ; a n d f o u r t h , a t a g e HO a n d over.
T h e g r o u p s for t h e offspring wero t h e flame*, e x c e p t t h a t
the first w a s divided into two p a r t s , n a m e l y , thosie d y i n g
u n d e r 20 a n d those d y i n g between 20 a n d 40, T h e result-
i n g figures a r e exhibited in T a b l e 14.
T h e r e s u l t s f o r f a t h e r a n d o f f s p r i n g a r e s h o w n in
F i g u r e 42, b a s e d upon, t h e d a t a of T a b l e 14. I n each
of t h e 5 polygons, one f o r each offspring g r o u p , t h e first
d o t shows t h e p e r c e n t a g e of f a t h e r s d y i n g u n d e r 4 0 ;
t h e second d o t t h e p e r c e n t a g e of f a t h e r s d y i n g hot ween
40 a n d 6 0 ; a n d so on, t h e l a s t d o t in each c u r v e s h o w i n g
t h e p e r c e n t a g e of f a t h e r s d y i n g a t a g e 80 a n d o v e r . I t
THE INHERITANCE OF DURATION 155
TABLE 14
Analysis of the Hyde family data by person's age at death, showing the number
and percentage having (a) fathers and (b) mothers who died
at the age periods named. (From Bell)
Father's age at death
Person's age at death
Stated -40 40-60 60-80 80-}-
Stated. .. 2,287 66 522 1,056 643
Under 20 669 20 189 299 161
20 and under 40 538 18 140 269 111
40 and under 60 . . . 467 12 116 215 124
60 and under 80 428 13 57 196 162
80 and over 185 3 20 77 85
Percentages
Stated 100.0 2.9 22.8 46.2 28.1
Under 20 100.0 3.0 28.2 44.7 24.1
20 and under 40 . . . . 100.0 3.4 26.0 50.0 20.6
40 and under 60 100.0 2.6 24.8 46.0 26.6
60 and under 80 100.0 3.0 13.3 45.8 37.5
80 and over 100.0 1.6 10.8 41.6 46.0
40 to so
Via. 42.Influence
42.—Influence of fntWa a« a«f*AtAt<i^ftth
<i«^ftth uptm
uptm bngevit
bngevity of tiiUprtua Iirst <it»t in
each di
diagram «how»
h th th« n<*r<!imtag«»lli&vtna;f fftth«Ts
h who
h i cited
d tntMi mmml < h g»t«twai»
I«*t thr
<It g»ttwai
nge having fath«r» who dwd from 4t>0J;4t>-0fJ; third dot thu purmmUkWn
U h hftvinjc
falhrra
fh who
h dm!d!
f 00-«U;
from 00«U futh
fourth ddot
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thepp«re«»iUg@
« U hhaving i fth father* who <H«d HO f (Aftrfllil
21 p e r cent, of t h e f a t h e r s of o f f s p r i n g d y i n g b e t w e e n 20
a n d 40 lived t o be 80 y e a r s o r over. F o r t h e n e x t longer-
lived g r o u p of offspring, d y i n g between 40 a n d 60, the
p e r c e n t a g e of f a t h e r s l i v i n g to 80 o r o v e r roue t o 27 p e r
cent. I n t h e n e x t h i g h e r g r o u p , t h e p e r c e n t a g e is n e a r l y
38, and finally t h e e x t r e m e l y long-lived g r o u p of offspring,
t h e 185 p e r s o n s w h o died a t a g e s of 80 a n d o v e r , h a d 46
p e r cent, o r n e a r l y one-half of t h e i r f a t h e r s l i v i n g t o the
s a m e g r e a t a g e . I n o t h e r w o r d s , w e see in g e n e r a l t h a t
t h e longer-lived a g r o u p of o f f s p r i n g i s , on t h e a v e r a g e ,
t h e longer-lived a r e t h e i r f a t h e r s , o n t h e a v e r a g e ; or,
p u t in a n o t h e r w a y , t h e h i g h e r t h e p e r c e n t a g e of v e r y
T H E I N H E R I T A N C E OF DURATION 157
long-lived f a t h e r s w h i c h this g r o u p will h a v e a s com-
pared with shorter-lived individuals.
F i g u r e 43 shows the s a m e s o r t of d a t a f o r m o t h e r s
a n d offspring. H e r e w e see t h e c u r v e of g r e a t l o n g e v i t y
of p a r e n t s r i s i n g in an even m o r e m a r k e d m a n n e r t h a n
w a s t h e c a s e w i t h f a t h e r s of offspring. T h e g r o u p of
30
- 40 60 4O 6Q SO - 40 AO £0 • 4O 60 BO -
4O 60 00 OO m O & 0 t i 0 B 40 60
CO AQ
Fifl, 43.—Influence of mother's agn at death upon longevity of offspring. First dot in
each diAfr&m tihow* tho per centAge having mother* who died at 40; second dot the per-
0ntift«f« having mother* who died at 40-60; third dot the percentage having mothers who
difid eb-SO; fourth dot the perotntftge having mothern who died 80f (After Bell).
o f f s p r i n g d y i n g a t a g e s u n d e r 20 h a d only 19 p e r c e n t
of t h e i r m o t h e r s l i v i n g t o 80 a n d o v e r , w h e r e a s t h e
g r o u p of o f f s p r i n g who lived to 80 a n d b e y o n d h a d 41
p e r c e n t of t h e i r m o t h e r s a t t a i n i n g t h e s a m e g r e a t a g e .
A t t h e s a m e t i m e w e n o t e f r o m t h e d o t t e d line a t t h e bot-
t o m of t h e c h a r t t h a t a s t h e a v e r a g e a g e a t d e a t h of t h e
o f f s p r i n g i n c r e a s e s , t h e p e r c e n t a g e of m o t h e r s d y i n g a t
e a r l y a g e s , n a m e l y , u n d e r 40, a s g i v e n b y t h e first d o t s ,
s t e a d i l y decreases f r o m 17 p e r c e n t a t t h e first g r o u p t o
j n s t o v e r 5 p e r cent, f o r t h e o f f s p r i n g d y i n g a t v e r y
advanced ages.
158 BIOLOGY OF D E A T H
T h e s e s t r i k i n g r e s u l t s d e m o n s t r a t e a t once t h a t t h e r e
is a definite a n d close connection b e t w e e n the a v e r a g e
longevity of p a r e n t s a n d t h a t of t h e i r children. E x -
t r e m e l y long-lived c h i l d r e n h a v e a m u c h h i g h e r percent-
age of e x t r e m e l y long-lived p a r e n t s t h a n d o s h o r t e r lived
children. W h i l e t h e d i a g r a m s d e m o n s t r a t e t h e fact of
this connection, t h e y d o n o t m e a s u r e i t s i n t e n s i t y with
as great precision as can be obtained by other methods
of dealing w i t h t h e d a t a . A little f a r t h e r on we shall
take u p t h e c o n s i d e r a t i o n of t h i s m o r e p r e c i s e m e t h o d
of m e a s u r e m e n t of t h e h e r e d i t a r y influence in respect
of longevity.
I n the p r e c e d i n g d i a g r a m s w e h a v e considered each
p a r e n t s e p a r a t e l y i n connection w i t h t h e offspring in
TABLE 15
Longevity of parents of persons dying at 80 and over. {From Bell)
Lived to be 80+
Neither parent 827 44 5.3
One parent (not other) 583 57 9.8
Both parents 184 38 20.6
Father (not mother) 337 38 11.3
Mother (not father) 246 19 7.7
r e g a r d to longevity. W e shall, of c o u r s e , get precisely
t h e s a m e k i n d of r e s u l t if we c o n s i d e r b o t h p a r e n t s to-
gether. F o r t h e s a k e of simplicity, t a k i n g only the cases
of e x t r e m e longevity, n a m e l y , p e r s o n s l i v i n g t o 80 or
over—the e s s e n t i a l d a t a a r e g i v e n in T a b l e 15.
F r o m this t a b l e i t is seen t h a t w h e r e n e i t h e r p a r e n t
lived t o b e 80, only 5.3 p e r cent, of t h e offspring lived to
be 80 o r over, t h e p e r c e n t a g e b e i n g b a s e d u p o n 827
THE INHERITANCE OF DURATION 159
cases. W h e r e one p a r e n t , b u t n o t t h e o t h e r , lived to be
80 or older, 9.8 p e r cent, of t h e o f f s p r i n g lived to be 80
or older, t h e p e r c e n t a g e h e r e b e i n g b a s e d u p o n 583 cases.
W h e r e b o t h p a r e n t s lived t o be 80 o r o l d e r 20.6 p e r cent, of
the p e r s o n s lived t o t h e s a m e g r e a t a g e , t h e p e r c e n t a g e be-
i n g b a s e d u p o n 184 cases. T h u s i t a p p e a r s t h a t i n t h i s
g r o u p of p e o p l e f o u r t i m e s as m a n y a t t a i n e d g r e a t l o n g e v -
i t y if both t h e i r p a r e n t s lived t o a n a d v a n c e d a g e , a s
a t t a i n e d this a g e w h e n n e i t h e r p a r e n t exhibited g r e a t
longevity. T h e figures f r o m t h e H y d e f a m i l y seem f u r -
t h e r to i n d i c a t e t h a t t h e t e n d e n c y of l o n g e v i t y i s i n h e r i t e d
more strongly through the father t h a n through the
m o t h e r . W h e r e t h e f a t h e r , b u t n o t t h e m q t h e r , lived t o
be 80 o r older, 11.3 p e r cent, of t h e p e r s o n s lived t o a g e
80 or m o r e , t h e r e b e i n g 337 cases of t h i s kind. W h e r e
t h e m o t h e r , b u t n o t t h e f a t h e r lived t o b e 80 o r older,
only 7.7 p e r cent., o r n e a r l y 4 p e r cent, f e w e r of t h e
p e r s o n s lived t o t h e a d v a n c e d a g e of 80 o r m o r e , t h e r e
being 246 c a s e s of t h i s s o r t . T o o m u c h s t r e s s is n o t ,
however, t o be l a i d u p o n t h i s p a r e n t a l difference b e c a u s e
t h e samples a f t e r all a r e q u i t e small.
One o t h e r p o i n t i n t h i s t a b l e d e s e r v e s c o n s i d e r a t i o n .
Out of t h e 1,594 cases a s a whole, l e s s t h a n 9 p e r cent,
of the p e r s o n s lived t o t h e a d v a n c e d a g e of 80 o r m o r e .
B u t o u t of this^ n u m b e r t h e r e a r e 767, o r 48.1 p e r cent.,
n e a r l y one-half of t h e whole, w h o h a d p a r e n t s w h o lived
to 80 o r m o r e y e a r s .
A n o t h e r i n t e r e s t i n g a n d significant w a y i n w h i c h one
m a y see t h e g r e a t influence of t h e age of t h e p a r e n t s a t
d e a t h u p o n t h e l o n g e v i t y of t h e offspring, is i n d i c a t e d
in T a b l e 16, w h e r e we h a v e t h e a v e r a g e d u r a t i o n of
life of i n d i v i d u a l s w h o s e f a t h e r s a n d m o t h e r s d i e d a t
the specified a g e s .
160 BIOLOGY OF D E A T H
W e see t h a t t h e l o n g e s t a v e r a g e d u r a t i o n of life, or
e x p e c t a t i o n of life, w a s of t h a t g r o u p which h a d b o t h
m o t h e r s a n d f a t h e r s living to a g e 80 a n d o v e r . T h e
a v e r a g e d u r a t i o n of life of t h e s e p e r s o n s w a s 52.7 y e a r s .
C o n t r a s t this w i t h t h e a v e r a g e d u r a t i o n of life of t h o s e
whose p a r e n t s b o t h died u n d e r 60 y e a r s of a g e , w h e r e
TABLE 16
Showing the influence of a considerable degree of longevity in both father
and mother upon the expectation of life of the off Hairing. (After Bui).
(/n each cell of the table the open figure i$ the average duration of
life of the offspring arid the bracketedfigureis the number of
cases upon which the average in hatted).
Prominence of Ratio of
Causes of death disease. Percent. preventability.
of all deaths Per cent.
43 Alcoholism .4 85
44 Hemorrhage of lungs .1 80
45 Diseases of the thyroid body... .02 10
46 Ovarian tumor .07 0
47 Uterine tumor .1 60
48 Rheumatism .5 10
49 Gangrene of lungs .03 0
50 Anaemia, leukaemia .4 50
51 Chronic poisonings .05 70
52 Congestion of lungs .4 50
53 Ulcer of stomach .2 50
54 Carbuncle .03 50
55 Pericarditis .1 10
56 Cancer of female genital organs .6 0
57 Dysentery .5 80
58 Gastritis .65 50
59 Cholera nostras .09 50
60 Cirrhosis of liver .9 60
61 General paralysis of insane .3 75
62 Hyatid tumors of liver .002 75
63 Endocarditis .8 25
64 Locomotor ataxia .17 35
65 Diseases of veins .04 40
66 Cancer of breast .4 0
67 Diabetes .8 10
68 Biliary calouli .17 40
69 Hernia .27 70
70 Cancer not specified .9 0
71 Tumor .08 0
72 Blight's disease 5.6 40
73 Embolism and thrombosis .26 0
74 Cancer of intestines .55 0
75 Cancer of stomach and liver 1.7 0
76 Calculi of urinary tract .03 10
77 Cancer of mouth .1 0
78 Heart disease 8.1 25
79 Influenza .7 50
80 Asthma and emphysema .23 30
81 Angina pectoris .4 25
82 Apoplexy 4.4 35
83 Cancer of skin .2 0
84 Chronic bronchitis .8 30
85 Paralysis 1.0 50
86 Softening of brain .2 0
87 Diseases of arteries .83 10
88 Diseases of bladder .2 45
89 Gangrene .25 60
90 Old age 2.0 0
164 BIOLOGY OF D E A T H
I t will be seen t h a t these r a t i o s of p r e v e n t a b i l i t y a r e
n o t all 100 p e r cent. T h e y a r e n o t t h e wild overstate-
m e n t s of t h e p r o p a g a n d i s t . B u t t h e y do, r e p r e s e n t , if
t h e y could b e realized, s u b s t a n t i a l r e d u c t i o n s f r o m exist-
ing mortality rates.
TABLE 16 b
Complete expectations of life as based upon the two assumptions that deaths
are and are not prevented according to the ratios given in Table 16a
PEABSON'S WOBK
T h e m a t e r i a l u s e d b y P e a r s o n a n d h i s s t u d e n t , Miss
Beeton, who w o r k e d w i t h h i m on t h e p r o b l e m , came from
a n u m b e r of different s o u r c e s . T h e i r first s t u d y dealt
with t h r e e series f r o m which all d e a t h s r e c o r d e d as due
to accident w e r e excluded. T h e first s e r i e s included one
t h o u s a n d cases of t h e a g e s of f a t h e r s a n d sons a t
death, t h e l a t t e r b e i n g o v e r 22.5 y e a r s of age, t a k e n
T H E I N H E R I T A N C E OF DURATION 167
from F o s t e r ' s " P e e r a g e . " T h e second s e r i e s c o n s i s t e d
of a t h o u s a n d p a i r s of f a t h e r s a n d s o n s , t h e l a t t e r
dying b e y o n d t h e a g e of 20, t a k e n f r o m B u r k e ? s
" L a n d e d G e n t r y . " T h e t h i r d series c o n s i s t e d of a g e s
a t d e a t h of one t h o u s a n d p a i r s of b r o t h e r s d y i n g
beyond the a g e of 20 t a k e n f r o m t h e " P e e r a g e . " It
will be noted t h a t all t h e s e s e r i e s c o n s i d e r e d i n t h i s first
study dealt only w i t h i n h e r i t a n c e in t h e m a l e line. T h e
r e a s o n for t h i s w a s s i m p l y t h a t in such b o o k s of r e c o r d
as t h e " P e e r a g e " a n d " L a n d e d G e n t r y " sufficiently ex-
act account is n o t g i v e n of t h e d e a t h s of f e m a l e r e l a t i v e s .
I n a second s t u d y t h e m a t e r i a l w a s t a k e n f r o m t h e p e d i -
g r e e r e c o r d s of m e m b e r s of the! E n g l i s h Society of F r i e n d s
and from the F r i e n d s ' P r o v i d e n t A s s o c i a t i o n . T h i s m a -
terial included d a t a o n i n h e r i t a n c e of l o n g e v i t y in t h e
female line a n d also p r o v i d e d d a t a f o r d e a t h s of i n f a n t s ,
which w e r e l a c k i n g i n t h e e a r l i e r u s e d m a t e r i a l . T h e
investigation w a s g r o u n d e d u p o n t h a t i m p o r t a n t b r a n c h
of m o d e r n s t a t i s t i c a l calculus k n o w n a s t h e m e t h o d of
correlation. F o r each p a i r of r e l a t i v e s b e t w e e n w h o m i t
w a s desired t o s t u d y t h e i n t e n s i t y of i n h e r i t a n c e of longe-
vity a table of double e n t r y w a s f o r m e d , like t h e o n e shown
h e r e as T a b l e 17.
T h e figures i n each cell o r c o m p a r t m e n t of t h i s t a b l e
denote t h e f r e q u e n c y of occurrence of p a i r s of f a t h e r s
a n d a d u l t sons h a v i n g r e s p e c t i v e l y t h e d u r a t i o n s of life
indicated b y t h e figures i n t h e m a r g i n s . T h u s w e see,
examining t h e first line of t h e t a b l e , t h a t t h e r e w e r e 11
cases in which t h e a v e r a g e d u r a t i o n of life of t h e f a t h e r
w a s 48 y e a r s a n d t h a t of t h e a d u l t son 23 y e a r s . F a r t h e r
down a n d t o t h e r i g h t i n t h e t a b l e t h e r e w e r e 13 c a s e s i n
which the a v e r a g e d u r a t i o n of life of t h e f a t h e r a n d t h e
son w a s in e a c h c a s e 83 y e a r s . T h e s e c a s e s a r e m e n -
168 BIOLOGY OF D E A T H
t i o n e d m e r e l y a s i l l u s t r a t i o n s . T h e w h o l e t a b l e is t o be
r e a d in t h e s a m e m a n n e r .
F r o m such a t a b l e a s t h i s i t i s p o s s i b l e t o calculate,
b y well-known m a t h e m a t i c a l m e t h o d s , a single n u m e r i c a l
c o n s t a n t of s o m e w h a t u n i q u e p r o p e r t i e s k n o w n a s the
TABLE 17
Correlation table showing the correlation between father and son in respect
of duration of life
DURATION OF LIFE OF FATHER
23 28 33 38 43 48 53 58 63 68 73 78 83 88 93 98 103 Totals
23 1 1 2 5 3 11 6 7 11 9 6 12 8 2 2 86
> 28 1 6 4 5 12 15 10 13 10 7 1 1
§m 33 1 2 2 5 7 8 7 10 7 8 8 4 1 70
38 1 1 2 2 8 5 3 9 11 11 9 5 2 1 70
§ 43 1 1 5 1 5 6 11 10 10 17 5 72
* 48 1 1 2 5 5 4 6 9 12 15 5 3 68
S 53 1 3 5 7 3 2 11 11 14 10 1 1 1 70
3 58 1 3 4 5 10 8 10 5 8 9 3 2 68
% 63 2 1 3 5 1 4 8 13 9 11 11 11 5 84
I8 68 1 6 3 6 7 5 5 6 14 16 12 7 2 90
73 1 2 1 6 5 4 7 9 10 14 13 8 8 1 1 90
§ 78 1 1 2 2 4 4 4 10 5 8 9 4 3 57
« 83 1 1 5 3 1 2 3 7 10 13 3 2 2 53
§ 88 1 2 3 1 4 7 5 1 2 2 28
° 93 1 2 2 5
98 1 1 1 1 4
Totals 1 8 9 30 26 65 70 76 90 122 131 153 132 53 18 15 1 1000
coefficient o f c o r r e l a t i o n , w h i c h m e a s u r e s t h e d e g r e e of
a s s o c i a t i o n o r m u t u a l d e p e n d e n c e of t h e t w o v a r i a b l e s
i n c l u d e d i n s u c h d o u b l e e n t r y t a b l e s . T h i s coefficient
m e a s u r e s t h e a m o u n t of r e s e m b l a n c e o r a s s o c i a t i o n b e -
t w e e n characteristics of individuals o r things. I t i s
stated in t h e f o r m of a decimal which m a y take a n y value
b e t w e e n 0 a n d 1. A s t h e c o r r e l a t i o n coefficient r i s e s t o
1 w e a p p r o a c h a c o n d i t i o n o f a b s o l u t e d e p e n d e n c e of t h e
variables one u p o n t h e other. A s i t falls t o zero
w e a p p r o a c h a c o n d i t i o n of a b s o l u t e i n d e p e n d e n c e , w h e r e
the one variable h a s n o relation t o the other i n the amount
T H E I N H E R I T A N C E OF D U R A T I O N 169
o r direction of i t s v a r i a t i o n . T h e significance of a cor-
r e l a t i o n coefficient i s a l w a y s to be j u d g e d , i n a n y p a r t i c -
u l a r case, b y t h e m a g n i t u d e of a c o n s t a n t a s s o c i a t e d
w i t h i t called t h e p r o b a b l e e r r o r . A c o r r e l a t i o n coeffi-
cient m a y be r e g a r d e d as c e r t a i n l y significant w h e n it h a s
a v a l u e of 4 o r m o r e t i m e s t h a t of i t s p r o b a b l e e r r o r ,
w h i c h i s always s t a t e d a f t e r t h e coefficient w i t h a com-
b i n e d plus a n d m i n u s s i g n between t h e t w o . T h e coeffi-
cient is p r o b a b l y significant w h e n i t h a s a v a l u e of n o t
less t h a n 3 t i m e s i t s p r o b a b l e e r r o r . B y " s i g n i f i c a n t "
i n t h i s connection is m e a n t t h a t t h e coefficient p r o b a b l y
i s n o t m e r e l y a r a n d o m chance r e s u l t .
