MONOGRAPHS ON EXPERIMENTAL BIOLOGY

T H E B I O L O G Y O F

D E A T H

Being a Scries of lectures Delivered at the Jewell Institute

in Boston in December 1920

BY
R A Y M O N D P E A R L
TMR JOHNS tmvKtm vmvumtrt

PHILADELPHIA AND LONDON

J. B. LIPPINCOTT COMPANY
COPYRIGHT, 1Q22, BY J. B. LJPPINCOTT COMPANY

PRINTBP BY J. B. LIPPINCOTT COMPANY

AT THB WASHINGTON SQUAJLE FKSSS
PHILADELPHIA, U. 8, A,
 TO
MY WISEST COUNSELLOR
 EDITOR'S ANNOUNCEMENT

T H E rapidly increasing specialization mak<! it im-

possible for one author to cover s a t i s f a c t o r i l y thci whola
field of modern B i o l o g y . T h i s situation, which e x i s t * in
all the sciences, has induced Englieh a u t h o r s t o itiBUc*
series of m o n o g r a p h s in Biochemistry, P h y s i o l o g y , and
Physics. A number of American biologists h a v e decided
to provide the s a m e opportunity for the study of
Experimental B i o l o g y .
B i o l o g y , which not l o n g a g o w a s purely dencriptivo
and speculative, has begun to a d o p t the method** of the
exact sciences, recognizing that for p e r m a n e n t p r o g r a m
n o t only e x p e r i m e n t s a r e required but t h a t the* experi-
ments should be of a quantitative character. It will \w
the purpose of tins s e r i e s of m o n o g r a p h s to vrnphnniw
and further a s much a s possible this d e v e l o p m e n t of
Biology.
E x p e r i m e n t a l B i o l o g y and General P h y s i o l o g y a m one
and the s a m e science, by method it wall an by c o n t e n t s ,
since both aim a t explaining life from the phyntco-ehomtcftl
constitution of living matter. T h e aerie* of m o n o g r a p h s
on E x p e r i m e n t a l B i o l o g y will therefore include the fluid
of traditional General Physiology.
JACSQUKII L o t f
T 1L MOMUUI,
W , J . V . OUTKIIIIOUT.
 AUTHOR'S PREFACE
I N preparing the material of a series of lectures, given
at the Lowell Institute in Boston in December 1920, for
book publication, I have deemed it on the whole best to
adhere rather closely to the original lecture mode of pre-
sentation with all its informality. E x c e p t for the fact
that the matter is here set forth in somewhat greater
detail than was possible under the rigid time limitations
of the Lowell Institute, and that the breaking into chap-
ters is slightly different, the whole is substantially as it
was presented in Boston.
What I tried to do in these lectures w a s to bring
together under a unified viewpoint some of the more im-
portant contributions which have been made to our know-
ledge of natural death, from three widely scattered
sources: namely general biology, experimental biology,
and statistical and actuarial science. I t will be obvious
to anyone who knows the literature from these fields
regarding natural death and the duration of life that in
such an amount of space as is here used, no one could
hope to cover a field so wide with anything approaching
completeness. To do so would require a series of volumes
in place of one small one. B u t this has i n no wise been
my object; I have instead hoped that the very incomplete-
ness itself of this work, necessitated by m y limitations
of space and knowledge, might stimulate the reader to
penetrate for himself further into the literature of this
fascinating and important field of biology. To help him
to start upon this excursion a brief bibliography is
appended. I t by no means completely covers the field,
but may perhaps serve as an introduction.
9
10 AUTHOR'S P R E F A C E
I am indebted to a number of authors and publishers
for permission to use illustrations and wish here to ex-
press my great appreciation of this courtesy. The indi-
vidual sources for these borrowed figures are in every
case indicated in the legends. To Dr. J. McKeen Cattail
I am especially grateful for allowing me the* une of the
blocks from the magazine publication of thin material in
the Scientific Monthly; to Dr. Alexis Carrel for permis-
sion to use unpublished photographs of his tiAHUO culture*;
and, finally, to Professor T. H. Morgan for critically
reading the manuscript and making many helpful
suggestions.
B . P.
BALTIMORE,
April 19, 1022.
 CONTENTS