I n T a b l e 18 a r e t h e n u m e r i c a l r e s u l t s f r o m t h e first
s t u d y based u p o n t h e " P e e r a g e " a n d " L a n d e d G e n t r y . "
TABLE 18
Inheritance of duration of life in male line. Data from "Peerage" and
"Landed Gentry." (Beeton and Pearson).
Batio of
Eelatives Correlation coefficient to
coefficient its probable
error
T
X y xy "*" **r
Father ("Peerage") Son, 25 years and over .115 db .021 5.5
Father ("Landed Gentry") Son, 20 years and over .142 ±.021 6.8
Father ("Peerage") Son, 52.5 years and over .116 db .023 5.0
Father ("Landed Gentry") Son, 50 years and over .113 ±.024 4.7
Brother ("Peerage") Brother .260 ±.020 13.0
I t i s seen a t once t h a t all of t h e coefficients a r e signifi-
cant in comparison with their probable errors. The
l a s t column of t h e t a b l e gives t h e r a t i o of t h e coefficient
t o i t s p r o b a b l e e r r o r , and in t h e w o r s t c a s e t h e
coefficient is 4.7 times its p r o b a b l e e r r o r . T h e o d d s
a g a i n s t such a c o r r e l a t i o n h a v i n g a r i s e n f r o m chance
alone a r e a b o u t 655 t o 1. Odds s u c h a s t h e s e m a y
be certainly t a k e n a s d e m o n s t r a t i n g t h a t t h e r e s u l t s r e p -
170 BIOLOGY OF D E A T H
r e s e n t t r u e o r g a n i c r e l a t i o n s h i p a n d n o t m e r e chance.
All of t h e o t h e r coefficients a r e c e r t a i n l y significant, hav-
i n g r e g a r d t o t h e i r p r o b a b l e e r r o r s . F u r t h e r m o r e , they
a r e all p o s i t i v e i n sign, which implies t h a t a v a r i a t i o n in
t h e direction of i n c r e a s e d d u r a t i o n of life i n one relative
of t h e p a i r is a s s o c i a t e d w i t h a n i n c r e a s e in expectation
of life in t h e other. I t will be n o t e d t h a t the magnitude
of t h e c o r r e l a t i o n b e t w e e n b r o t h e r a n d b r o t h e r is about
twice as g r e a t a s in t h e case of c o r r e l a t i o n of f a t h e r with
son. F r o m t h i s it is p r o v i s i o n a l l y concluded t h a t the
i n t e n s i t y of t h e h e r e d i t a r y influence in r e s p e c t of duration
of life is g r e a t e r i n t h e f r a t e r n a l r e l a t i o n s h i p t h a n i n the
p a r e n t a l . I t e v i d e n t l y m a k e s n o difference, broadly
speaking, so f a r as t h e s e t w o sets of m a t e r i a l a r e con-
cerned, w h e t h e r t h e r e a r e included i n t h e c o r r e l a t i o n table
all a d u l t s o n s , w h a t e v e r t h e i r age, or only a d u l t sons over
50 y e a r s of age. T h e coefficients in b o t h cases a r e es-
sentially of t h e s a m e o r d e r of m a g n i t u d e .
P e r h a p s someone will* b e inclined t o believe t h a t the
c o r r e l a t i o n b e t w e e n f a t h e r a n d son, a n d brother! and
b r o t h e r , i n r e s p e c t of t h e d u r a t i o n of life arises as a
r e s u l t of s i m i l a r i t y of t h e e n v i r o n m e n t s to which they
a r e exposed. P e a r s o n ' s comments on t h i s p o i n t are
p e n e t r a t i n g , a n d I believe absolutely sound. H e s a y s :
There may "be some readers who will be inclined to consider that much
of the correlation of duration of life between brothers is due to there "being
a likeness of their environment, and that thus each pair of brethren is
linked together and differentiated from the general population. But it is
difficult to believe that this really affects adult brothers or a father and his
adult offspring. A man who dies between 40 and 80 can hardly be said
to have an environment more like that of his brother or father, who died
also at some such age, than like any other member of the general popula-
tion. Of course, two brothers have usually a like environment in infancy,
and their ages at death, even if they die adults, may be influenced by their
rearing. But if this be true, we ought to find a high correlation in ages
T H E I N H E R I T A N C E OF D U R A T I O N 171
at death of brethren who die as minors. As a matter of fact this correla-
tion for minor and minor is 40 to 50 per cent, less than in the case
of adult and adult. It would thus seem that identity of environment is
not the principal factor in the correlation between ages of death, for this
correlation is far less in youth than in old age.
TABLE 19
Inheritance of duration of life. Data from Quaker records.
(Beeton and Pearson)
T h e r e s u l t s r e g a r d i n g m i n o r s t o which P e a r s o n r e f e r s
a r e shown i n T a b l e 19. T h i s t a b l e gives t h e r e s u l t s of
t h e second s t u d y m a d e b y B e e t o n a n d P e a r s o n on inher-
i t a n c e of d u r a t i o n of life, based u p o n t h e r e c o r d s of the
172 BIOLOGY OF D E A T H
F r i e n d s ' Societies. I t a p p e a r s i n t h e u p p e r half of the
table that wherever a parent, father or mother, appears
w i t h a m i n o r son or d a u g h t e r t h e c o r r e l a t i o n coefficients
a r e small i n m a g n i t u d e . I n some cases t h e y a r e j u s t
b a r e l y significant i n c o m p a r i s o n w i t h t h e i r p r o b a b l e e r r o r s
as f o r example, t h e c o r r e l a t i o n of f a t h e r a n d minor
son, a n d t h a t of m o t h e r a n d m i n o r d a u g h t e r . I n the
other cases i n v o l v i n g m i n o r s t h e coefficients a r e so small
as to be insignificant. O n t h e o t h e r h a n d , i n e v e r y case
of c o r r e l a t i o n b e t w e e n p a r e n t a n d adult offspring of
either sex, t h e coefficient is 6 o r m o r e t i m e s i t s p r o b a b l e
e r r o r , a n d m u s t c e r t a i n l y b e r e g a r d e d a s significant. I t
will f u r t h e r be n o t e d t h a t t h e m a g n i t u d e of t h e coefficients
obtained f r o m t h e s e Q u a k e r r e c o r d ^ is of t h e same general
o r d e r as w a s seen i n t h e p r e v i o u s t a b l e b a s e d on t h e
" P e e r a g e " and " L a n d e d G e n t r y " material.
T h e l o w e r p a r t of t h e t a b l e gives t h e r e s u l t s for
various fraternal relationships. I n general the frater-
n a l c o r r e l a t i o n s a r e h i g h e r t h a t t h e p a r e n t a l . T h e coeffi-
cients f o r m i n o r s o r f o r m i n o r s w i t h a d u l t s a r e v e r y low
a n d in m o s t cases n o t significantly different f r o m zero.
I n four cases—namely, adult brother with minor brother;
adult s i s t e r w i t h m i n o r s i s t e r ; a d u l t b r o t h e r with m i n o r
s i s t e r ; a n d a d u l t s i s t e r w i t h m i n o r b r o t h e r — t h e coeffi-
cients a r e all n e g a t i v e i n sign, a l t h o u g h i n no one of t h e
cases is t h e coefficient significant i n c o m p a r i s o n with
its probable e r r o r . A m i n u s s i g n before a c o r r e l a t i o n
coefficient m e a n s t h a t a n i n c r e a s e i n t h e v a l u e of
one of t h e v a r i a b l e s is associated w i t h a decrease
in t h e v a l u e of t h e o t h e r . S o t h a t t h e s e n e g a t i v e
coefficients w o u l d m e a n , if t h e y w e r e significant, t h a t
t h e g r e a t e r t h e a g e a t d e a t h of a n a d u l t b r o t h e r , the lower
t h e a g e a t d e a t h of h i s m i n o r b r o t h e r o r sister. B u t t h e
coefficients a r e a c t u a l l y sensibly e q u a l t o zero. P e a r s o n
THE I N H E R I T A N C E OF DURATION 173
p o i n t s out t h a t t h e m i n u s s i g n i n the case of these correla-
tions of a d u l t w i t h m i n o r exhibits the effect of the i n h e r i -
tance of the m o r t a l i t y of y o u t h . M i n o r s d y i n g f r o m 16
t o 20 a r e associated w i t h a d u l t s d y i n g f r o m 21 t o 25.
T h a t is, m i n o r s d y i n g l a t e c o r r e s p o n d t o a d u l t s d y i n g
e a r l y . T h i s s i t u a t i o n m a y be a p e c u l i a r i t y of t h e Q u a k e r
m a t e r i a l w i t h which t h i s w o r k deals. T h e r e is u r g e n t
n e e d f o r f u r t h e r s t u d y of t h e i n h e r i t a n c e of t h e d u r a t i o n
of life on m o r e a n d b e t t e r m a t e r i a l t h a n a n y which h a s
h i t h e r t o been u s e d for t h e p u r p o s e . I h a v e u n d e r w a y
i n m y own l a b o r a t o r y a t t h e p r e s e n t t i m e a n extensive
i n v e s t i g a t i o n of t h i s kind, i n which t h e r e will be h u n d r e d s
of t h o u s a n d s of p a i r s of r e l a t i v e s in t h e individual
c o r r e l a t i o n tables i n s t e a d of t h o u s a n d s , a n d all t y p e s of
c o l l a t e r a l kinship will be r e p r e s e n t e d . B e c a u s e of t h e
m a g n i t u d e of t h e i n v e s t i g a t i o n , however, i t will be still
a n u m b e r of y e a r s b e f o r e t h e r e s u l t s will be in h a n d
f o r discussion.
T h e facts which h a v e b e e n p r e s e n t e d leave n o doubt
a s t o t h e r e a l i t y of t h e i n h e r i t a n c e f a c t o r as a p r i m e
d e t e r m i n a n t of t h e l e n g t h of t h e life s p a n .
A t t h e b e g i n n i n g i t w a s p a i n t e d o u t t h a t i t w a s on
a priori g r o u n d s h i g h l y p r o b a b l e t h a t d u r a t i o n of life
i s influenced b y b o t h h e r e d i t y a n d environment, a n d t h a t
t h e r e a l p r o b l e m is t o m e a s u r e the c o m p a r a t i v e effect of
t h e s e t w o g e n e r a l s e t s of f a c t o r s . W e h a v e seen t h a t t h e
i n t e n s i t y of i n h e r i t a n c e of d u r a t i o n of life, t a k i n g a v e r -
a g e s , is of the o r d e r i n d i c a t e d b y t h e following coefficients.
Parental correlation (adult children) r = . 1365
Fraternal correlation (adults) r = .2831
N o w we h a v e t o ask t h i s q u e s t i o n : W h a t a r e t h e v a l u e s
of p a r e n t a l a n d f r a t e r n a l c o r r e l a t i o n for c h a r a c t e r s b u t
slightly if a t all affected i n t h e i r values by the environ-
ment? Happily, P e a r s o n h a s p r o v i d e d such values in h i s
174 BIOLOGY OF D E A T H
extensive investigations on the inheritance of physical
characters in man.
I n Table 20 are given the values of the parental
correlations for the four physical characters—stature,
span, forearm length, and eye color. N o w i t is obvious
TABLE 20
Parental inheritance of physical characters in man, (Pearson)
Pair Organ Correlation
Father and Son Stature 51
Father and Son Span .45
Father and Son Forearm 42
Father and Son Eye color 55
Father and Daughter Stature 51
Father and Daughter Span .45
Father and Daughter Forearm 42
Father and Daughter Eye color 44
Mother and Son Stature .49
Mother and Son Span 46
Mother and Son Forearm 41
Mother and Son Eye color 48
Mother and Daughter Stature .51
Mother and Daughter Span 45
Mother and Daughter Forearm s 42
Mother and Daughter Eye color 51
that the differences of environmental forces impinging
upon the various members of a homogeneous group of
middle class English families (from which source the
data for these correlations were drawn) can by no pos-
sibility be great enough to affect sensibly the stature, the
arm-length, or the eye color of the adults of such families.
It would be preposterous to assert that the resemblance
between parents and offspring i n respect of eye color is
due solely, or even sensibly, to similarity of environment.
It is due to heredity and substantially nothing else.
Now the average value of the 16 parental coefficients for
the inheritance of physical characters shown i n the table is
r=.4675
T H E I N H E R I T A N C E OF DURATION 175
Table 21 shows t h e coefficients for t h e f r a t e r n a l in-
h e r i t a n c e of six p h y s i c a l c h a r a c t e r s , cephalic i n d e x (the
r a t i o of h e a d l e n g t h a n d h e a d b r e a d t h ) a n d h a i r color
h a v i n g been a d d e d t o t h o s e given i n t h e p a r e n t a l table.
A g a i n i t is seen t h a t t h e coefficients h a v e all a b o u t t h e
TABLE 21
Fraternal inheritance of physical characters in man. (Pearson)
Pair Organ Correlation
Brother and Brother Stature 51
Brother and Brother Span 55
Brother and Brother Forearm 49
Brother and Brother Eye color 52
Brother and Brother Cephalic index .49
Brother and Brother Hair color 59
Sister and Sister Stature 54
Sister and Sister Span 56
Sister and Sister Forearm 51
Sister and Sister Eye color 45
Sister and Sister Cephalic index .54
Sister and Sister Hair color 56
Brother and Sister Stature 55
Brother and Sister Span 53
Brother and Sister Forearm 44
Brother and Sister Eye color 46
Brother and Sister Cephalic index .43
Brother and Sister Hair color .56
s ^ a e values, a n d i t is as a p p a r e n t as before t h a t t h e
r e s e m b l a n c e between b r o t h e r a n d sister, f o r example, i n
eye-color, or a r m length, o r s h a p e of h e a d c a n n o t f o r
a m o m e n t , because of t h e n a t u r e of t h e c h a r a c t e r s t h e m -
selves, b e s u p p o s e d t o h a v e a r i s e n because of t h e simi-
l a r i t y of e n v i r o n m e n t . T h e a v e r a g e v a l u e of all t h e s e
f r a t e r n a l coefficients is
r=.5156
F r o m these d a t a , w i t h t h e help of a m e t h o d due t o
P e a r s o n , it is possible to d e t e r m i n e t h e p e r c e n t a g e of t h e
176 BIOLOGY OF D E A T H
death rate dependent u p o n the inherited constitution, and
the percentage not so dependent. If pN be the number
of deaths i n N cases which depend in no w a y upon the
inherited constitution of the individual, then (1-p) will
represent the chance of an individual dying because of
his inherited constitutional makeup, and ( l - p ) a will be
the chance of a pair of individuals, say t w o brothers, both
dying from causes determined by inheritance. If further
r denotes the observed correlation between individuals in
respect of duration of life, and rQ the correlation between
the same kin i n respect of such measured physical charac-
ters as those j u s t discussed, i n the determination of which
it is agreed that environment can play only a small part,
we have the following relation:
so
A5
§ *°
35
b
30
25
:§ 20
15 MOTHER
FATHER AND
AND CHILDREN
CHILDREN
10
16 26 36 56 66 76 66 and over
MALES
.7
.3
I I I
J67O 1671 /ssi 1682.
ENGLISH PRUSSIAN
FIG. 45.—Snow's results on seleotive death rate in man. The cross-hatched area may
be taken, in comparison with the small clear area at the bottom, as indicating the influence
of the selective death rate in increasing the correlations.
Our investigation substantiates for a general population the results
found by Pearson and Ploetz for more restricted populations, and disagrees
with many statements of health officers. It is with great reluctance that
T H E I N H E R I T A N C E OF DURATION 183
we point out this disagreement, and assert a doctrine which, in the present
sentiment of society, is bound to be unpopular. We have no feelings of
antagonism towards the efforts which have been made in recent years to
save infant life, but we think that the probable consequences of such actions,
so far as past experience can indicate them, should be completely under-
stood. All attempts at the reduction of mortality of infancy and childhood
should be made in the full knowledge of the facts of heredity. Everybody
knows the extreme differences in. constitutional fitness which exist in men
and women. Few intelligent people can be ignorant of the fact that this
constitutional fitness is inherited according to laws which are fairly
definitely known. At the same time marriage is just as prevalent among
those of weak stocks as among those of the vigorous, while the fertility
of the former is certainly not less than that of the latter. Thus a propor-
tion of the infants born every year must inevitably belong to the class
referred to in the report as "weaklings," and, with Pearson's results before
us, we are quite convinced that true infantile mortality (as distinct from
the mortality due to accident, neglect, etc.—no small proportion of the
whole) finds most victims from among this class. Incidentally we would
here suggest that no investigation into the causes of infant and child
mortality is complete until particulars are gathered by the medical officers
of the constitutional tendencies and physical characters of the parents.
Our work has led us to the conclusion that infant mortality does effect
a "weeding out" of the unfit; but, though we would give this conclusion
all due emphasis, we do not wish to assert that any effort, however small,
to the end of reducing this mortality is undesirable. Nobody would suggest
that the difference between the infant rates in Oxfordshire and Glamorgan-
shire (73 and 154 per 1,000 births respectively, in 1908) was wholly due
to the constitutional superiority of the inhabitants of the former county.
The "weeding-out" process is not uniform. In the mining districts of
South Wales, accident, negligence, ignorance and insanitary surroundings
account for much. By causing improvements under these heads it may
be possible to reduce the infant mortality of Glamorganshire by the sur-
vival of many who are not more unfit than are those who survive in
Oxfordshire, and the social instincts of the community insist that this
should be done.
T h i s w o r k of S n o w ' s a r o u s e d g r e a t i n t e r e s t , a n d soon
after it appearance was controverted, as it seems to me
q u i t e unsuccessfully, b y B r o w n l e e , S a l e e b y a n d o t h e r s .
H a p p i l y t h e r e s u l t s of P e a r s o n , P l o e t z a n d S n o w on
t h e selective d e a t h r a t e h a v e r e c e n t l y b e e n a c c o r d e d a
confirmation a n d e x t e n s i o n t o still a n o t h e r g r o u p of
184 BIOLOGY OF D E A T H
people—the D u t c h — i n some i n v e s t i g a t i o n s c a r r i e d out
by D r . F . S. C r a m of the P r u d e n t i a l L i f e I n s u r a n c e Com-
pany, with t h e a s s i s t a n c e of t h e d i s t i n g u i s h e d m a t h e -
matical statistician, M r . A r n e F i s h e r .
The D u t c h G o v e r n m e n t publishes a n n u a l l y d a t a which
undoubtedly f u r n i s h t h e best available m a t e r i a l now exist-
ing in the w o r l d f o r t h e p u r p o s e of d e t e r m i n i n g w h e t h e r
or not t h e r e is a positive o r n e g a t i v e c o r r e l a t i o n between
infant m o r t a l i t y a n d t h e m o r t a l i t y i n t h e immediately
subsequent y e a r s of life. F i s h e r ' s m a t h e m a t i c a l analy-
sis embraces a v e r y l a r g e b o d y of m a t e r i a l , including
n e a r l y a million a n d a half b i r t h s , a n d n e a r l y a q u a r t e r
of a million d e a t h s of m a l e s o c c u r r i n g i n t h e first five
y e a r s of life. T h e H o l l a n d d a t a m a k e i t possible to
develop life tables f o r e v e r y c o h o r t of b i r t h s a n d this
h a s been done i n the 16 c o h o r t s of m a l e s d u r i n g the y e a r s
1901-1916. T h e d a t a also m a k e i t p o s s i b l e to w o r k u p
these life t a b l e s f o r u r b a n a r e a s a n d f o r r u r a l a r e a s .
After carefully e l i m i n a t i n g s e c u l a r d i s t u r b a n c e s the
Holland m a t e r i a l a p p e a r s t o p r o v e quite conclusively for
the r u r a l d i s t r i c t s t h a t t h e r e is a definite n e g a t i v e corre-
lation, of significant m a g n i t u d e , b e t w e e n i n f a n t m o r t a l i t y
and t h e m o r t a l i t y in t h e i m m e d i a t e l y s u b s e q u e n t y e a r s of
life. The only place w h e r e p o s i t i v e c o r r e l a t i o n a p p e a r s is
i n the four l a r g e cities of t h e c o u n t r y w i t h m o r e t h a n a
hundred t h o u s a n d i n h a b i t a n t s each. F i s h e r m a k e s t h e
following p o i n t (in a l e t t e r t o t h e p r e s e n t w r i t e r ) in ex-
planation of t h e s e positive c o r r e l a t i o n s . H e s a y s :
The larger cities are better equipped with hospital and * clinical
facilities than the smaller cities and the rural districts. More money
is also spent on child welfare. Is it therefore not possible that many feeble
lives who in the course of natural circumstances would have died in the
first year of life are carried over into the second year of life by means
of medical skill? But medicine cannot always surpass nature, and it
THE INHERITANCE OF DURATION 185
might indeed be possible that among cohorts with a low mortality during
the first two years of life there will be an increase of death rate in the
following three years of life.
A l t o g e t h e r , we m a y r e g a r d t h e w e i g h t of p r e s e n t evi-
dence a s a l t o g e t h e r p r e p o n d e r a n t i n f a v o r of t h e v i e w
that the death rate of the earliest period of life is selec-
tive—eliminating the weak and leaving the strong. From
o u r p r e s e n t p o i n t of view i t a d d s a n o t h e r b r o a d class of
evidential m a t e r i a l t o t h e p r o o f of t h e p r o p o s i t i o n t h a t
i n h e r i t a n c e is one of t h e s t r o n g e s t elements, if n o t i n d e e d
t h e d o m i n a t i n g f a c t o r , i n d e t e r m i n i n g t h e d u r a t i o n of
life of h u m a n b e i n g s .
CHAPTER VII
EXPERIMENTAL S T U D I E S ON T H E DURATION
OF LIFE
cf
Fio. 46—
. Mael and female fruit fly. (J>roaophila melanog aster). (From M
banana it is sometimes called the "banana" fly. While
it lives on decaying fruit surfaces its food is mainly not
the fruit itself, but the yeast which is always growing
in such places.
The life cycle of theflyis as follows: The egg laidby
the female on some fairly dry spot on the food develops
in about 1 day into a larva. TMs larva or maggot crawls
about and feeds in the rich medium in which it finds
itself for about 3 to 4 days and then forms a pupa. From
the pupa the winged imago or adult form emerges in
about 4 or 5 days. The female generally begins to lay
eggs within the first 24 hours after she is hatched. So
188 BIOLOGY OF D E A T H
then we h a v e a b o u t 8 to 10 d a y s a s t h e m i n i m u m t i m e
d u r a t i o n of a g e n e r a t i o n . T h e whole cycle f r o m egg to
egg, a t o r d i n a r y r o o m t e m p e r a t u r e , falls w i t h i n t h i s 10-
d a y period w i t h s t r i k i n g a c c u r a c y a n d p r e c i s i o n .
T h e d u r a t i o n of life of t h e a d u l t v a r i e s i n a n o r d e r l y
m a n n e r from less t h a n 1 d a y to o v e r 90 d a y s . T h e s p a n
AOL IN OAY3
F i a . 47.—lAie tinea tor Droaophila melanooaster; showing the Burvivor» a t differeat mftm out
of 1 0 0 0 born a t t h e s a m e time.
of l i f e o f DrosophUa q u a n t i t a t i v e l y p a r a l l e l s in a n e x t r a -
o r d i n a r y w a y t h a t o f m a n , w i t h o n l y t h e difference t h a t
life's d u r a t i o n is m e a s u r e d w i t h d i f f e r e n t y a r d s t i c k s i n
is i n h u m a n society.
T h i s p a r a l l e l i s m i n t h e d u r a t i o n of life of DrosophUa
a n d m a n is well s h o w n i n F i g u r e 47, w h i c h r e p r e s e n t s a
s h i p , o r lx figures, a x e t h e o n e s p l o t t e d . T h e c u r v e s d e a l
S T U D I E S O N T H E D U R A T I O N OF L I F E 189
only with flies i n the adult or i m a g o stage, after the com-
pletion of the larval and pupal periods. T h e curve is
based upon 3,216 female and 2,620 male flies, large enough
numbers to give reliable and smooth results. W e n o t e at
once that in general the curve has the same form as the
corresponding lx curve from human mortality tables. The
most striking difference i s i n the absence f r o m the fly
curves of the h e a v y infant mortality which characterizes
the human curve. There is no specially sharp drop i n the
curve at the beginning of the life cycle, such a s h a s been
seen in the lx curve for jnan i n an earlier chapter in this
book. This m i g h t at first be thought t o be accounted
for by the fact that the curve begins after the infantile life
of the fly, but i t m u s t be remembered that the human I x
line begins at birth, and n o account is taken of the mortal-
ity in utero. R e a l l y the larval and pupal stages of the
fly correspond rather to the foetal life of a human being
than t o the infant life, so- that one may perhaps fairly take
the curves as covering comparable portions of the life s p a n
in the two cases and reach the conclusion that there i s n o t
in the fly an especially heavy incidence of mortality i n
the infant period of life, as there i s i n man. The explana-
tion of this fact i s , without doubt, that the fly when i t
emerges from the pupal stage is completely able to take
care of itself. T h e baby i s , on the contrary, i n an almost
totally helpless condition at the same relative age.
I t i s further evident that at practically all ages i n
Drosophila the number of survivors at any g i v e n age
is higher among the female than among the males. This,
it will be recalled, i s exactly the state of the case in h u m a n
mortality. The speed of the descent of the Drosophila
curve slows off in old age, j u s t a s happens in the human
life curve. T h e rate of descent of the curve in early
middle life is somewhat more rapid w i t h the flies t h a n
190 BIOLOGY OF D E A T H
i n the case of human beings, but a s will p r e s e n t l y a p p e a r
there are some strains of flies which give c u r v e s a l m o s t
identical i n this respect with the human m o r t a l i t y c u r v e s .
I n the life curves of F i g u r e 47, all different d e g r e e s of
inherited or constitutional variation in l o n g e v i t y a r e in-
cluded together. More accurate pictures of the t r u e s t a t e
of affairs will appear when w e come, as w e p r e s e n t l y
shall, t o deal with groups of individuals m o r e h o m o g e -
neous in respect of their hereditary constituents.
H a v i n g now demonstrated that the incidence of m o r -
tality is in general similar in the fly Drosophila t o "what
it i s i n man, w i t h a suitable change of unit of m e a s u r e ,
we mayproceed to examine some of the evidence r e g a r d i n g
the inheritance of duration of life in this o r g a n i s m .
The first step in such an examination i s to d e t e r m i n e
what degree of natural variation of an h e r e d i t a r y
sort exists in a general fly population in r e s p e c t o f t h i s
characteristic. I n order to do this it i s n e c e s s a r y to
isolate individual pairs, male and female, b r e e d t h e m
together and see whether, between the g r o u p s of o f f s p r i n g
so obtained, there are genetic differences in r e s p e c t of
duration of life which persist through an indefinite n u m -
ber of generations. This approaches closely t o t h e p r o -
cess called by geneticists the t e s t i n g of p u r e l i n e s . In
such a process the purpose is t o reduce t o a m i n i m u m
the genetic diversity which can possibly b e e x h i b i t e d in
the material. I n a case like the present, the w h o l e a m o u n t
of genetic variation in respect of duration of life w h i c h can
appear in the offspring of a single pair of p a r e n t s i s o n l y
that which can arise by virtue of i t s prior e x i s t e n c e i n the
parents themselves individually, and from the c o m b i n a -
tion of the germinal variation existing i n t h e t w o p a r e n t s
one with another. W e m a y call the offspring, t h r o u g h
successive generations, of a single p a i r of p a r e n t s a l i n e
STUDIES ON T H E D U R A T I O N OF L I F E 191
of descent. If, w h e n k e p t u n d e r identical e n v i r o n m e n t a l
conditions, such lines exhibit widely different a v e r a g e
d u r a t i o n s of life, a n d if t h e s e differences r e a p p e a r w i t h
constancy i n successive g e n e r a t i o n s , i t m a y be j u s t l y
concluded t h a t t h e b a s i s of t h e s e differences is h e r e d i -
t a r y in n a t u r e , since by h y p o t h e s i s t h e e n v i r o n m e n t of
all the lines is k e p t t h e s a m e . I n consequence of t h e
e n v i r o n m e n t a l e q u a l i t y , w h a t e v e r differences do a p p e a r
m u s t be i n h e r e n t l y genetic.
The manner in which these experiments are performed
m a y be of i n t e r e s t . A n e x p e r i m e n t s t a r t s b y p l a c i n g
t w o flies, b r o t h e r a n d s i s t e r , selected f r o m a stock bottle,
t o g e t h e r in a h a l f - p i n t milk bottle. A t t h e b o t t o m of t h e
bottle is a solidified, jelly-like m i x t u r e of a g a r - a g a r a n d
boiled a n d p u l p e d b a n a n a . On t h i s is sown, a s food, some
d r y y e a s t . A b i t of folded filter p a p e r i n t h e b o t t l e f u r -
n i s h e s t h e l a r v a e o p p o r t u n i t y t o p u p a t e on a d r y s u r -
face. A b o u t t e n d a y s a f t e r t h e p a i r of flies h a v e b e e n
placed i n t h i s b o t t l e , fully developed offspring i n t h e
i m a g o s t a g e begin to e m e r g e . T h e d a y before t h e s e off-
s p r i n g flies a r e d u e t o a p p e a r , t h e o r i g i n a l p a r e n t p a i r
of flies a r e r e m o v e d t o a n o t h e r bottle p r e c i s e l y like t h e
first, a n d t h e f e m a l e is allowed to l a y a n o t h e r b a t c h of
eggs over a p e r i o d of a b o u t n i n e d a y s . I n t h e o r i g i n a l
bottle t h e r e will b e offspring flies e m e r g i n g e a c h d a y ,
h a v i n g developed f r o m t h e eggs l a i d b y t h e m o t h e r on
each of t h e successive d a y s d u r i n g which she w a s i n t h e
bottle. E a c h m o r n i n g t h e offspring flies w h i c h h a v e
emerged d u r i n g t h e p r e c e d i n g t w e n t y - f o u r h o u r s a r e
t r a n s f e r r e d to a s m a l l bottle. T h i s h a s , j u s t a s t h e
l a r g e r one, food m a t e r i a l a t t h e b o t t o m a n d like t h e l a r g e r
one is closed w i t h a c o t t o n s t o p p e r . A l l of t h e offspring
flies in one of t h e s e s m a l l b o t t l e s a r e obviously of t h e
s a m e age, because t h e y w e r e b o r n a t t h e s a m e t i m e ,
192 BIOLOGY OF D E A T H
using this term " b o r n " to denote emergence from the
pupal stage as imagines. Each following day these small
bottles are inspected. Whenever a dead fly is found, it
is removed and a record m a d e i n proper form of the
fact that its death occurred, and its age and sex are noted.
Finally, when all the flies in a given small bottle have
died, that bottle i s discarded, as the record of the duration
1.000
900
600 \ ,
TOO
6O0
\ \ \
\
\
ZOO
00
0 \
30 3642 46 54 60 66 73 84 90
A6C tN Q4YS
F I G . 4 8 . — L i f e l i n e s for different i n b r e d lines of d e s c e n t i n Drosophila.
o f l i f e o f e a c h i n d i v i d u a l i s t h e n c o m p l e t e . A l l t h e
b o t t l e s a r e k e p t i n e l e c t r i c i n c u b a t o r s a t a c o n s t a n t
t e m p e r a t u r e o f 2 5 ° C , t h e s m a l l b o t t l e s b e i n g p a c k e d f o r
c o n v e n i e n c e i n w i r e b a s k e t s . A l l h a v e t h e s a m e f o o d
m a t e r i a l , b o t h i n q u a l i t y a n d q u a n t i t y , s o t h a t t h e e n v i -
r o n m e n t a l c o n d i t i o n s s u r r o u n d i n g t h e s e flies d u r i n g t h e i r
l i f e m a y b e r e g a r d e d a s s u b s t a n t i a l l y c o n s t a n t a n d u n i -
f o r m f o r a l l .
F i g u r e 4 8 s h o w s t h e s u r v i v a l f r e q u e n c y , o r lx l i n e
o f a l i f e t a b l e , f o r s i x d i f f e r e n t l i n e s o f D r o s o p h i l a , w h i c h
h a v e b e e n b r e d i n m y l a b o r a t o r y . E a c h l i n e r e p r e s e n t s
STUDIES ON T H E DURATION OF LIFE 193
the s u r v i v a l d i s t r i b u t i o n of t h e offspring of a single
brother and sister p a i r mated together. I n forming a
line a b r o t h e r a n d s i s t e r a r e t a k e n a s t h e i n i t i a l s t a r t
because by so d o i n g t h e a m o u n t of genetic v a r i a t i o n p r e s -
ent in t h e line a t t h e b e g i n n i n g is r e d u c e d t o t h e lowest
possible m i n i m u m . I t should be said t h a t i n all of t h e
curves i n F i g u r e 48, b o t h m a l e a n d f e m a l e offspring a r e
lumped t o g e t h e r . T h i s is justifiable f o r i l l u s t r a t i v e p u r -
poses because of t h e s m a l l difference i n t h e e x p e c t a t i o n
of life a t a n y a g e b e t w e e n t h e sexes. T h e line of descent,
No. 55, figured a t t h e t o p of t h e d i a g r a m , gives a n lx
line e x t r a o r d i n a r i l y like t h a t f o r m a n , w i t h t h e exception
of the omission of t h e s h a r p d r o p d u e to i n f a n t i l e m o r -
tality a t the b e g i n n i n g o£ t h e c u r v e . T h e e x t r e m e d u r a -
tion of life i n t h i s line w a s 81 d a y s , r e a c h e d b y a f e m a l e
fly. T h e h line d r o p s off v e r y slowly u n t i l a g e 36 d a y s .
F r o m t h a t time on, t h e descent is m o r e r a p i d u n t i l 72 d a y s
of a g e a r e r e a c h e d w h e n i t slows u p a g a i n . L i n e s 50, 60,
and 58 show h c u r v e s all d e s c e n d i n g m o r e r a p i d l y i n t h e
early p a r t of t h e life cycle t h a n t h a t f o r line 55, a l t h o u g h
the m a x i m u m d e g r e e of l o n g e v i t y a t t a i n e d is a b o u t t h e
same in all of t h e f o u r first c u r v e s . T h e g e n e r a l s h a p e
of the lx c u r v e s c h a n g e s h o w e v e r , a s i s c l e a r l y seen if
we c o n t r a s t line 55 w i t h l i n e 58. T h e f o r m e r is concave
to t h e base t h r o u g h n e a r l y t h e w h o l e of i t s c o u r s e , w h e r e a s
the lx curve f o r line 58 is convex t o t h e b a s e p r a c t i c a l l y
t h r o u g h o u t its c o u r s e . "While, a s i s c l e a r f r o m t h e dia-
g r a m , t h e m a x i m u m longevity a t t a i n e d is a b o u t t h e s a m e
for all of t h e s e u p p e r f o u r lines, i t is equally obvious
t h a t the m e a n d u r a t i o n of life exhibited b y t h e lines falls
off a s we go down t h e d i a g r a m . T h e s a m e p r o c e s s , which
is in o p e r a t i o n between lines 55 a n d 58, i s c o n t i n u e d i n
a n even m o r e m a r k e d d e g r e e i n lines 61 a n d 64. H e r e
n o t only is t h e d e s c e n t m o r e r a p i d in t h e e a r l y p a r t of t h e
194 BIOLOGY OF D E A T H
lx c u r v e , b u t t h e m a x i m u m d e g r e e of longevity a t t a i n e d
is m u c h smaller, a m o u n t i n g to a b o u t half of t h a t a t t a i n e d
in t h e o t h e r f o u r lines. B o t h lines 61 a n d 64 t e n d t o show
in g e n e r a l a c u r v e convex to t h e b a s e , especially in the
l a t t e r half of t h e i r c o u r s e .
Since each of t h e s e l i n e s of d e s c e n t continues t o show
t h r o u g h successive g e n e r a t i o n s , f o r a n indefinite time,
t h e same t y p e s of m o r t a l i t y c u r v e s a n d a p p r o x i m a t e l y
t h e s a m e a v e r a g e d u r a t i o n s of life, i t m a y safely be con-
cluded t h a t t h e r e a r e well m a r k e d h e r e d i t a r y differences
in different s t r a i n s of t h e s a m e species of Drosophila in
r e s p e c t of d u r a t i o n of life. P a s s i n g f r o m t h e t o p to the
b o t t o m of t h e d i a g r a m t h e a v e r a g e e x p e c t a t i o n of life is
reduced by about two-thirds in these representative
curves. F o r p u r p o s e s of e x p e r i m e n t a t i o n , each one of
these lines of d e s c e n t becomes c o m p a r a b l e t o a chemical
r e a g e n t . T h e y h a v e s t a n d a r d d u r a t i o n s of life, each
p e c u l i a r to its own line a n d d e t e r m i n e d b y t h e h e r e d i t a r y
c o n s t i t u t i o n of t h e i n d i v i d u a l i n r e s p e c t of t h i s charac-
t e r . W e m a y , w i t h e n t i r e justification, speak of the
flies of line 64 a s h e r e d i t a r i l y short-lived, a n d those of
line 55 a s h e r e d i t a r i l y long-lived.
H a v i n g e s t a b l i s h e d so much, t h e n e x t s t e p in the analy-
sis of t h e m o d e of i n h e r i t a n c e of t h i s c h a r a c t e r is ob-
viously t o p e r f o r m a M e n d e l i a n e x p e r i m e n t by c r o s s i n g
a n h e r e d i t a r i l y s h o r t - l i v e d line w i t h a n h e r e d i t a r i l y long-
lived line, a n d follow t h r o u g h i n t h e p r o g e n y of succes-
sive g e n e r a t i o n s t h e d u r a t i o n of life. If t h e c h a r a c t e r
follows t h e o r d i n a r y c o u r s e of M e n d e l i a n i n h e r i t a n c e , we
should expect t o g e t i n t h e second offspring g e n e r a t i o n a
s e g r e g a t i o n of different t y p e s of flies in r e s p e c t of t h e i r
d u r a t i o n of life.
F i g u r e 49 shows t h e r e s u l t of such Mendelian experi-
STUDIES ON T H E DURATION OF LIFE 195
m e n t p e r f o r m e d o n a l a r g e scale. I n t h e second line f r o m
t l i e t o p of t h e d i a g r a m , labeled " T y p e I L," w e see t h e
m o r t a l i t y c u r v e f o r a n h e r e d i t a r i l y long-lived p u r e s t r a i n
o f i n d i v i d u a l s . A t t h e b o t t o m of t h e d i a g r a m t h e ' ' T y p e I V
lx" line gives t h e m o r t a l i t y c u r v e for one of o u r h e r e d i t a -
r i l y short-lived s t r a i n s . I n d i v i d u a l s of T y p e I a n d T y p e I V
1.000
300 [ X
SCO \
\ \
7O0
\ X
6O0
\
\
300
400 V \ \
soo A
• \
zoo
\
too N\ \
0
c 6 Z l9 &t 30 Ji5 4Z 46 J4 6O 66 72
AQC IN DAYS
F I G . 49.—^Life lines s h o w i n g t h e result of M e n d e l i a n e x p e r i m e n t s o n t h e d u r a t i o n of life in
Drosophila. Explanation in text.
w e r e m a t e d t o g e t h e r . T h e r e s u l t i n t h e first o f f s p r i n g
h y b r i d g e n e r a t i o n i s s h o w n b y t h e l i n e a t t h e t o p o f d i a -
i i
g r a m m a r k e d F 1 l x . " T h e F x d e n o t e s t h a t t h i s i s t h e m o r -
t a l i t y c u r v e o f t h e first filial g e n e r a t i o n f r o m t h e c r o s s .
I t i s a t o n c e o b v i o u s t h a t t h e s e first g e n e r a t i o n h y b r i d s
h a v e a g r e a t e r e x p e c t a t i o n o f l i f e a t p r a c t i c a l l y a l l a g e s
t h a n d o e i t h e r o f t h e p a r e n t s t r a i n s m a t e d t o g e t h e r t o
p r o d u c e t h e h y b r i d s . T h e r e s u l t i s e x a c t l y c o m p a r a b l e
t o t h a t w h i c h h a s f o r s o m e t i m e b e e n k n o w n t o o c c u r
i n p l a n t s , f r o m t h e r e s e a r c h e s p a r t i c u l a r l y o f E a s t a n d
o t h e r s w i t h m a i z e . E a s t a n d h i s s t u d e n t s h a v e w o r k e d
196 BIOLOGY OF D E A T H
out v e r y t h o r o u g h l y t h e cause of t h i s i n c r e a s e d v i g o r of
the first h y b r i d g e n e r a t i o n a n d show t h a t it is directly
due to t h e m i n g l i n g of different g e r m p l a s m s .
The a v e r a g e d u r a t i o n of life of t h e T y p e I original
p a r e n t stock is 44.2 =t .4 d a y s . T h e a v e r a g e d u r a t i o n of
life of t h e short-lived T y p e I V flies is 14.1 zfc -2 d a y s , or
only a b o u t one t h i r d a s g r e a t as t h a t of t h e o t h e r stock.
The a v e r a g e d u r a t i o n of life of t h e first h y b r i d genera-
tion shown i n t h e F j lx line i s 51.5 ± .5 d a y s . So t h a t
t h e r e is a n i n c r e a s e i n a v e r a g e d u r a t i o n of life in t h e
first h y b r i d g e n e r a t i o n , o v e r t h a t of t h e long-lived p a r e n t ,
of a p p r o x i m a t e l y 7 d a y s . I n e s t i m a t i n g t h e significance
of this, one should r e m e m b e r t h a t a d a y i n t h e life of a
fly c o r r e s p o n d s , a s h a s a l r e a d y b e e n p o i n t e d out, almost
exactly t o a y e a r i n t h e life of a m a n .
W h e n i n d i v i d u a l s of t h e first h y b r i d g e n e r a t i o n a r e
mated t o g e t h e r t o g e t t h e second, o r F 2 h y b r i d g e n e r a t i o n
we get a g r o u p of flies which, if taken all together, give
the m o r t a l i t y c u r v e s h o w n i n t h e line a t a b o u t t h e middle
of the d i a g r a m , labelled " A l l F 2 lx." I t , however, tells
us little a b o u t t h e m o d e of i n h e r i t a n c e of t h e c h a r a c t e r
if we c o n s i d e r all t h e i n d i v i d u a l s of t h e second h y b r i d
generation t o g e t h e r , because r e a l l y t h e r e a r e s e v e r a l
kinds of flies p r e s e n t i n t h i s second h y b r i d g e n e r a t i o n .
T h e r e a r e s h a r p l y s e p a r a t e d g r o u p s of long-lived flies a n d
of short-lived flies. T h e s e h a v e b e e n l u m p e d t o g e t h e r t o
give t h e " A l l F 2 lx" line. If w e c o n s i d e r s e p a r a t e l y t h e
long-lived second g e n e r a t i o n g r o u p a n d t h e short-lived
second g e n e r a t i o n g r o u p w e g e t t h e r e s u l t s shown i n t h e
two lines labelled " L o n g - l i v e d F 2 S e g r e g a t e s lX9" a n d
" S h o r t - l i v e d F 2 S e g r e g a t e s h." I t will be noted t h a t t h e
long-lived F 2 s e g r e g a t e s h a v e a m o r t a l i t y c u r v e which al-
m o s t e x a c t l y coincides w i t h t h a t of t h e o r i g i n a l p a x e n t T y p e
I stock. I n o t h e r w o r d s , i n t h e second g e n e r a t i o n after
STUDIES ON T H E DURATION OF LIFE 197
t h e cross of t h e long-lived a n d s h o r t - l i v e d t y p e s , a g r o u p
of animals a p p e a r s h a v i n g a l m o s t i d e n t i c a l l y t h e s a m e
f o r m of m o r t a l i t y c u r v e as t h a t of one of t h e o r i g i n a l
p a r e n t s in t h e c r o s s . T h e m e a n d u r a t i o n of life of this
long-lived second g e n e r a t i o n g r o u p is 43.3 db .4 d a y s ,
while t h a t of t h e o r i g i n a l long-lived stock w a s 44.2 =b -4
d a y s . T h e short-lived F 2 s e g r e g a t e s , s h o w n a t t h e b o t t o m
of t h e d i a g r a m , g i v e a m o r t a l i t y c u r v e e s s e n t i a l l y like
t h a t of t h e o r i g i n a l s h o r t - l i v e d p a r e n t s t r a i n . T h e two
curves wind i n a n d a b o u t each o t h e r , t h e F 2 flies s h o w i n g
a m o r e r a p i d d e s c e n t i n t h e first half of t h e c u r v e a n d a
slower descent i n t h e l a t t e r half. I n g e n e r a l , however,
t h e two a r e v e r y c l e a r l y of t h e s a m e f o r m . T h e a v e r -
age d u r a t i o n of life of t h e s e short-lived second g e n e r a t i o n
s e g r e g a t e s is 14.6 db .6 d a y s . T h i s , i t will b e recalled,
is a l m o s t identically t h e s a m e a v e r a g e d u r a t i o n of life
as the o r i g i n a l p a r e n t T y p e I V gave, w h i c h w a s 14.1 ±
.2 d a y s .
I t m a y occur t o one t o w o n d e r h o w i t is possible to
pick out the long-lived a n d short-lived s e g r e g a t e s i n t h e
second g e n e r a t i o n . T h i s i s done b y v i r t u e of t h e c o r r e -
lation of t h e d u r a t i o n of life of these flies w i t h c e r t a i n
e x t e r n a l bodily c h a r a c t e r s , p a r t i c u l a r l y t h e f o r m of
the wings, so t h a t t h i s a r r a n g e m e n t of t h e m a t e r i a l c a n
be m a d e w i t h p e r f e c t e a s e a n d c e r t a i n t y .
T h e s e r e s u l t s s h o w i n a c l e a r m a n n e r t h a t d u r a t i o n of
life, in DrosopMla a t least, is i n h e r i t e d e s s e n t i a l l y in
accordance w i t h M e n d e l i a n l a w s , t h u s fitting i n w i t h a
wide r a n g e of o t h e r p h y s i c a l c h a r a c t e r s of t h e a n i m a l
which have, b e e n t h o r o u g h l y s t u d i e d p a r t i c u l a r l y b y
M o r g a n a n d h i s s t u d e n t s . S u c h r e s u l t s as t h e s e j u s t
shown c o n s t i t u t e t h e b e s t k i n d of proof of t h e e s s e n t i a l
p o i n t which w e a r e e x a m i n i n g — n a m e l y , t h e f a c t t h a t
198 BIOLOGY OF D E A T H
d u r a t i o n of life is a n o r m a l l y i n h e r i t e d c h a r a c t e r . I d o
n o t w i s h a t this time to go into a n y d i s c u s s i o n of t h e
details of t h e Mendelian m e c h a n i s m f o r t h i s c h a r a c t e r ,
i n the first place, because it is too c o m p l i c a t e d a n d t e c h -
nical a m a t t e r f o r discussion h e r e * a n d , i n t h e s e c o n d
place, because t h e i n v e s t i g a t i o n s a r e f a r f r o m b e i n g c o m -
pleted yet. I wish h e r e a n d n o w m e r e l y t o p r e s e n t t h e
d e m o n s t r a t i o n of t h e b r o a d g e n e r a l f a c t t h a t d u r a t i o n
of life is i n h e r i t e d i n a n o r m a l M e n d e l i a n m a n n e r i n
these fly p o p u l a t i o n s . T h e first evidence t h a t this w a s
t h e case came f r o m some work of D r . E . R. H y d e w i t h
Drosophila some y e a r s ago. T h e n u m b e r s involved i n
his experiment, however, were m u c h s m a l l e r t h a n t h o s e
of the p r e s e n t e x p e r i m e n t s , a n d t h e p r e l i m i n a r y d e m o n -
s t r a t i o n of t h e existence of p u r e s t r a i n s r e l a t i v e to d u r a -
tion of life i n Drosophila w a s n o t u n d e r t a k e n by h i m .
Hyde's results and those here presented are e n t i r e l y
i n accord.
W i t h the evidence which h a s now been p r e s e n t e d r e -
g a r d i n g the i n h e r i t a n c e of life in m a n a n d in Drosophila
we m a y let t h a t p h a s e of the subject r e s t . T h e e v i d e n c e
is conclusive of the b r o a d fact, beyond a n y q u e s t i o n I
think, coining as it does f r o m such widely different t y p e s
of life, and a r r i v e d a t b y such totally different m e t h o d s a s
t h e statistical, on t h e one hand, a n d t h e e x p e r i m e n t a l , o n
t h e other. W e m a y safely conclude t h a t t h e p r i m a r y a g e n t
concerned i n t h e w i n d i n g u p of t h e v i t a l clock, a n d b y
t h e winding d e t e r m i n i n g p r i m a r i l y a n d f u n d a m e n t a l l y
how long i t shall r u n , i s heredity. T h e b e s t i n s u r a n c e
of longevity is beyond question a c a r e f u l selection o f
one's p a r e n t s and g r a n d p a r e n t s .
* Full technical details and all the numerical data regarding th«*»e and
other Droaophila experiments referred to in this book will shortly be
published elsewhere.
S T U D I E S ON T H E D U R A T I O N O F L I F E 199
BACTERIA AN"D DUBATION OF LIFE IN DBOSOPHILA
B u t clocks m a y be s t o p p e d i n o t h e r w a y s t h a n by
r u n n i n g d o w n . I t will be w o r t h while to c o n s i d e r w i t h
s o m e c a r e a c o n s i d e r a b l e m a s s of m o s t i n t e r e s t i n g , a n d
i n s o m e r e s p e c t s even s t a r t l i n g , e x p e r i m e n t a l d a t a , r e -
g a r d i n g v a r i o u s w a y s in which l o n g e v i t y m a y be influenced
b y e x t e r n a l a g e n t s . Since w e h a v e j u s t been c o n s i d e r i n g
JDrosophila i t m a y be well to c o n s i d e r t h e e x p e r i m e n t a l
evidence r e g a r d i n g t h a t f o r m first. I t i s a n obviously
w e l l - k n o w n f a c t t h a t b a c t e r i a a r e r e s p o n s i b l e i n all h i g h e r
o r g a n i s m s for much o r g a n breakdown and consequent
d e a t h . A n infection of s o m e p a r t i c u l a r o r g a n o r o r g a n
s y s t e m occurs, a n d t h e d i s t u r b a n c e of t h e b a l a n c e of t h e
w h o l e s o b r o u g h t a b o u t finally r e s u l t s i n d e a t h . B u t is
i t n o t p o s s i b l e t h a t w e o v e r r a t e t h e i m p o r t a n c e of b a c t e r -
ial invasion in determining, in general and in the broad-
e s t s e n s e , t h e a v e r a g e d u r a t i o n of life? M a y i t n o t be
t h a t w h e n a n o r g a n s y s t e m b r e a k s down, u n d e r s t r e s s
of b a c t e r i a l t o x i n s , i t is i n p a r t a t least, p e r h a p s
primarily, because for internal organic reasons the resis-
t a n c e of t h a t o r g a n s y s t e m to b a c t e r i a l i n v a s i o n h a s n o r -
m a l l y a n d n a t u r a l l y r e a c h e d such a low p o i n t t h a t i t s
defenses are no logger adequate? All higher animals
live c o n s t a n t l y in a n e n v i r o n m e n t f a r f r o m s t e r i l e . O u r
m o u t h s a n d t h r o a t s h a r b o r p n e u m o n i a g e r m s m u c h of
t h e t i m e , b u t w e d o n o t all o r a l w a y s h a v e p n e u m o n i a .
A g a i n i t m a y f a i r l y be e s t i m a t e d t h a t of all p e r s o n s who
a t t a i n t h e a g e of 35, p r o b a b l y a t l e a s t 95 p e r cent, h a v e
a t s o m e t i m e o r o t h e r b e e n infected w i t h t h e t u b e r c l e
bacillus, y e t f e w e r t h a n o n e in t e n b r e a k d o w n w i t h
active t u b e r c u l o s i s .
W h a t p l a i n l y is n e e d e d in o r d e r t o a r r i v e a t a j u s t
e s t i m a t e of t h e r e l a t i v e influence of b a c t e r i a a n d t h e i r
200 BIOLOGY OF D E A T H
toxins i n d e t e r m i n i n g t h e a v e r a g e d u r a t i o n of life is an
e x p e r i m e n t a l i n q u i r y i n t o t h e effect of a bacteria-free,
sterile m o d e of life. Metchnikoff h a s s t u r d i l y advocated
t h e view t h a t d e a t h i n g e n e r a l is a r e s u l t of bacterial
intoxication. N o w a b a c t e r i a - f r e e existence is n o t pos-
sible f o r m a n . B u t i t i s possible f o r c e r t a i n insects,
as w a s first d e m o n s t r a t e d b y B o g d a n o w , a n d l a t e r con-
firmed b y D e l c o u r t a n d Guyenot. If one carefully washes
either t h e egg o r t h e p u p a of Drosophila for 10 m i n u t e s
i n a s t r o n g a n t i s e p t i c solution, s a y 85 p e r cent, alcohol,
h e will kill a n y g e r m which m a y be u p o n t h e surface. If
t h e b a c t e r i a - f r e e e g g o r p u p a is t h e n p u t i n t o a sterile
receptacle, c o n t a i n i n g only s t e r i l e food m a t e r i a l a n d a
p u r e c u l t u r e of y e a s t , d e v e l o p m e n t will occur a n d p r e -
sently a n a d u l t i m a g o will e m e r g e . A d u l t flies r a i s e d in
this w a y a r e s t e r i l e . T h e y h a v e n o b a c t e r i a i n s i d e or
out. N o r m a l h e a l t h y p r o t o p l a s m i s n o r m a l l y sterile, so
w h a t is inside t h e fly i s b o u n d to be sterile on t h a t account,
a n d b y t h e u s e of t h e a n t i s e p t i c solution w h a t b a c t e r i a
w e r e on t h e o u t s i d e h a v e b e e n killed.
T h e p r o b l e m n o w i s , h o w l o n g on the a v e r a g e do such
sterile s p e c i m e n s of Drosophila live in c o m p a r i s o n w i t h
t h e o r d i n a r y fly, w h i c h is t h r o u g h o u t i t s a d u l t life as
much b e s e t b y b a c t e r i a r e l a t i v e l y a s is m a n himself, i t
being p r e m i s e d t h a t i n b o t h cases a n a b u n d a n c e of p r o p -
e r food is f u r n i s h e d a n d t h a t i n g e n e r a l t h e environ-
m e n t a l conditions, o t h e r t h a n bacterial, a r e m a d e t h e same
f o r t h e t w o sets? F o r t u n a t e l y , t h e r e a r e some d a t a to
t h r o w l i g h t u p o n t h i s q u e s t i o n f r o m t h e e x p e r i m e n t s of
L o e b a n d his a s s o c i a t e N o r t h r o p on the d u r a t i o n of life
i n this f o r m , t a k e n i n connection w i t h e x p e r i m e n t s i n t h e
writer's laboratory.
L o e b a n d N o r t h r o p show t h a t a s a m p l e of 70 flies, of
t h e Drosophila w i t h which t h e y w o r k e d , which w e r e
STUDIES ON T H E DURATION OF LIFE 201
p r o v e d by the m o s t careful a n d critical of t e s t s t o h a v e
r e m a i n e d entirely f r e e of b a c t e r i a l c o n t a m i n a t i o n t h r o u g h -
o u t t h e i r lives, exhibited, w h e n g r o w n a t a c o n s t a n t t e m -
p e r a t u r e of 25° C. an average duration of life of 28.5
days. I n our e x p e r i m e n t s 2,620 m a l e flies, of all s t r a i n s
of Drosophila i n o u r c u l t u r e s t a k e n t o g e t h e r , t h u s giv-
i n g a f a i r r a n d o m s a m p l e of genetically t h e whole Droso-
phila p o p u l a t i o n , g a v e a n a v e r a g e d u r a t i o n of life a t t h e
s a m e c o n s t a n t t e m p e r a t u r e of 25° C. of 31.3 =±=.3 d a y s ,
a n d 3,216 females u n d e r t h e s a m e t e m p e r a t u r e lived a n
a v e r a g e of 33.0 ± . 2 d a y s T h e s e w e r e all n o n - s t e r i l e
flies, subject t o all t h e b a c t e r i a l c o n t a m i n a t i o n i n c i d e n t
t o t h e i r n o r m a l l a b o r a t o r y e n v i r o n m e n t , w h i c h we h a v e
s e e n t o be a d e c a y i n g g e r m - l a d e n m a s s of b a n a n a p u l p a n d
a g a r . I t i s t h o u g h t t o b e f a i r e r t o c o m p a r e a s a m p l e of
a g e n e r a l p o p u l a t i o n w i t h t h e L o e b a n d N o r t h r o p figures
r a t h e r t h a n a p u r e s t r a i n b e c a u s e p r o b a b l y t h e i r Droso-
phila m a t e r i a l w a s f a r f r o m h o m o z y g o u s i n r e s p e c t of
t h e genes f o r d u r a t i o n of life.
T h e detailed c o m p a r i s o n s a r e s h o w n i n T a b l e 23.
TABLE 23
Average duration of ; oj Drosophila in the imago stage at 85° C.
too
90
60
9 7 I 17 6 10 2 3 5 4 12 14 15 18 II 13 B 20 n m ET PARIS
i
CLASSES OF
ARZ0NDIS5EMWS
ARRONDISSEMOrrS
F I G . 50.—Distribution of p o v e r t y i n Paris (1911-13) a s i n d i c a t e d b y exemption from personal
p r o p e r t y t a x . (After H e r s c h ) .
that e n v i r o n m e n t a l c o n d i t i o n s i n g e n e r a l can, a n d u n d e r
sojne c i r c u m s t a n c e s , d o e x e r t a m a r k e d influence u p o n
e x p e c t a t i o n of life. A r e c e n t s t u d y of g r e a t interest a n d
s u g g e s t i v e n e s s , if p e r h a p s s o m e l a c k of critical s o u n d -
be cited i n this c o n n e c t i o n . H e r s c h b e c a m e i n t e r e s t e d
i n the r e l a t i o n of p o v e r t y to m o r t a l i t y . H e g a t h e r e d
STUDIES ON T H E DURATION OF LIFE 203
d a t a f r o m the 20 a r r o n d i s s e m e n t s of t h e city of P a r i s i n
respect of the following p o i n t s , a m o n g o t h e r s :
a. Percentage of families not paying a personal property tax.
b. Death rate per 1000 from all causes.
c. Stillbirths per 1000 living births.
F i g u r e 50 shows i n t h e black t h e p e r c e n t a g e of f a m -
ilies too p o o r t o h a v e a n y p e r s o n a l p r o p e r t y t a x a s s e s s e d ,
first f o r each a r r o n d i s s e m e n t s e p a r a t e l y , t h e n a t t h e
MORTALITY IN PARIS 1911 - 1913
\6 I 7 17 6 ZlO3GIZ4SIII5i4- 19 20 13
ARR0ND1SSCMENTS
FIG. 51.—Death rates in Paris (1911-13) from all causes. (After Hersch).
o n Ufcxe " b a s i s of t h e B a v a r i a n d a t a of G r o t h a n d H a h i i .
206 BIOLOGY OF D E A T H
Now this is a statistically insignificant n e t c o r r e l a t i o n
being less even t h a n 3 times its p r o b a b l e e r r o r . I t m e a n
that, when the b i r t h r a t e a n d artificial f e e d i n g r a t e a r
held constant, differences i n t h e i n f a n t d e a t h r a t e a r
n o t sufficiently influenced o r d e t e r m i n e d b y d i f f e r e n c e
in the p o v e r t y r a t e to lead to a coefficient of c o r r e l a t i o
significantly different f r o m z e r o , so f a r as B a v a r i a
populations a r e indicative.
This r e s u l t i s f u r t h e r confirmed b y a n a n a l y s i s w h i c
Greenwood a n d B r o w n m a d e of H e r o n ' s L o n d o n m a t *
rial, showing t h a t in t h a t case
of population (1911)
Death-rate
Domestic servants
from all causes, fr<>m
1911-13 tub ere ulosis,
1918-19 1911-lc 1911-13
Legitimate
Standard-
Standard-
Illegiti-
Crude
Crude
Total
mate
ized
ized
Kensington 16.67 13.7 13.6 83 221 112 1.32 1.32
Hampstead 16.40 10.4 11.0 64 236 72 0.81 0.80
Westminster . 15.17 12 6 13 3 77 235 94 1.49 1.42
Chelsea 14.96 14.7 14.0 78 155 91 1.64 1.61
Marylebone 12.98 14.3 14.6 79 250 98 1.70 1.66
Paddington 10.42 13.3 13.2 88 237 109 1.33 1.31
Group I 14.38 13.2 13.4 80 228 100 1.39 1.36
City 6.46 14 0 14.6 122 278 97 1.95 1.84
Lewisham 5.71 11.3 11.1 51 294 84 1.09 1.01
Wandsworth 5.67 11.5 11.6 72 240 96 1.20 1.18
Stoke Newington.... 4.98 13.0 12.4 71 231 85 1.30 1.28
Holborn 4.38 15.1 15.2 99 267 102 2.30 2.17
Greenwich 4.10 13.9 13.7 93 349 102 1.60 1.59
Group II 5.34 12.2 12.1 73 272 94 1.32 1.30
Fulham 3.52 13.6 14.1 85 222 105 1.73 1.69
Haramersmith 3.30 14.5 14.3 91 207 114 1.62 1.58
Lambeth 3.18 14.3 14.0 82 217 105 1.71 1.68
St. Pancras 3.12 15.1 15.1 81 226 98 1.91 1.85
Hackney.. .. 2.95 13.5 13.6 83 355 100 1.68 1.67
Woolwich 2.81 12.6 12.9 88 220 84 1.67 1.65
Camberwell. 2.68 13.8 13.6 83 268 99 1.61 1.60
Deptford 2.64 15.0 14.8 87 176 117 1.73 1.70
Battersea 2.62 13.6 13.7 74 276 107 1.56 1.53
Islington 2.48 14.9 14.5 88 258 107 1.69 1.66
Group III 2.90 14.1 14.0 84 243 103 1.70 1.67
SteDnev 1.33 15.8 16.5 90 314 121 2.15 2.12
Finsbury 1.24 18.6 18.4 91 229 137 2.47 2.45
Southwark 1.23 17.5 17.6 101 225 122 2.23 2.17
Poplar 1.18 17.2 17.0 92 216 125 1.88 1.86
Bermondsey 0.97 17.8 17.8 105 360 133 2.35 2.31
Shoreditch 0.91 18.9 19.5 124 255 150 2.47 2.46
Bethnal Green 0.77 16.4 17.1 101 263 123 2.21 2.21
Group IV 1.13 17.1 17.4 99 260 128 2.21 2.18
County of London... 4.74 14.4 14.4 86 247 109 1.71 1.68
208 BIOLOGY OF D E A T H
richest group. Indeed, the difference between the first three London
groups is alight, significant excess only being apparent for the poorent
group. And whereas the excess of mortality of the poorest over the
richest group in Paris ia 104 per cent., in London it is only 30 per cent."
H e t h e n e x a m i n e s t h e q u e s t i o n a s t o w h e t h e r t h e dis-
c r e p a n c i e s m a y be d u e t o differences i n t h e m e t h o d of con-
s t r u c t i o n of t h e t w o s e t s of m o r t a l i t y figures a n d c o n c l u d e s :
"That the remarkable contrast in experience between the two cities
cannot be explained, except possibly in a very minor degree, by any
differences of method in compilation of the statistics compared/'
S t e v e n s o n t h e n g o e s on to, t h e discussion of i n f a n t
mortality and s a y s :
"The conclusion just arrived at applies still more to infant than to
total mortality, for, in its case, the contrast between rich and poor quartern
of Paris assumes dimensions which, in the light of London exptmtttiee, «tH*m
quite fantastic.1'
Regarding mortality from tuberculosis the London
experience a g a i n fails t o a g r e e w i t h t h e P a r i s e x p e r i e n c e ,
a n d H e r s c h ' s conclusions f r o m t h e d a t a of t h e l a t t e r city
would be a b s u r d if a p p l i e d to t h e f o r m e r .
H o w a r e t h e s e r e s u l t s to be reconciled w i t h t h e p r e -
vious finding t h a t h e r e d i t y is a p r i m a r y f a c t o r i n the
d e t e r m i n a t i o n of d u r a t i o n of life of Drosophilaf We
h a v e h e r e , on first i m p r e s s i o n a t least, a n excellent exam-
ple of w h a t one a l w a y s e n c o u n t e r s in critical genetic
i n v e s t i g a t i o n s : t h e c o m p l e m e n t a r y r e l a t i o n s of h e r e d i t y
a n d e n v i r o n m e n t . I n o u r e x p e r i m e n t s a g e n e r a l mixed
p o p u l a t i o n of Drosophila k e p t u n d e r constant environ-
ment w a s s h o w n t o be s e p a r a b l e by selection i n t o a n u m -
b e r of v e r y d i v e r s e s t r a i n s in r e s p e c t of d u r a t i o n of life.
I n L o e b a n d N o r t h r o p ' s e x p e r i m e n t s , a g e n e r a l mixed
p o p u l a t i o n of Drosophila, b u t of p r e s u m a b l y constant
genetic constitution, a t l e a s t a p p r o x i m a t e l y such, t h r o u g h -
out the e x p e r i m e n t , w a s s h o w n t o e x h i b i t c h a n g e s of
d u r a t i o n of life w i t h c h a n g i n g e n v i r o n m e n t s . I t i s the
old f a m i l i a r deadlock. H e r e d i t y c o n s t a n t p l u s c h a n g i n g
STUDIES ON T H E DURATION OF LIFE 211
environment equals diversity. E n v i r o n m e n t constant
p l u s v a r y i n g h e r e d i t a r y c o n s t i t u t i o n also e q u a l s d i v e r s i t y .
C a n we p e n e t r a t e n o f a r t h e r t h a n t h i s i n t o t h e m a t t e r T
I think in t h e p r e s e n t c a s e we can. I n L o e b a n d N o r t h r o p ' s
e x p e r i m e n t s , t e m p e r a t u r e a n d d u r a t i o n of life w e r e n o t
t h e only two t h i n g s t h a t v a r i e d . T h e different t e m p e r a -
t u r e g r o u p s also differed f r o m each o t h e r — b e c a u s e of t h e
t e m p e r a t u r e differences, t o b e s u r e , b u t n o t less r e a l l y —
in r e s p e c t of g e n e r a l m e t a b o l i c activity, e x p r e s s e d in
m u s c u l a r m o v e m e n t a n d e v e r y o t h e r w a y . I n t h e gene-
tic e x p e r i m e n t s m e t a b o l i c activity w a s s u b s t a n t i a l l y equal
in all t h e h e r e d i t a r i l y different lines. T h e i d e a s u g g e s t s
itself, both on a priori g r o u n d s a n d also u p o n t h e b a s i s
of c e r t a i n e x p e r i m e n t a l d a t a p r e s e n t l y t o be i n p a r t r e -
viewed, t h a t p o s s i b l y d u r a t i o n of life m a y be a n i m p l i c i t
function of only t h e t w o v a r i a b l e s
a. Genetic constitution
b. Rate of metabolic activity.
T h e functional r e l a t i o n s of metabolic a c t i v i t y w i t h
t e m p e r a t u r e , food, l i g h t a n d o t h e r e n v i r o n m e n t a l fac-
t o r s a r e all well known. F o r p r e s e n t p u r p o s e s w e do
n o t need to g o i n t o t h e q u e s t i o n of t h e i r e x a c t f o r m . T h e
essential p o i n t is t h a t all t h e s e e n v i r o n m e n t a l f a c t o r s
s t a n d in definite functional r e l a t i o n s to r a t e of metabolic
activity, a n d do n o t so s t a n d i n r e l a t i o n to genetic consti-
tution. Genetic c o n s t i t u t i o n is n o t a f u n c t i o n of t h e
environment, b u t is, f o r a n y i n d i v i d u a l , a c o n s t a n t , a n d
only v a r i e s b e t w e e n i n d i v i d u a l s .
T h i s m a y be t h o u g h t m e r e l y t o b e a n i n v o l v e d w a y of
s a y i n g w h a t one k n o w s a priori: n a m e l y , t h a t d u r a t i o n
of life, i n g e n e r a l a n d i n p a r t i c u l a r , d e p e n d s only u p o n
h e r e d i t y a n d e n v i r o n m e n t . S o in one s e n s e i t i s . B u t
the essential p o i n t I w o u l d m a k e h e r e is t h a t t h e manner
212 BIOLOGY OF D E A T H
in which the environmental forces (of sub-lethal inten-
sity, of course) chiefly act in determining duration of
life, appears to be by changing the rate of metabolism of
the individual. F u r t h e r m o r e one w o u l d s u g g e s t , on t h i s
view, t h a t w h a t h e r e d i t y does in r e l a t i o n t o d u r a t i o n of
S T U D I E S ON T H E D U R A T I O N O F L I T E 213
m o v e a b o u t to a limited d e g r e e , b u t o t h e r w i s e under
c o n d i t i o n s , i n c l u d i n g t e m p e r a t u r e , identical w i t h those
i n t h e r e v o l v i n g cages, a t t a i n e d a n a v e r a g e age a t death
o f 40.3 m o n t h s . A l l w e r e s t a t e d to h a v e d i e d of "old
TABLE 26
Relation of longevity to muscular activity in rats (Slonaker)
TOTAX NUMBER OP MtfLES EUN" DURING UFB
SUMMARY OP RESULTS
I h a v e a t t e m p t e d t o r e v i e w some of t h e i m p o r t a n t
biological a n d s t a t i s t i c a l c o n t r i b u t i o n s w h i c h h a v e b e e n
m a d e to the k n o w l e d g e of n a t u r a l d e a t h a n d t h e d u r a t i o n
of life, a n d to s y n t h e s i z e t h e s e s c a t t e r e d r e s u l t s i n t o
a c o h e r e n t unified whoje. I n t h e p r e s e n t c h a p t e r I shall
endeavor to summarize, in the briefest way, the scattered
f a c t s which h a v e b e e n p a s s e d i n review, a n d t o follow a
p r e s e n t a t i o n of t h e g e n e r a l r e s u l t s to w h i c h t h e y l e a d
w i t h some discussion of w h a t w e m a y r e a s o n a b l y r e g a r d
t h e f u t u r e a s h a v i n g i n s t o r e f o r u s , so f a r a s m a y be
j u d g e d f r o m o u r p r e s e n t k n o w l e d g e of t h e t r e n d of e v e n t s .
"What a r e t h e g e n e r a l r e s u l t s of o u r r e v i e w of t h e gen-
e r a l biology of d e a t h ? I n t h e first place, one p e r c e i v e s t h a t
n a t u r a l d e a t h is a r e l a t i v e l y n e w thing, w h i c h a p p e a r e d
first i n evolution w h e n d i f f e r e n t i a t i o n of cells f o r p a r t i c -
u l a r functions c a m e i n t o existence. U n i c e l l u l a r ani-
m a l s a r e , a n d a l w a y s h a v e been, i m m o r t a l . T h e cells of
h i g h e r organisms, set a p a r t for reproduction in the
c o u r s e of differentiation d u r i n g evolution, a r e i m m o r t a l .
T h e only r e q u i s i t e c o n d i t i o n s t o m a k e t h e i r p o t e n t i a l im-
m o r t a l i t y a c t u a l a r e physico-chemical i n n a t u r e a n d a r e
n o w f a i r l y well u n d e r s t o o d , p a r t i c u l a r l y a s a r e s u l t of
t h e i n v e s t i g a t i o n s of L o e b u p o n artificial p a r t h e n o g e n e s i s
a n d related phenomena. T h e essential a n d i m p o r t a n t
223
224 BIOLOGY OF D E A T H
somatic cells of the body, h o w e v e r m u c h differentiated,
a r e also potentially i m m o r t a l ; b u t t h e c o n d i t i o n s neces-
s a r y f o r t h e a c t u a l r e a l i z a t i o n of t h e p o t e n t i a l immor-
t a l i t y are, i n t h e n a t u r e of the case, a s h a s b e e n shown
by t h e brilliant r e s e a r c h e s of L e o L o e b , HarriHon a n d
C a r r e l on t i s s u e c u l t u r e , s u c h as c a n n o t be realized so
long a s t h e s e cells a r e actually i n a n d a p a r t of t h e h i g h e r
m e t a s o a n body. T h e r e a s o n w h y thin i» BO, a n d why in
consequence d e a t h r e s u l t s i n t h e m e t a z o a , in t h a t , in such
o r g a n i s m s t h e specialization of s t r u c t u r e a n d function
necessarily m a k e s the s e v e r a l p a r t s of the b o d y m u t u a l l y
d e p e n d e n t f o r t h e i r life u p o n each o t h e r . If one o r g a n
or g r o u p , f o r a n y accidental r e a s o n b e g i n s to function
a b n o r m a l l y a n d finally b r e a k s down, t h e b a l a n c e of t h e
whole is u p s e t a n d d e a t h e v e n t u a l l y follows. B u t t h e
individual cells, themselves, could g o o n l i v i n g indefinitely,
if t h e y w e r e freed, a s t h e y a r e in c u l t u r e s , of t h e neces
sity of d e p e n d i n g u p o n t h e p r o p e r f u n c t i o n i n g of o t h e r
cells for t h e i r food, oxygen, etc.
S o t h e n we see e m e r g i n g , a s o u r first g e n e r a l r e s u l t ,
t h e fact t h a t n a t u r a l d e a t h i s n o t a n e c e s s a r y o r inevit-
able consequence of life. I t is n o t a n a t t r i b u t e of t h e
cell. I t is a b y - p r o d u c t of p r o g r e s s i v e e v o l u t i o n — t h e
price we p a y f o r differentiation a n d s p e c i a l i s a t i o n of
s t r u c t u r e a n d function.
T h i s first r e s u l t i n d i c a t e s logically, in a n y p a r t i c u -
l a r o r g a n i s m such a s m a n , t h e g r e a t i m p o r t a n c e of
a q u a n t i t a t i v e a n a l y s i s of t h e m a n n e r i n w h i c h dif-
f e r e n t p a r t s of t h e b o d y b r e a k d o w n a n d lead t o d e a t h .
S u c h a n analysis, carefully w o r k e d t h r o u g h , d e m o n s t r a t e s
that this breaking down is not a haphazard process, b u t
a highly o r d e r l y one r e s t i n g u p o n a f u n d a m e n t a l biolog-
ical basis* T h e p r o g r e s s of t h e b a s i c tissue e l e m e n t s
NATURAL DEATH, PUBLIC HEALTH 225
of the body along the evolutionary pathway appears to be
an important factor in determining the time when the
organ systems in which they are chiefly involved shall
break down. Those organ systems that have evolved
farthest away from original primitive conditions are
the soundest and most resistant, and wear the longest
under the strain of functioning. So then, the second
large result is that it is the way potentially immortal
cells are put together in mutually dependent organ sys-
tems that immediately determines the time relations of
the life span.
But it was possible to penetrate more deeply into the
problem than this by finding that the duration of life is
an inherited character of an individual, passed on from
parent to offspring, just as is eye color or hair color, and
with a relatively high degree of precision. This has
been proved in a variety of ways, first directly for man
(Pearson) and for a lower animal, Drosophila, (Hyde,
Pearl) by measuring the degree of hereditary transmis-
sion of duration of life, aad indirectly by showing that
the death rate was selective (Pearson, Snow, Bell, Ploetz)
and had been, since nearly the beginning of recorded his-
tory, at least. It is heredity which determines the way
the organism is put together—the organization of the
parts. And it is when parts break down and the organ-
ization is upset that death comes. So the third large re-
sult is that heredity is the primary and fundamental
determiner of the length of the span of life.
Finally, it is possible to say probably, though not as
yet definitely because the necessary mass of experimen-
tal evidence is still lacking, but will, I believe, be shortly
provided, that environmental circumstances play their
15
226 BIOLOGY OF D E A T H
p a r t in d e t e r m i n i n g the d u r a t i o n of life l a r g e l y , if not
in principle entirely, b y influencing t h e rate a t which the
vital p a t r i m o n y is spent. If we live r a p i d l y , like Loeb
a n d N o r t h r o p ' s Drosophila a t t h e h i g h t e m p e r a t u r e s , our
lives m a y be m o r e i n t e r e s t i n g , b u t t h e y will n o t be so
long. The fact a p p e a r s t o be, t h o u g h r e s e r v a t i o n of
final j u d g m e n t is n e c e s s a r y till m o r e r e t u r n s a r e in,
t h a t heredity d e t e r m i n e s t h e a m o u n t of c a p i t a l placed in
the vital bank u p o n which w e d r a w t o continue life, and
which when all u s e d u p spells d e a t h ; while environment,
using the t e r m in the b r o a d e s t sense to include habits
of life as well as physical s u r r o u n d i n g s , d e t e r m i n e s the
r a t e a t which d r a f t s a r e p r e s e n t e d a n d cashed. The
case seems in principle like w h a t obtains i n r e s p e c t of the
d u r a t i o n of life of a m a n - c o n s t r u c t e d machine. I t is
self-evident t h a t if, of t w o automobiles of t h e s a m e make
leaving the factory t o g e t h e r n e w a t t h e s a m e time, one is
r u n a t the r a t e of 1,000 miles p e r y e a r a n d t h e o t h e r at
the r a t e of 10,000 miles p e r y e a r , t h e useful life of the
f o r m e r is bound t o be m u c h l o n g e r in time t h a t t h a t of
t h e latter, accidents being excluded i n b o t h cases. Again,
a v e r y high p r i c e d car, well-built of t h e finest m a t e r i a l ,
m a y have a s h o r t e r d u r a t i o n of life t h a n t h e poorest
and cheapest machine, p r o v i d e d the a n n u a l m i l e a g e output
of the f o r m e r is m a n y t i m e s t h a t of the l a t t e r .
The first t h r e e of these conclusions seem to be firmly
grounded. T h e l a s t r e s t s , a t p r e s e n t , u p o n a less secure
footing. Because i t does, i t offers a n extremely promis-
ing field for b o t h statistical a n d e x p e r i m e n t a l research.
W e need a wide v a r i e t y of investigations, like those of
Loeb and N o r t h r o p , of S l o n a k e r a n d of E u b n e r , on the
experimental side. On t h e s t a t i s t i c a l side, well-conceived
NATURAL DEATH, PUBLIC HEALTH 227
a n d careful studies, by the most refined of modern meth-
ods, upon occupational mortality seem likely to yield
la,rge returns.
PUBLIC HEALTH ACTIVITIES
Fortunately, it is possible to get some light on the
environmental side from existing statistical data by con-
sidering, in a broad general way, the results of public
health activities. Any public health work, of course,
deals, and can deal in the present state of public senti-
ment and enlightenment, only with environmentalmatters.
-Attempts at social control of the germ-plasm—the innate
inherited constitutional make-up—of a people, by eugenic
legislation, have not been conspicuously successful. And
there is a good deal of doubt, having regard to all factors
necessarily involved,, whether they have always been
even well-conceived. As an animal breeder of some
years' experience, I have no doubt whatever that almost
any breeder of average intelligence, if given omnipotent
oontrol over the activities of human beings, could, in a
few generations, breed a race of men on the average con-
siderably superior—by our present standards—to any
x*ace of men now existing in respect of many qualities or
attributes. But, as a practical person, I am equally sure
that nothing of the sort is going to be dojie by legislative
action or any similar delegation of powers. Before any
sensible person or society is going to entrust the control
of its germ-plasm to politics or to science, there will be
demanded that science know a great deal more than it
mow does about the vagaries of germ-plasms and how to
control them. Another essential difficulty is one of stan-
dards. Suppose it to be granted that our knowledge of
228 BIOLOGY OF D E A T H
genetics w a s sufficiently ample and profound to make it
possible to make a racial germ-plasm exactly whatever
one pleased; what individual or group of individuals
could possibly be trusted to decide what it should be?
Doubtless many persons of uplifting tendencies would
promptly come forward prepared to undertake such a
responsibility- B u t what of history? If it teaches us
anything, it is that social, moral and political standards
are not fixed and absolute, but vary, and v a r y radically
in both space and time. A n d further, history teaches
that a great many of the most valuable people, in the
highest and best sense, whom the world has ever known,
were so constituted, physically, morally} or otherwise,
as to make it certain that under a strict eugenic regime
they never would have existed at all. One cannot but
feel that man's instinctive wariness about experimental
interferences with his germ-plasm i s i n considerable
degree, well-f ojunded.
But because of the altogether more impersonal na-
ture of the case, most men individually and society in
general are perfectly willing to let anybody do anything
they like in the direction of modifying the environment in
what is believed, or hoped t o be, the direction of improve-
ment, or trying to, quite regardless of whether science
i s able to give any slightest inkling on the basis of ascer-
tained facts as to whether the outcome will be good, bad
or indifferent. Hence many kinds of weird activities and
propaganda flourish like the proverbial bay tree.
Of all organized activities looking towards the direct
modification of the environment to the benefit of mankind,
that group comprised under the terms sanitation, hygiene
NATURAL DEATH, PUBLIC HEALTH 229
a n d public h e a l t h h a v e , b y all odds, t h e b e s t c a s e w h e n
m e a s u r e d i n t e r m s of accomplishment. M a n ' s e x p e c t a -
tion of life h a s i n c r e a s e d a s he h a s come d o w n t h r o u g h
t h e c e n t u r i e s (cf. P e a r s o n a n d Macdonell.) A l a r g e
p a r t of this i m p r o v e m e n t m u s t surely b e c r e d i t e d t o h i s
i m p r o v e d u n d e r s t a n d i n g of h o w t o cope w i t h a n a l w a y s
m o r e o r less inimical e n v i r o n m e n t a n d a s s u a g e i t s a s p e r -
ities t o his g r e a t e r c o m f o r t a n d well-being. T o fail t o
give this c r e d i t w o u l d b e m a n i f e s t l y a b s u r d .
B u t it would be equally absurd to attempt t o main-
t a i n t h a t all decline i n t h e d e a t h - r a t e which h a s o c c u r r e d
h a s been d u e t o t h e efforts of h e a l t h officials, w h e t h e r
conscious or u n c o n s c i o u s , as i s often a s s e r t e d a n d still
m o r e often i m p l i e d i n t h e i m p a s s i o n e d o u t p o u r i n g s of
zealous p r o p a g a n d i s t s . T h e open-minded s t u d e n t of t h e
n a t u r a l h i s t o r y of d i s e a s e knojws p e r f e c t l y well t h a t a
l a r g e p a r t of t h e i m p r o v e m e n t i n t h e r a t e of m o r t a K t y
c a n n o t possibly h a v e b e e n d u e t o a n y s u c h efforts. To
i l l u s t r a t e t h e point, I h a v e p r e p a r e d a series of i l l u s t r a -
tions d e a l i n g w i t h conditions i n t h e R e g i s t r a t i o n A r e a
of t h e U n i t e d S t a t e s i n t h e i m m e d i a t e p a s t . A l l t h e s e
d i a g r a m s ( F i g u r e s 52, 53, a n d 54) give d e a t h - r a t e s p e r
100,000 f r o m v a r i o u s c a u s e s of d e a t h i n t h e p e r i o d of
1900-1918, inclusive, b o t h sexes f o r simplicity b e i n g t a k e n
t o g e t h e r . T h e lines a r e all p l a t t e d o n a l o g a r i t h m i c
scale. T h e r e s u l t of t h i s m e t h o d of p l o t t i n g i s t h a t t h e
slope t r e n d of each, l i n e i s d i r e c t l y c o m p a r a b l e w i t h t h a t
of a n y other, n o m a t t e r w h a t t h e a b s o l u t e m a g n i t u d e of
t h e r a t e s concerned. I t is t h e s e slopes, m e a s u r i n g im-
p r o v e m e n t i n m o r t a l i t y , t o which I w o u l d especially
direct attention.
230 BIOLOGY OF D E A T H
100
10
0/ 1900
i i i i i i i i i i i i i i i i i
01 02 03 04- 05 06 O7 06 09 1O
12 13II14- 15 16 17 18
YEAR
FIG. 52.—Trend of death rates for four causes of death against which public health activities
have been particularly directed.
I n figure 52 a r e given t h e t r e n d s of t h e d e a t h - r a t e s
f o r f o u r diseases a g a i n s t which public h e a l t h a n d s a n i -
t a r y activities h a v e been p a r t i c u l a r l y a n d v i g o r o u s l y
NATURAL DEATH, PUBLIC HEALTH 231
directed, with, as we are accustomed to say, most grati-
fying results. The diseases are:
1. Tuberculosis of the lungs.
2. Typhoid fever.
3. Diphtheria and croup.
4. Dysentery.
We note at once that the death-rates from these
diseases have all steadily declined in the 19 years tinder
review. But the rate of drop has been slightly unequal.
Uemembering that the slopes are comparable, where-
ever the lines may lie, and that an equal slope means a
relatively equally effective diminution of the mortality
of the disease, we note that the death-rate from tuber-
culosis of the lungs has decreased slightly less than any
of the other three. Yet it may fairly be said that so
strenuous a warfare, or one engaging in its ranks so many
earnest and active workers, has probably never in the
liistory of the world been waged against any disease as
that which has been fought in the United States against
tuberculosis in the period covered. The rates of decline
of the other three diseases are all practically identical.
Figure 53 shows entirely similar trends for four
other causes of death—namely:
1. Bronchitis (acute and chronic).
2. Paralysis without specified cause.
3. Purulent infection and septicaemia.
4. Softening of the brain.
Now it will be granted at once, I think, that public
health and sanitation can have had, at the utmost, ex-
tremely little, if anything, to do with the trend of mor-
tality from these four causes of death. For the most
part they certainly represent pathological entities far
"beyond the present reach of the health officer. Yet the
232 BIOLOGY OF DEATH
at 1 1 1 ! 1 1 1 1 1 1 1 1 1 1 1 I 1 11
19000102 0304050607060910 IZ 0 f+ 15 ft f7 IS
FIG. 53.—Trend of death rates from four oawm YEARof 4m,th upon which no dlr*rtfttlrrnptat
oontrol has been made.
outstanding fact is that their rates of mortality havo de-
clined and are declining just as did those in the control-
lable group shown in Figure 52. I t is of no moment
NATURAL DEATH, PUBLIC HEALTH 233
1,000
I0O —
O.I I 1 1 I 1 I I 1 1 1 I 1
1900 01 02 03 04- 05 06 OT 08 09 10 II IZ 13 15 16 17 id
YEAR
FIG. 54.—Trend of combined death rate from the four causes ehown in Figure 52 as compared
with the four causes shown in Figure 53.
to say that the four causes of d e a t h in t h e s e c o n d group
are absolutely of l e s s i m p o r t a n c e than some of those in
the first group, because w h a t w e a r e here discussing
is not relative force of mortality from different causes,
234 BIOLOGY OF DEATH
b u t r a t h e r t h e trend of m o r t a l i t y from p a r t i c u l a r c a u s e s .
T h e rate of decline is j u s t a s significant, w h a t e v e r the
a b s o l u t e p o i n t f r o m which t h e c u r v e s t a r t s .
I t is difficult to c a r r y in t h e mind a n exact i m p r e s s i o n
of t h e slope of a line, so, in o r d e r t h a t a c o m p a r i s o n m a y
be m a d e , I h a v e p l o t t e d in F i g u r e 54, first, the* total r a t ft
of m o r t a l i t y f r o m t h e f o u r controllable causes of d e a t h
t a k e n t o g e t h e r and, second, t h e total r a t e of m o r t a l i t y
f r o m t h e four uncontrolled causes t a k e n t o g e t h e r . T h e
r e s u l t is i n t e r e s t i n g . T h e t w o lines w e r e actually n e a r e r
t o g e t h e r in 1900 t h a n t h e y w e r e in 1918. T h e y h a v e
d i v e r g e d because the recorded m o r t a l i t y from the uncon-
trolled four h a s actually decreased f a s t e r in the 19 y e a r s
t h a n h a s t h a t f r o m the f o u r a g a i n s t which we have W n
actively fighting. T h e divergence is n o t g r o a t , however.
P e r h a p s we a r e only justified in s a y i n g t h a t the m o r t a l i t y
in each of the t w o g r o u p s h a s notably declined, and a t not
f a r from identical r a t e s .
N o w the f o u r diseases in t h i s g r o u p , I ehoae quite* at.
r a n d o m from a m o n g the causes of d e a t h whose r a t OH T
k n e w t o be declining, to u s e a s a n i l l u s t r a t i o n solely. I
could easily pick out eight o t h e r causes of death which
would i l l u s t r a t e t h e s a m e point. I d o not wish too much
s t r e s s to be laid u p o n t h e s e e x a m p l e s . If t h e y m a y nerve
merely t o drive s h a r p l y h o m e i n t o the m i n d t h a t it i« only
t h e t y r o or the reckless p r o p a g a n d i s t , long a g o a n t r a n g ' T
to t r u t h , who will v e n t u r e t o a s s e r t t h a t a declining d e a t h -
r a t e in cmd of itself m a r k s the successful r e s u l t of h u m a n
effort, I shall be a b u n d a n t l y sat-infied.
I t h a s been objected t h a t t h e decline shown by t h e
f o u r " n o n - c o n t r o l l e d " causes in the e x a m p l e jimt d e a l t
w i t h is due wholly, o r n e a r l y so, to changen in the p r a c t i c e
of physicians r e l a t i v e t o t h e r e p o r t i n g of t h e c a u s e of
NATURAL DEATH, PUBLIC H E A L T H 235
d e a t h , a n d t h a t , t h e r e f o r e , t h e decline i s s p u r i o u s . I h a v e
n o t been able to find t h a t t h e r e is a n y good evidence t h a t
t h i s is t h e f a c t ; t h a t , i n s h o r t , c h a n g e s i n r e p o r t i n g p r a c -
tice h a v e affected t h e " n o n - c o n t r o l l e d " g r o u p m o r e t h a n
t h e " c o n t r o l l a b l e " g r o u p . B u t a n o t h e r k i n d of e x a m p l e
m a y be cited t o i l l u s t r a t e t h e s a m e g e n e r a l p o i n t . S u p p o s e
w e c o m p a r e t h e c o u r s e of m o r t a l i t y f r o m c e r t a i n well-
defined causes, a b o u t t h e r e p o r t i n g of w h i c h t h e r e c a n b e
n o c o n t r o v e r s y , in ( a ) a g r o u p of c o u n t r i e s s t a n d i n g i n a n
a d v a n c e d position i n m a t t e r s of p u b l i c h e a l t h , s a n i t a t i o n ,
etc., a n d (b) a g r o u p of c o u n t r i e s r e l a t i v e l y b a c k w a r d a n d
undeveloped in these r e s p e c t s . S u c h a c o m p a r i s o n is im-
possible t o m a k e o v e r a n y l o n g p e r i o d of t i m e b e c a u s e of
lack of c o m p a r a b l e d a t a . I have, succeeded i n g e t t i n g com-
parable statistics on two diseases, namely typhoid fever
a n d d i p h t h e r i a , f o r t h e p e r i o d 1898 t o 1912 inclusive, f o r
t h e following c o u n t r i e s :
A. Countries having (in period B. Countries having (in period
covered) highly developed covered) less highly developed
public health and sanitation. public health and sanitation
Australia than those in group A.
Austria Italy
England and Wales Jamaica
Germany Roumania
W i t h o u t going i n t o d e t a i l e d c o m p a r i s o n s , w h i c h m i g h t
be t h o u g h t invidious, i t is e v i d e n t on t h e face of t h e case,
I think, t h a t t h e c o u n t r i e s i n t h e A g r o u p w e r e , o n t h e
a v e r a g e d u r i n g t h e p e r i o d covered, m u c h m o r e a d v a n c e d
i n all p r a c t i c a l public h e a l t h m a t t e r s t h a n w e r e t h e coun-
tries in group B.
I n F i g u r e s 55 a n d 56 a r e s h o w n t h e t r e n d s of t h e
weighted average death r a t e s from typhoid fever and
d i p h t h e r i a r e s p e c t i v e l y i n t h e two g r o u p s of c o u n t r i e s .
I t is evident f r o m t h e s e d i a g r a m s t h a t t h e d e a t h r a t e s
236 BIOLOGY O F DEATH
from these two causes d e c l i n e d , during the period cov-
ered, in both the A and t h e B groups of countries and
at not far from the same r a t e . There is no such large
difference aa would be e x p e c t e d if organized human inter-
ference with the natural h i s t o r y of disease always played
100
Mill
TYPHOID F£V£R
^ *
\
1 1 1 1 1 II
10 —
^ ^ ~
\
Mil 1 1 1 1 1 1 1 I 11 |
99 1900 Ot OH OS 04 OS O6 07 06 09 fO // IZ
F J O . 55,-—-Counse of the weighted a v e r a g e d e a t h r a t e , f o r t h e countries i n t h e A (solid line)
* o d B (broken l i n e ) g r o u p s , f r o m t y p h o i d fever.
p r o p a g a n d i s t a s s e r t s t h a t i t d o e s .
T o g u a r d a g a i n s t t h e p o s s i b i l i t y of a n y m i s u n d e r -
tihat t h e a b o v e is n o t i n t e n d e d i n a n y w a y t o c o n v e y t h e
IS&8 39 1900 01
OS 04OS 0607 08 00
VCAR.
Via. 50.—Likefigure55, but for diphtheria and croup.
extraordinarily complex biological phenomenon, in
which many factors besides the best efforts of health
officials are involved.
2. That for many causes of death a vast lot needs to
be added to our knowledge of etiology, in the broadest
sense, before really efficient control can be hoped for.
This knowledge can come only through scientific investi-
238 BIOLOGY OF D E A T H
gation, a n d n o t t h r o u g h t h e complacent acceptance of t h e
p r o p a g a n d i s t ' s a s s u r a n c e t h a t " i f w h a t knowledge we
now h a v e is applied, all will be w e l l . " *
M a n y o t h e r s h a v e , of course, p e r c e i v e d t h a t , in the
n a t u r a l h i s t o r y of disease-, m o r t a l i t y f r o m p a r t i c u l a r
causes m a y decline o v e r long p e r i o d s of time without any
relation t o w h a t h e a l t h d e p a r t m e n t s h a v e done, o r t r i e d
to d o a b o u t it. F o r example, Given h a s r e c e n t l y pointed
out t h a t t h e r e i s no evidence t h a t a n y t h i n g t h a t m a n h a s
done h a s affected, i n e i t h e r one w a y o r t h e other, the
decline in t h e m o r t a l i t y of tuberculosis, which h a s been
continuous f o r n e a r l y t h r e e - q u a r t e r s of a century.
P e a r s o n h a s discussed t h e s a m e point.
T h e r e i s m u c h i n o u r public h e a l t h w o r k t h a t i s w o r t h y
of the h i g h e s t p r a i s e . W h e n based u p o n a sound founda-
tion of a s c e r t a i n e d f a c t i t m a y , a n d does, p r o c e e d w i t h a
step as firm a n d i n e x o r a b l e a s t h a t of F a t e itself, to the
wiping out of p r e v e n t a b l e m o r t a l i t y . T w o recent ex-
amples m a y b e cited h e r e , by w a y of specific i l l u s t r a t i o n
of w h a t real a n d r e a s o n a b l y complete scientific knowledge
can accomplish i n public h e a l t h work. B o t h examples
a r e t a k e n f r o m t h e w o r k of t h e I n t e r n a t i o n a l H e a l t h
B o a r d of t h e Eockef eller F o u n d a t i o n , w i t h t h e p e r m i s s i o n
of i t s director, M r . Wickliffe E o s e .
T h e first concerns m a l a r i a . T h e life cycle of the
m a l a r i a p a r a s i t e is definitely known, a n d furnished a
* One can but wonder if the many scientific men, who permit, and to
some extent approve, such assertions, have ever thought of the menace to
the continued support of research in science in general which inheres in
this attitude of mind. The support of research comes finally back always
to society in general—to the "average citizen" in short. Is it the part
of wisdom to leave his education as to the meaning and significance of
science for his happiness and well-being, so entirely in the hands of the
propagandist as we now do? Has anti-vivisection taught no lesson?
NATURAL DEATH, PUBLIC HEALTH 239
definite scientific basis fox control procedure. "It is
•well understood, not only by scientists, but also by intel-
ligent laymen, that the spread of the infection may be
prevented by mosquito control, by protecting people from
being bitten by mosquitoes, or by destroying the parasite
in the blood of the human carrier. It has been shown,
moreover, by repeated demonstrations, that by applica-
tion of any one of these measures, or of any combination
of them, the amount of malaria in a community may be
reduced indefinitely. There are few diseases that pre-
sent so many vulnerable points of attack and none per-
haps the control of which may be made more definite
or certain." (Eose).
In 1916 the International Health Board undertook
some experiments in control at Crossett, Ark. In des-
cribing the work Rose says:
"Effort has been made to test the feasibility of malaria control in
small communities by resort to such simple anti-mosquito measures as
would fall within the limits of expenditure that such communities might
well afford. The habits of the three* mosquitoes—A. quadrimaculatus Say,
A. punctipenms Say, and A. oruzians Wiedermann—which are responsible
for the infection in these communities have been made the subject of
constant study with a view to eliminating all unnecessary effort, and thereby
reducing cost.
"Experiment at Crossett, 1910—The first of these tests was undertaken
at Crossett, a lumber town of 2,129 inhabitants, situated in Ashley County
in south-eastern Arkansas, about 12 miles north of the Louisiana line.
Crossett lies at the edge of the so-called "uplands," in a level, low-lying
region (elevation 165 feet), with sufficient undulation to provide reason-
ably good natural drainage. Climatic conditions and abundant breeding
places favor the propagation of anopheles. Malaria, in its severe form,
is widely prevalent as an endemic infection, and according to the estimate
of local physicians, is the cause of about 60 per cent, of all illness through-
out the region. Within the town itself the malaria rate was high, and
was recognized by the lumber corporation and the people as a serious
menace to health and working efficiency.
"The initial step in the experiment was a survey of the community
to determine the malaria incidence, to ascertain in the species of mosquitoes
240 BIOLOGY OF DEATH
responsible for the spread of the infection, and to locate the breeding places
of these mosquitoes. Breeding places were exhibited on a community
map, and organized effort was centered on their destruction or control.
The program of simple measures excluded all major drainage. Barrow
pits and shallow ponds were filled or drained; streams were cleared of
undergrowth when necessary to let the sunlight in; their margins and beds
wore cleared of vegetation and obstruction; and they were trained to a
narrow channel, thus providing an unobstructed off-flow. Artificial con-
tainers were removed from premises; water barrels on bridges were treated
with nitre cake. All remaining breeding places were regularly treated by
removing vegetation, opening up shallow margins to give free access to
small fish, and spraying once a week with road oil by means of automatic
drips or a knapaack sprayer. All operations were under the supervision
of a trained lay inspector. Care was exercised to eliminate all unnecessary
effort and to secure, not the elimination of the last mosquito, but a rea-
sonably high degree of control at a minimum cost/*
T h e r e s u l t s a r e s h o w n i n F i g u r e 57, as m e a s u r e d by
a n u m b e r of p h y s i c i a n s 7 c a l l s f o r t h e t r e a t m e n t of ma-
l a r i a in the community.
T h e second example s h o w s t h e effectiveness of con-
t r o l of yellow fever, a n o t h e r d i s e a s e f o r which definite
scientific knowledge e x i s t s a s t o etiology and m o d e
of t r a n s m i s s i o n .
N o t h i n g could m o r e c o n v i n c i n g l y d e m o n s t r a t e t h a n
does F i g u r e 58 the e f f e c t i v e n e s s w i t h which this disease
can be controlled. T h e d i a g r a m s h o w s the r e s u l t s of
the I n t e r n a t i o n a l H e a l t h B o a r d ' s y e l l o w f e v e r w o r k in
Guayaquil in 19184920.
JULY
MAY
JUNE
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JUNE
MARCH
MARCH
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FEB.
JAN.
OCT.
AUG.
SEPT.
AUG.
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JAN.
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NATURAL DEATH, PUBLIC HEALTH 243
born, t h a n w a s f o r m e r l y t h e case. I t is f u r t h e r m o r e
p l a i n t h a t if n o t h i n g h a p p e n s to t h e b i r t h - r a t e t h e r e m u s t
eventually be as m a n y p e r s o n s living u p o n t h e h a b i t a b l e
p a r t s of the globe a s can p o s s i b l y b e s u p p o r t e d w i t h
food a n d the o,ther necessities of life. M a l t h u s , w h o m
e v e r y one discusses b u t few t a k e t h e t r o u b l e to r e a d ,
p o i n t e d out m a n y y e a r s a g o t h a t t h e p r o b l e m of p o p u -
l a t i o n t r a n s c e n d s , in its d i r e c t i m p o r t a n c e to t h e w e l f a r e
of h u m a n beings a n d f o r m s of social o r g a n i z a t i o n , all
o t h e r p r o b l e m s . L a t e l y we h a v e h a d a d e m o n s t r a t i o n on
a g h a s t l y g i g a n t i c scale of t h e t r u t h of M a l t h u s ' conten-
tion. F o r , in l a s t a n a l y s i s , i t c a n n o t be d o u b t e d t h a t one
i m p o r t a n t u n d e r l y i n g c a u s e of t h e g r e a t w a r , t h r o u g h
which we h a v e j u s t p a s s e d , w a s t h e e v e r - g r o w i n g p r e s -
s u r e of p o p u l a t i o n u p o n subsistence.
A n y s y s t e m o r f o r m of a c t i v i t y w h i c h t e n d s , by how-
e v e r slight a n a m o u n t , t o k e e p m o r e p e o p l e a l i v e a t a given
i n s t a n t of time t h a n w o u l d o t h e r w i s e r e m a i n alive, a d d s
t o t h e difficulty of t h e p r o b l e m of p o p u l a t i o n . W e h a v e
j u s t seen t h a t t h i s is p r e c i s e l y w h a t o u r public-health
activities a i m to do, a n d i n w h i c h t h e y succeed in a n o t
inconsiderable d e g r e e . B u t s o m e o n e will s a y a t once
t h a t , while i t i s t r u e t h a t t h e d e a t h - r a t e is f a l l i n g m o r e
o r less generally, still t h e b i r t h - r a t e i s f a l l i n g concomi-
t a n t l y , so we n e e d n o t w o r r y a b o u t t h e p o p u l a t i o n p r o b -
lem. I t i s evident t h a t if w e r e g a r d t h e p o p u l a t i o n
p r o b l e m i n t e r m s of w o r l d - a r e a , r a t h e r t h a n t h a t of a n y
p a r t i c u l a r c o u n t r y , its d e g r e e of i m m e d i a c y d e p e n d s u p o n
t h e r a t i o of b i r t h s to d e a t h s i n a n y given t i m e u n i t . I f
w e examine, as I h a v e r e c e n t l y done, t h e s e d e a t h - b i r t h
r a t i o s f o r different c o u n t r i e s , we find t h a t t h e y give u s
little h o p e of a n y solution of t h e p r o b l e m of p o p u l a t i o n
244 BIOLOGY OF DEATH
by virtue of a supposed general positive correlation be-
tween birth-rates and death-rates.
The relation of birth-rate and death-rate changes to
population changes is a simple one and may be put this
way. If, neglecting migration as wo are justified in
doing in the war period and in considering the world prob-
lem, in a given time unit the percentage
100 Deaths
Births
has a value less than 100, it means that the births exceed
the deaths and that the population is increasing within
the specified time unit. If, on the other hand, the per-
centage is greater than 100, it means that the deaths are
more frequent than the births and that the population
is decreasing, again within the specified time unit. The
TABLE 29
Percentage of Deaths to Births
77 non-invaded Bavaria Knaland ami
Year departments Prussia
of France
1913 97 per cent. 58 percent. 67 per cent.
1914 110 per cent. 66 per cent. 74 per cent, 59 per cent.
1915 169 per cent. 101 per cent, 98Jper cent 09 per cent.
1916 193 per cent. 117 per cent. 131 per cent. 05 per cent.
1917 179 per cent. 140 per cent. 127 per cent. 75 per cent.
1918 198 per cent. 132* per cent. 146 per cent. 02 per mnt*
1919 154 per cent. 78 par cent.
1920 42* per eeut.
* First three-fourths of year only.
ratio of deaths to births may be conveniently designated
as the vital index of a population.
F r o m the raw data of births and deaths, I have cal-
culated the percentage which the deaths were of tha births
for (a) the 77 non-invaded departments of F r a n c e ; (b)
NATURAL DEATH, PUBLIC HEALTH 245
Prussia; (c) Bavaria; and (d) England and Wales, from
1913 to 1920 by years. The results are shown in Table 29.
The points to be especially noted in Table 29 are:
1. In all the countries here dealt with the death-birth
ratio in general rose throughout the war period. This
means that the proportion of deaths to births increased
so long as the war continued.
2. But in England it never rose to the 100 per cent,
mark. In other words, in spite of all the dreadful effects
of war, England's population wenfi on making a net
increase throughout the war.
3. Immediately after the war was over, the death-
birth ratio began to drop rapidly in all countries. In
England in 1919 it had dropped back from the high figure
of 92 per cent, in 1918 to 73 per cent. In France it dropped
from the high figure of 198 in 1918 to 154 in 1919, a
lower figure than France had shown since 1914. In all
the countries the same change is occurring at a rapid pace.
Perhaps the most striking possible illustration of this
is the history of the death-birth ratip of the city of
Vienna, shown in Figure 4, with data from the United
States and England and Wales for comparison. Prob-
ably no single large city in the world was so hard hit by
the war as Vienna. Yet observe what has happened to
its death-birth ratio. Note how sharp is the decline in
1919 after the peak in 1918. In other words, we see
how promptly the growth of population tends to regulate
itseK back towards the normal after even so disturbing
an upset as a great war.
In the United States, the death-birth ratio was not
affected at all by the war, though it was markedly altered
by the influenza epidemic. The facts are shown in Fig-
ure 59 for the only years for which data are available.
246 BIOLOGY OF DEATH
The area covered is the United States birth registration
area. We see that with the very low death-birth ratio
of 56 in 1915, there was no significant change till the
influenza year 1918, when the ratio rose to 73 per cent.
2Z5
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ZOO \
173 A \
450
/
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\
125
' A
100
/ Af if*
75
50.
—
1
WEAR
1
^n! change in percentage which deaths were of births in each of the
>;»<*
175
UNITED STATE'S
150
115
100
76
60
Z3
1 I _.LJ_
/7OO 20 4O 60 80 J6OO JX>4O«O IOOO 2060 &> 2OO0W 40 6O gO £>QO
YCARS
Fxa. 81.—Showing the curve of growth of the population of the United State*. For further
explanation of this and the two following diagrams, see text.
tion of t h e comet. H e h a s , f r o m N e w t o n i a n p r i n c i p l e s ,
a g e n e r a l m a t h e m a t i c a l e x p r e s s i o n of t h e l a w s of m o t i o n
of heavenly bodies. H e m u s t t h e n c o n s t r u c t his whole
c u r v e f r o m t h e d a t a given b y t h e few o b s e r v a t i o n s . S o ,
similarly, the s t a t i s t i c i a n h a s b u t a r e l a t i v e l y few p o p u -
l a t i o n o b s e r v a t i o n s because census t a k i n g h a s been p r a c -
tised a l o n g p r e s e n t lines only a little m o r e t h a n a c e n t u r y .
A c c o r d i n g to t h e s t a g e i n h i s t o r i c a l d e v e l o p m e n t of t h e
c o u n t r y dealt with, he m a y h a v e given a n e a r l y , a l a t e , o r
a middle s h o r t piece of t h e p o p u l a t i o n " o r b i t " o r his-
t o r y . F r o m this he m u s t construct, on t h o b a s i s of h i s
g e n e r a l t h e o r y of " p o p u l a t i o n o r b i t s , " t h e whole h i s t o r y ,
p a s t a n d future, of t h e p o p u l a t i o n in q u e s t i o n .
T o d e m o n s t r a t e h o w successful t h e p o p u l a t i o n c u r v e
shown in F i g u r e 60 is in d o i n g this, t h r e e d i a g r a m s a r e
p r e s e n t e d , each i l l u s t r a t i n g t h e g r o w t h of t h e p o p u l a t i o n
NATURAL DEATH, PUBLIC HEALTH 251
i n a different c o u n t r y . T h e h e a v y solid p o r t i o n of e a c h
c u r v e shows t h e r e g i o n for w h i c h c e n s u s d a t a exist. T h e
l i g h t e r b r o k e n p a r t of t h e c u r v e shows t h e p o r t i o n s out-
side t h i s o b s e r v e d r a n g e . T h e circles s h o w t h e a c t u a l ,
k n o w n o b s e r v a t i o n s . T h e first c u r v e d e a l s w i t h t h e p o p u -
FRANCL
YEA2S
F i a . 6 2 . — S h o w i n g t h e c u r v e of g r o w t h o f t h e p o p u l a t i o n of F r a n c e .
l a t i o n o f t h e U n i t e d S t a t e s . H e r e t h e o b s e r v a t i o n s c o m e
f r o m t h e f i r s t p a r t o f t h e c u r v e , w h e n t h e p o p u l a t i o n w a s
l e a v i n g t h e l o w e r a s y m p t o t e . F i r s t s h o u l d b e n o t e d t h e
e x t r a o r d i n a r y a c c u r a c y w i t h w h i c h t h e m a t h e m a t i c a l
t h e o r y d e s c r i b e s t h e k n o w n f a c t s . I t w o u l d b e e x t r e m e l y
d i f f i c u l t , b y a n y p r o c e s s , t o d r a w a c u r v e t h r o u g h t h e o b -
s e r v e d c i r c l e s a n d c o m e n e a r e r t o h i t t i n g t h e m a l l t h a n
t h i s o n e d o e s .
B e f o r e c o n s i d e r i n g t h e d e t a i l e d c o n s e q u e n c e s o f t h i s
U n i t e d S t a t e s c u r v e i n r e l a t i o n t o t h e w h o l e p o p u l a t i o n
h i s t o r y o f t h e c o u n t r y , l e t u s first e x a m i n e s o m e c u r v e s
f o r o t h e r c o u n t r i e s , w h e r e t h e o b s e r v e d d a t a f e l l i n q u i t e
d i f f e r e n t p o r t i o n s o f t h e " p o p u l a t i o n o r b i t . " F i g u r e 6 2
252 BIOLOGY OF D E A T H
gives t h e c u r v e f o r F r a n c e . Since b e f o r e t h e t i m e w h e n
definite census r e c o r d s began, F r a n c e h a s been a r a t h e r
densely p o p u l a t e d c o u n t r y . All t h e d a t a w i t h which w e
h a d to work, b e l o n g t h e r e f o r e , t o w a r d s t h e final end of
t h e whole p o p u l a t i o n h i s t o r y curve. T h e k n o w n p o p u l a -
tion d a t a f o r F r a n c e a n d f o r t h e U n i t e d S t a t e s s t a n d a t
opposite ends of t h e whole h i s t o r i c a l c u r v e . One in an
old c o u n t r y whose p o p u l a t i o n is n e a r i n g t h e u p p e r l i m i t ;
t h e o t h e r a n e w c o u n t r y whose p o p u l a t i o n s t a r t e d f r o m
n e a r the lower a s y m p t o t e only a b o u t a c e n t u r y a n d a half
ago. B u t i t i s seen f r o m t h e d i a g r a m t h a t t h e g e n e r a l
t h e o r y of p o p u l a t i o n g r o w t h fits perfectly t h e known f a c t s
r e g a r d i n g F r a n c e ' s p o p u l a t i o n i n t h e 120 y e a r s f o r w h i c h
r e c o r d s exist. W h i l e t h e r e a r e some i r r e g u l a r i t i e s in t h e
observation, due p r i n c i p a l l y to t h e effects of t h e F r a n c o -
P r u s s i a n war, i t is p l a i n t h a t on t h e whole it would be
p r a c t i c a l l y impossible t o g e t a b e t t e r fitting line t h r o u g h
t h e o b s e r v a t i o n a l circles t h a n t h e p r e s e n t one.
W e h a v e seen t h a t t h e g e n e r a l t h e o r y of p o p u l a t i o n
describes with equal a c c u r a c y t h e r a t e of g r o w t h in a
y o u n g country, w i t h r a p i d l y i n c r e a s i n g p o p u l a t i o n , a n d
a n old country, w h e r e t h e p o p u l a t i o n is a p p r o a c h i n g close
t o the absolute s a t u r a t i o n p o i n t . L e t u s now see h o w i t
w o r k s for a c o u n t r y i n a n i n t e r m e d i a t e p o s i t i o n in r e s p e c t
of p o p u l a t i o n . F i g u r e 63 shows t h e p o p u l a t i o n h i s t o r y
of S e r b i a . H e r e i t will b e n o t e d a t once t h a t t h e h e a v y
line, which denotes t h e r e g i o n of known c e n s u s d a t a , lies
a b o u t i n t h e m i d d l e of t h e whole c u r v e . A g a i n t h e fit
of t h e o r y to; o b s e r v a t i o n is e x t r a o r d i n a r i l y close. N o
b e t t e r fit, by a g e n e r a l l a w involving n o m o r e t h a n 3 con-
s t a n t s , could possibly be hoped, for.
I t h i n k t h a t t h e s e t h r e e e x a m p l e s , which could be
m u l t i p l i e d to include p r a c t i c a l l y e v e r y c o u n t r y f o r wMeh
NATURAL DEATH, PUBLIC HEALTH 253
accurate population data exist, furnish a cogent demon-
stration of the essential soundness and accuracy of this
theory of population growth. Indeed, the facts warrant,
I believe, our regarding this as a first approximation to
the true natural law of population growth. We now are
4.388
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POO £0 40 60 60 J&OO £0 40 60 60 1900 2O 40 60 60 2000 2O -40 60 60 2/0O
YEARS
F I G . 6 3 . — S h o w i n g t h e c u r v e o f g r o w t h o f t h e . p o p u l a t i o n of S e r b i a .
a p p r o a c h i n g t h e p r o p e r m a t h e m a t i c a l f o u n d a t i o n o n
w h i c h t o b u i l d s o c i o l o g i c a l d i s c u s s i o n s o f t h e p r o b l e m
o f p o p u l a t i o n .
A s a f u r t h e r d e m o n s t r a t i o n o f t h e s o u n d n e s s o f t h i s
t h e o r y o f p o p u l a t i o n g r o w t h , l e t a t t e n t i o n b e d i r e c t e d f o r
a m o m e n t t o a n e x a m p l e o f i t s e x p e r i m e n t a l v e r i f i c a t i o n .
T o a f r u i t fly { D r o s o p h i l a ) i n a h a l f p i n t m i l k b o t t l e , s u c h
a s i s u s e d i n e x p e r i m e n t a l w o r k o n t h e s e o r g a n i s m s , t h e
i n t e r i o r o f t h e b o t t l e r e p r e s e n t s a d e f i n i t e l y l i m i t e d u n i -
v e r s e . H o w d o e s t h e fly p o p u l a t i o n g r o w i n s u c h a u n i -
v e r s e ? W e s t a r t a b o t t l e w i t h a m a l e a n d f e m a l e fly,
a n d a s m a l l s a m p l e , s a y 1 0 , o f t h e i r o f f s p r i n g o f d i f f e r e n t
a g e s ( l a r v s e a n d p u p a e ) . T h e r e s u l t s a r e s h o w n i n F i g -
254 BIOLOGY OF DEATH
u r e 64. T h e circles give t h e o b s e r v e d p o p u l a t i o n g r o w t h ,
o b t a i n e d by census c o u n t s a t 3-day i n t e r v a l s . T h e r e c a n
b e n o d o u b t t h a t t h i s p o p u l a t i o n h a s g r o w n in a c c o r d a n c e
w i t h t h e equation. T h e t w o final o b s e r v a t i o n s He below
the curve, because of t h e difficulty experienced, i n t h i s
346. IS.
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tzs 7
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F i o . 6 4 . — S h o w i n g t h e growth of a Drotophila nopuUtlon kept under controlled
experimental conditions.
p a r t i c u l a r e x p e r i m e n t , of k e e p i n g the f o o d s u p p l y in g o o d
c o n d i t i o n after s o l o n g a p e r i o d f r o m t h e start.
L e t u s r e t u r n to the f u r t h e r d i s c u s s i o n of t h e p o p u -
the curve.
T h e first q u e s t i o n w h i c h interests o n e is t h i s : W h e n
p o i n t of inflection a n d e x h i b i t a p r o g r e s s i v e l y d i m i n i s h i n g
i n s t e a d of i n c r e a s i n g r a t e of g r o w t h ? I t is easily d e t e r -
m i n e d that this p o i n t o c c u r r e d a b o u t A p r i l 1 , 1 9 1 4 , o n t h e
a s s u m p t i o n that o u r p r e s e n t n u m e r i c a l v a l u e s reliably r e p -
r e s e n t t h e r a t e of p o p u l a t i o n g r o w t h in this c o u n t r y .
I n o t h e r w o r d s , so f a r a s w e m a y rely u p o n p r e s e n t n u -
p e r i o d of m o s t r a p i d p o p u l a t i o n g r o w t h , u n l e s s t h e r e
NATURAL DEATH, PUBLIC HEALTH 255
comes into play some factor not now known, and which
has never operated during the past history of the country,
to make the rate of growth more rapid. The latter con-
tingency appears improbable. The 1920 census confirms
the result, indicated by the curve, that the period of most
rapid population growth was passed somewhere in the
last decade. The population at the point of inflection
works out to have been 98,637,000, which was, in fact,
about the population of the country in 1914.
The upper asymptote given by the equation has the
value of 197,274,000 roughly. This means that the maxi-
mum population which continental United States, as now
areally limited, will have, will be roughly twice the pres-
ent population; provided no fundamental new factor
comes into play in the meantime, different in its magni-
tude and mode of operation from any of the factors which
have influenced population growth in the past. This
state of affairs will be reached in about the year 2,100, a
little less than two centuries'hence. Perhaps it may be
thought that the magnitude of this number is not suffi-
ciently imposing. It is so easy, and most writers on
population have been so prone, to extrapolate population
by geometric series or by a parabola or some such purely
empirical curve, and arrive at stupendous figures, that
calm consideration of real probabilities is most difficult
to obtain. While we regard the numerical results as
only a rough first approximation, it remains a fact that
if anyone will soberly think of every city, every village,
every town in this country having its present population
multiplied by 2, and will further think of twice as many
persons on the land in agricultural pursuits, he will be
bound, we think, to conclude that the country would be
256 BIOLOGY OF DEATH
f a i r l y densely p o p u l a t e d . I t would h a v e a b o u t G6 p e r -
sons p e r s q u a r e mile of l a n d a r e a .
I t will a t once b e p o i n t e d o u t t h a t m a n y E u r o p e a n
c o u n t r i e s h a v e a m u c h g r e a t e r d e n s i t y of p o p u l a t i o n t h a n
66 p e r s o n s to t h e s q u a r e mile, a s , for e x a m p l e , B e l g i u m
w i t h 673, t h e N e t h e r l a n d s w i t h 499, etc. B u t i t m u s t n o t
b e f o r g o t t e n t h a t t h e s e c o u n t r i e s a r e f a r f r o m aelf-
s u p p o r t i n g i n r e s p e c t of p h y s i c a l m e a n s of s u b s i d e n c e .
T h e y a r e , o r w e r e before t h e w a r , economically self-
s u p p o r t i n g , which is a v e r y different t h i n g , because, b y
t h e i r i n d u s t r i a l d e v e l o p m e n t a t home a n d in t h e i r colo-
nies, t h e y p r o d u c e m o n e y e n o u g h to b u y physical m e a n s
of subsistence f r o m lees densely p o p u l a t e d portionB of
t h e world. W e can, of course, d o t h e s a m e t h i n g , p r o -
vided t h a t b y t h e time o u r p o p u l a t i o n g e t s BO dense a s to
m a k e i t n e c e s s a r y , t h e r e still r e m a i n p o r t i o n s of the globe
w h e r e food, clothing m a t e r i a l a n d fuel a r e p r o d u c e d in
excess of t h e n e e d s of t h e i r h o m e p o p u l a t i o n s .
N o w 197,000,000 people will r e q u i r e , on t h e basis of
o u r p r e s e n t food h a b i t s , a b o u t 260,000,000 million c a l o r i e s
p e r a n n u m . T h e U n i t e d S t a t e s , d u r i n g the seven y e a r s
1911-1918, p r o d u c e d a s a n a n n u a l a v e r a g e , in t h e f o r m of
h u m a n food, both primary mid secondary (i.e.f b r o a d l y
vegetable a n d a n i m a l ) , only 137,163,606 million c a l o r i e s
p e r y e a r . So t h a t , u n l e s s o u r food h a b i t s r a d i c a l l y c h a n g e ,
a n d a m a n is able t o do w i t h less t h a n 3,000 t o 3,500 c a l o r i e s
per day, or unless o u r agricultural production radically
increases, which i t a p p e a r s n o t likely t o d o f o r a v a r i e t y
of r e a s o n s which c a n n o t b e h e r e gone i n t o , i t will be
n e c e s s a r y , w h e n even o u r m o d e s t figure f o r t h e a s y m p t o t i c
p o p u l a t i o n is reached, t o i m p o r t n e a r l y o r q u i t e one-half
of t h e calories n e c e s s a r y f o r t h a t p o p u l a t i o n . I t s e e m s
i m p r o b a b l e t h a t t h e p o p u l a t i o n will g o o n i n c r e a s i n g a t
NATURAL DEATH, PUBLIC HEALTH 257
any very rapid rate after such a condition is reached.
East has shown that the United States has already entered
upon the era of diminishing returns in agriculture in this
country. Is it at all reasonable to suppose that by the time
this country has closely approached the asymptote here
indicated, with all the competition for means of sub-
sistence which the already densely populated countries of
Europe will then be putting up, there can be found any
portion of the globe producing food in excess of its own
needs to an extent to make it possible for us to find the
calories we shall need to import?
Altogether we believe it will be the part of wisdom
for anyone disposed to criticize our asymptotic value of
a hundred and ninety-seven and a quarter millions because
it is thought too small, to look further into all the rele-
vant facts. This point of view is sustained in a recent
paper by East in which the future agricultural resources
of the country are particularly examined.
The relation of this already pressing problem of popu-
lation to the problem of the duration of life is obvious
enough. For every point that the death rate is lowered
(or, what is the same thing, the average duration of life
increased) the problem of population is made more imme-
diate and more difficult unless there is a corresponding
decrease in the birth-rate. Is it to be wondered at that
most thoughtful students of the problem of population
are advocates of birth control? Or is it remarkable
that Major Leonard Darwin, president of the Eugenics
Education Society in England, should say in a carefully
considered memorandum to the new British Ministry of
Health: "In the interests of posterity it is most desirable
that parents should now limit the size of their families
by any means held by them to be right (provided such
17
258 BIOLOGY OF D E A T H
m e a n s a r e not i n j u r i o u s to health, n o r , like* a b o r t i o n , ai
offense a g a i n s t public m o r a l s ) to such an e x t e n t t h a t the
children could be b r o u g h t u p a s efficient citizens and with-
out d e t e r i o r a t i o n in t h e s t a n d a r d s of t h e i r civilization:
and t h a t p a r e n t s should n o t limit the size of the familv
for any other r e a s o n s except on account of definite hered-
i t a r y defects, or to secure a n a d e q u a t e interval between
births."
I a m able to m a k e no prediction a s to how civilized
countries will solve (if they do solve) the problems
a r i s i n g out of the i m p e n d i n g s a t u r a t i o n with h u m a n popu-
lation of t h e p o r t i o n of the e a r t h ' s s u r f a c e habitable by
m a n . T h e c e r t a i n t y a n d a s s u r a n c e with which v a r i o u s
ones of m y friends a d v a n c e solutions excites my wonder
a n d a d m i r a t i o n . B u t w h a t i m p r e s s e s me even m o r e
is t h a t scarcely any two of t h e m a g r e e on the n a t u r e
of the p a n a c e a . T o some it is b i r t h control, to other*
synthetic foods d e r i v e d from the a t m o s p h e r e or else-
where, and so on-
F o r myself, I a m content if I h a v e succeeded, hi avert
a small m e a s u r e , in i n d i c a t i n g t h a t population g r o w t h p r e -
sents a problem fast becoming u r g e n t ; a p r o b l e m that
in its overwhelming significance and almost infinite rami
fications touches upon v i r t u a l l y every p r e s e n t h u m a n ac-
tivity and i n t e r e s t , and in p a r t i c u l a r upon the activitif**
comprised in t h e t e r m s public health and hygiene.
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the periodicity of endomixis in Paramecium aurelia. Biol. Bui. Vol.
33, pp- 437-462, 1917.
268 BIBLIOGRAPHY
WOODRUFF, L. L. The present status of the long-continued pedigree culture
of Paramedum aurelia at Yale University. Proc. Nat Acad. Mci.
Vol. 7, pp. 41-44, 1921.
WOODRUFF, L. L. Amicronucleate infusoria. Proc. Boc. Exp. Hiol. Mcd.
Vol. 18, pp. 28-29, 1921.
WOODRUFF, L. L. and ERDMANN, K. A normal periodic reorganization pro-
cess without cell fusion in Paramecium. Jour, fisoper, Zool. Vol. 17,
pp. 425-518, W14.
WOODS, F. A. Origin and migration of germ-cells in Acanthiaa. Amcr,
Jour. Anat. Vol. I, p. 307.
YOUNG, T. E. On Centenarians j and the Duration of the Human Race.
London, 1905 (Keissue) pp. 147.
I N D E X
Acanthia8, 38 Bibliography, 259-268
Accidental deaths, 107 Bills of mortality, 79
Activity, metabolic, 211-217 Biological classification of causes of
Agamic reproduction, 33, 35, 37, 41, death, 104-137
77 Birds, longevity of, 22, 63
Alchemy, 19 Birth control, 257
Alimentary tract, 107, 108, 110, 112, injuries at, 121, 123
129-131 premature, 121, 123
Amicronucleate races, 72, 73 Blood, 107, 108, 111, 118, 119
Amma, K., 39, 259 Body size and longevity, 26, 68
Amphibia, longevity of, 22 weight, 68
Analysis of life tables, 94-101 Bogdanow, E. A., 200, 259
Animals, longevity of, 22, 68 Brain weight, 68
Anopheles cruzians, 239^ Brazil, 106
punctipennis{ 28£ Bright's disease, 161
quadrimaculatus, ~239 Bronchitis, 231, 232
Anti-vivisection, 238 Brown, J. W., 206, 261
Apple trees, 37, 74, 75 Brownlee, J., 183
Artificial parthenogenesis, 51-58,223 Budding, 37
Ascoritf, 39 Bulloch, W., 259
Aseptic life, 43, 200-202 Burrows, M. T., 59-62, 260
Astrology, 19
Australia, 235 Callithria;, 68
Austria, 235 Calories, 256
Autogamy, 73 Cancer cells, 61
Automobiles, 226 Carrel, A., 10, 61-65, 73, 74, 76-78,
224, 260
Bacteria, rOle of, in duration of life, Cat, 61
43, 199-202 Cattell, J. McK., 10
Bataillon, 52, 259 Causes of death, 102-137
Bavaria, 244, 245 biological classifi-
Bee, senility in, 28 cation of, 104
Beeton, M., 166, 169, 171, 259 international clas-
Belgium, 256 sification of, 103
Bell, A. G.,* 152-158, 165, 166, 225, non-controlled,
259 232, 233
Benedict, H. N., 44, 259 Cells, interstitial, 219
Bertillon, J., 216, 259 Cellular immortality, 51-78
269
270 INDEX
Centenarians, 23-26, 04 Dawson, M. M., 82, 200
Cephalic index, 175 Death, appearance of in evolution, 42
Cephalisation, 68 biological clarification of
Chances of death, 70-101 cauites of, 104-137
Changes in expectation of life, 82-04 chancei of, 79-101
Chick, 59-61, 76 causes of, 102-137
duration of life of, 63 the Mftrknman, 96-98
Child, C. M;, 34-36, 39, 43, 44, 260 theories of, 43-150
Child welfare, 112 Death birth ratio, 243-246
Chironomu8, 39 Death-rate, selective, 177-1H5
Circulatory system, 107, 108, 111, Death-rates, crude, 112
118, 119 «p«*dftef 112-137
Classification, biological, of causes DeDeeker, A., 216, 267
of death, 104 Deer, 68
international, of Delage, Y., 45, 260
causes of death, 103 Iteleourt, A., 200, 260
Clocks, analogy with living things, Descent, method of, 40>42
150, 151, 198 Diarrha*, 110, 112
Clonal reproduction, 37, 74 Differentiation, 45-47, 67, 75
Coefficient of correlation, 168 Diphtheria, 230, 231, 235, 237
Coelenterates, 62 Diabases, prevent ability of, Wl
Cohnheim, J., 44, 260 Doflein, F., 33, 260
Collis, E. L., 216, 260 Dog fteh, 39
Conjugation, 30-33, 71, 73 Dommtlc fowl, duration of lif# of, <$3
Conklin, E. G., 29, 44, 260 Donaldson, H. II, 20, tm
Controllable causes of death, 230,233 Drosophila mtUinopaMter, 180*202,
Correlation coefficient, 168 208-211, 214, 222, 225, 22S, Z&%,
Correlations in duration of life, 168- 2f>4
177 Dublin, U I., 82, 113, 260, 261
Crossett, 239, 241 du No«y, l\ L., 77
Croup, 230, 231 Duration of life, corn*!ntion in, 16B-
Crura, F. 8., 184, 260 177
Culture of tissues in vitro, 58-78
Curve, mortality, graduation of, 94- of, im^tt
101 \nfimtm* of activi-
specific death rate, 114, lltt ty mi, 2U-217
Curves, logarithmic plotting of, 114 of t«-ni»
Cyclops, 39 t on, 20B-
Cytomorphosis, 28 217
Cytoplasm, 29, 44 inh<»ritanc«i of, M,
Darwin, L., 257 imim
Davis, W. H., 113 In man, 7D-04, 150.
Dawson, J. A., 73, 260 185
INDEX 271
Duration of life of domestic fowl, 63 Expectation of life, effect of selection
of parents and off- on, 94
spring, 155-157 hypothetical, 164
rOle of bacteria in, in ancient Egypt,
43, 199-202 87-89
variation in, 21, 22, in ancient Home,
68, 80-82 90-92
Dysentery, 230, 231 in Hispania and
LuHitania, 91-
East, E. M., 195, 257, 261 92
Ebeling, A. 1L, 60, 61, 74, 76, 77, 78, in Roman
260, 261 Africa, 92-93
Ectoderm, 138-149 Experimental atudy of duration of
Effects of public health work, 112, life, 186-222
227-242 Eye color, 174, 175
Egypt, expectation of life in, 87-89
Elephant, longevity of, 22 Ferxnat, 82
Embryology and mortality, 138-149 Fertiltain, 67
Embryonic juice, 74 Finn, longevity of, 22
Endocrinal system, 107, 108, 112, FiHher, A., 101, 149, 184, 261
133, 134 PiHher, I., 161, 162, 165
Endoderm, 138-149 Kiftiiion, 32, 33, 35, 40, 41
Endomixis, 30, 33, 71-73 Fitting the mortality curve, 94-101
Energy, 213-217 Food requirements, 256
England, 106, 108-111, 139, 140, 235, Fowyth, C. H., 161, 164, 261
244-246 Fowl, duration of life of, 63
Enriquea, P., 73, 261 France, 244, 245, 251, 252
Environment, 2Z$t 226 Franco-Prussian war, 252
Epidemic, influenza, 245 Fraternal correlations, 171,172,175,
ETdrnan, K., 30, 2111, 268 176
Eudorvna elegant, 31, 73 Frkdenth&t, H,, 68, 69, 261
Eugenics, 227 Friends' Provident association, 167
Education Hociety, 257 Frog, 52, 58, 59
Evolutionary program in longevity,
87-94 Galvani, 68
Evolution of eetodtrm, 141 Genealogy of Hyde family, 152
of en<l<Ki<rm, 141 Genetic variation, 190
of mm0dmmf 141 Germany, 235
of workmanship of, 148 Orm cell*, 37-42, 51-58
Excretory organ*, 107, 108, 111, 120, layeru, 138
127 plasm, 227, 228
Exercise, 212, 213 Given, D. H. CL, 238, 261
Expectation of life, <Ufined, 82 Gland, pituitary, 220-222
change* in, 82-04 Glands, puberty, 217-210
272 INDEX
Glaucoma pyriformi8t 73 Immortality, cellular, 51-78
Glover, J. W., 80, 84, 88, 90-92, 201 human, 17-20
Gonads, 217-219 of protozoa, 30-33, 64
Gonococcus infection, 123, 124 of aomatic cells, 58-78
Graduation of mortality curve, 94- Industrial mortality, 216
101 Infant mortality, 205, 206, 208
Grafting, 37 Influence of activity on duration of
Graunt, J., 79 of life, 211-217
Greenwood, M., 205, 216, 259-261 of poverty on mortality,
Groth, 205, 261 202-208
Growth of Drosophila population, of fterum on tiftnue cul-
254 ture, 76, 77
of populations, 247-258 of temperature on dura-
Growth of United States, 250-252, tion of life, 208-217
254-257 Influenza epidemic, 245
Guayaquil, 240, 242 Inheritance of duration of Iif«», 1M
Guinea pig, 01 in Vrontt-
Guydnot, E., 200, 260, 261 p h i I n,
Guyer, M. F., 52, 262 IM-im
in m a n,
Hahn, 205, 261 150-1KA
Halley, R, 81, 82, 84, 262 of
Harper, M., 39, 262 174, 175
Harrison, R. G., 58-60, 63, 64, 224, at birth, VZh 1^3
262 In«i*ctH, long«»vity of, 22
Hartman, M., 31, 73, 262 International cla*wifiattion of
Heart muscle, 61 of death, 103
Hegner, It. W., 40, 262 Health Board, TM*
Henderaon, R., 99, 262 240, t\t
Heron, D., 206, 262 Intcrntitiftl cdl«f 210
Hersch, L., 202, 203, 205, 206, 208, In vertebra ten, longevity of, 22
262 In vitro culture of ti»*ti<**, f»H 7H
Hertwig, R., 44, 263 Italy, 23f>
HiHpania and LiiHitania, expectation Jamaica, 235
of life in, 91-02 Jenninxti, H. 8., 31, 3.1, 40, 41, M,
Hodge, C. F., 27, 28, 263 71? 72, 203
Holland, 184 i, C. F.f 44, tU
Homicide, 107
Homoiotoxin, 64
Howard, VV. T., 44, 263 Jonm, IX F.r
Hyde family, 152-166 t M,, -H, 203
Hyde, R. R., 108, 225, 263 Krimbahn, 40
Hygiene, 227 Kidneys 01, 107, 10H, 111, 120,
INDEX 273
Kopf, E. W., 113, 260 Metchnikoff, E., 43, 109, 200, 264
Korsclielt, E., 263 Method of descent, 40-42
Micronucleus, 72
Landed Gentry, 167, 169, 172 Minot, C. S., 27, 28, 44, 71, 264
Lankaster, E. R., 263 Mitchell, P. C, 264
Levasseur, E., 82, 263 Mitosis, 61
Legrand, M. A., 263 Montgomery, T. H., 44, 265
Lewis, M. R., 62, 263 Morgan, T. H., 10, 186, 197, 265
Lewis, W. H., 53, 54, 62, 263, 264 Mortality, bills of, 79
Life, aseptic, 43, 200-202 curve, graduation of, 94-
changes in expectation of, 82-94 101
curve of Hyde family, 153 embryological basis of,
cycle of Drosophila, 187, 188 138-149
prolonging, 17, 54, 218, 221 industrial, 216
table, 79-82 infant, 205, 206
analysis of, 94-101 influence of poverty on,
Breslau, 83, 84, 92 202-208
Carlisle, 83, 86 organ system in, 107,108
U. S., 1910, 83-86 Mosquito, 239, 240
Lillie, F. R., 57, 263 Most fatal organ systems, 136
List, International, 103 Mouse, 68, 220-222
Locomotor ataxia, 124 growth of, 69-70
Loeb, J., 47, 52-55, 57, 200, 201, 208- Miihlmann, M., 44, 265
211, 214, 215, 223, 226, 263, 264 Muller, J., 44
Loeb, L., 59, 64, 65, 67, 224, 264 Miiller, L. R., 265
Logarithmic plotting, 114 Muscular system, 107, 108, 112, 127,
London, 205-208 128
Longevity, body size and, 26
evolutionary progress in, Nascher, L, 26, 27, 265
87-04 Nerve cells, senile changes in, 27-29
of animals, 22 Nervous system, 107, 108, 130, 131
of parents, 158, 160 Netherlands, 256
Lowell Institute, 9, 27 Non-controlled causes of death, 232,
233
Macdonell, W. R., 87, 89-93, 229, 264 Northrop, J. H., 200, 201, 209-211,
Malaria, 238-241 214, 215, 226, 264, 265
Malthus, T. R., 243 Nucleus, 29, 30, 44
Mammals, longevity of, 22
Man, longevity of, 23-26, 80-94 Occupation, 216
Marmoset, 68 Ogle, W., 95
Mendelian inheritance, 194, 197, 198 Orbits, 250
Mesoderm, 138-149 Organ systems in mortality, 107,108
Metabolic activity, 211-217 most fatal, 136
Metazoa, 31, 33, 40, 46, 71 Oxytricha hymenostoma, 73
274 INDEX
Paralysis, 231, 232 Quaker record*, 171, 17.1
Faramccium, 30-32, 35, 40, 72
Parental correlations, 171, 172, 174, Rabbit, 68
176 Hut, 61, 212, 213, 21H
Parents and offspring, duration of Ratio, death-birth, 243-240
life of, 155-157 Kay, L. A., «9, 70, 220, 221, 266
longevity of, 158, 160 Reed, L. J., 247, 240, 266
Paris, 202-206 Ri»gi«tration Area, U. R, IO6, 108,
Parr, T., 24 100, 139, 140, 164* 2*29, tW, 246
Parthenogenesis, artificial, 51-58, 223 Reproduction, organic* 33, 41
Pascal, 82 by budding, #7
Pearl, R., 106, 201, 225, 249, 265 byftfttrion,32, 33, 41
Pearson, K., 19, 87-91, 03-101, 165, cloiml, 37
169-177, 179, 182, 183, 225, 229, uexuai, 37-40, 41
238, 259, 266 Reptile, longevity of, 22
Peerage, 167, 169, 172 Respiratory »y«tem, 107, 108, 110,
Pennaria, 62 112, 119, 120, 136, 1S7
Physical characters, inheritance of, Results, summary of, 225*227
174, 175 Richards, H. A., 86, 87, 2«0
Pituitary gland, 220-222 Ritter, W. K., 75, 266
Pixell-Goodrich, Mrs., 28 Robertnon, T. B., 6ft, 70,220,221,206
Planaria dorotocephala, 34, 35 Rockefeller Foundation, 2.18
Plants, senility in, 44 Intstitut*, 62, 61
Ploetz, A., 178, 179, 182, 183, 225, Rdle of bacteria In duration of lift*,
266 43, 199-202
Population, 240-258 Roman Africa, expectation *>f Hfn
Potassium cyanide, 53, 54 in, 92
Poverty, 202-208 Rome, expectation of life In, 90-92
Premature birth, 121, 123 Romei«, B., 219, 206
Preventability of diseases, 162 Rose, W., 238, 230, 241, 266
Pritchett, A. 8., 247, 266 Roumania, 235
Progress, evolutionary, in longevity, Roundworm, 39
87-94 Royal families, 177
Prolonging life, 17, 54, 218, 221 Rubner, M., 21S, 214, 226, 267
Prostate, 126, 219
Protozoa, 30-33, 40, 41, 46 Saleeby, 183
immortality of, 30-33, 41, Sanitation, 227, 235
64, 71 Bao Paulo, 106f 108-111, 139, 140
Prussia, 244, 245 Sea urchin, 52, 54, 57
Puberty glands, 217-219 Selection, effect of, on exptcUtion of
Public health work, effects of, 112, life, 94
227-242 Selective death rate, 177-185
Purulent infection, 231, 232 Seneca, 102
INDEX 275
HeneKcenee, 27-30, 4o\ 70-78 Table, life, 79-82
theorie* of, 43-50 Temperature, 208-217
Senile changes in nerve cells, 27-2D Tethelin, 70, 220-222
Senility as eaune of death, 100 Theories of death, 43-50
in plants, 44, 74, 75 Theory of population growth, 249
Septicaemia, 231, 232 Thyroid gland, 61
Serbia, 252, 253 TiHsue culture in vitro, 5H-78
Serum, influence on tiH*m* culture, Transplantation of tumors, 64, 05
76, 77 TutmrcukmiB, 161, 204, 208, 230, 2:U,
Sex organs, 107, 108, 111, 121-125, 238
217-219 Tumor tranHplaniation, 64, 65
Sexual reproduction, 37-41 Typhoid fever, 230, 231, 235, 2.'W
Shell, J., 26, 27
Skeletal nyHtem, 107, 108, 112, 127, United States, growth of, 250-252,
128 254-257
Skin, 107, 108, 110, 112, 131, 132 Urostyla grantlis, 72
Klonaker, J. H., 212, 213, 218, 228,
267 Van Buren, G. H., 113, 260
Klotnpohtki, BM 33, 267 Variation, genetic, 190
Snowf E. C, 179-183, 225, 267 Venereal diabases, 123, 124
Softening of th« hrain, 231 , 232 VerhulHt, I\ F., 249, 267
Soma, 40 Verworn, M., 44, 267
Somatic cells, immortality of, 58-78 Vienna, 245, 246
Span, 174, 175 Voronoff, 217
HpiegelWg, W.t 87
Spiritualmm, 18-20 Waller, A. I)., 216, 267
Kple<?n, 61 Waiworth, H. H., 152, 267
Sponge*, 02 War, 243
Stature, 174, 175 Wedekind, 33, 267
Htelnach, E., 217-210, 207 Weiamann, A., 26, 43, 65, 267
Btenoatomum, 35, 36 Whale, longevity of, 22
Btnvunnon, T. H. C., 206-208, 267 Wilton, H. V., 62, 267
Still hirthn, 205 Wittatein, 09
8trongylocentrotu$ purpuratus, 38, Woodruff, U L.t 30, 33, 72, 73, 267,
56 268
Sttmmary of renultw, 223-227 Woods, F. A., 38, 39, 268
gurvivornhip Hum of DromphHa,
188, 102, 195 Ydlow fever, 240, 242
Syphilis, 123 Voting, T. E., 23-25, 268