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

PROBABLY n o subject so deeply i n t e r e s t s h u m a n beings

a s t h a t of t h e d u r a t i o n of h u m a n life. P r e s u m a b l y j u s t
because t h e business of living w a s such a wonderfully
i n t e r e s t i n g a n d impo.rtant one f r o m t h e viewpoint of t h e
individual, m a n h a s endeavored, i n e v e r y w a y he could
think of, t o p r o l o n g i t a s m u c h as possible. H e h a s h a d
recourse t o b o t h n a t u r a l a n d s u p e r n a t u r a l schemes for
a t t a i n i n g t h i s objective. On t h e m u n d a n e p l a n e h e h a s
developed t h e sciences a n d a r t s of biology, medicine a n d
hygiene, w i t h t h e f u n d a m e n t a l p u r p o s e of l e a r n i n g the
u n d e r l y i n g p r i n c i p l e s of v i t a l p r o c e s s e s , so t h a t i t m i g h t
ultimately b e possible t o s t r e t c h t h e l e n g t h of each indivi-
d u a l ' s life on e a r t h t o t h e g r e a t e s t a t t a i n a b l e d e g r e e .
Recognizing p r a g m a t i c a l l y , however, t h a t a t b e s t t h e limi-
tations in t h i s direction w e r e distinctly n a r r o w , w h e n
conceived i n a n y h i s t o r i c a l sense, h e h a s w i t h s i n g u l a r l y
w i d e - s p r e a d u n a n i m i t y , deemed i t wise t o seek a n o t h e r
means of s a t i s f y i n g his desires. M a n ' s b o d y p l a i n l y a n d
palpably r e t u r n s to dust, a f t e r t h e b r i e f e s t of i n t e r v a l s ,
m e a s u r e d i n t e r m s of cosmic evolution. B u t , p a t e n t as
this fact is i t h a s n o t precluded the p o s t u l a t i o n of a n infin-
ite c o n t i n u a t i o n of t h a t i m p a l p a b l e p o r t i o n of m a n ' s be-
ing whieh i s called t h e soul. W i t h t h e field t h u s open we
2 17
18 T H E BIOLOGY O F D E A T H
see some s o r t of notion of i m m o r t a l i t y i n c o r p o r a t e d in
an integral p a r t of almost all folk philosophies of which
any record exists.
Now, p e r h a p s unfortunately, p e r h a p s f o r t u n a t e l y , it
has u p to the p r e s e n t time p r o v e d impossible absolutely
to demonstrate, for r e a s o n s which will p r e s e n t l y a p p e a r ,
by any scientifically valid method of e x p e r i m e n t a t i o n or
reasoning, t h a t any r e a l p o r t i o n of t h a t t o t a l i t y of being
which is an individual living m a n p e r s i s t s a f t e r he dies.
Equally, for t h e same reasons, science cannot absolutely
demonstrate t h a t such persistence does n o t occur. The
l a t t e r fact h a s h a d two i m p o r t a n t consequences. I n the first
place, it h a s p e r m i t t e d m a n y millions of people to derive
a real comfort of soul in sorrow, a n d a f a i r l y abiding t r a n -
quility of m i n d i n general from the belief t h a t i m m o r t a l i t y
is a reality. E v e n the m o s t cynical of scoffers can find lit-
tle fault with such a result, t h e w o r l d a n d h u m a n n a t u r e
being constituted as they a r e . T h e other consequence of
science's p r e s e n t inability t o lay b a r e , i n final a n d i r r e -
fragable t e r m s , the t r u t h about t h e course, if any, of
events subsequent to death is m o r e serious. I t opens the
w a y for r e c u r r i n g m e n t a l epidemics of t h a t i n t i m a t e mix-
t u r e of hyper-credulity, h y p e r - k n a v e r y , a n d mysticism,
which used t o be called spiritualism, b u t now u s u a l l y p r e -
fers m o r e seductive titles. W e a r e a t t h e m o m e n t i n the
midst of p e r h a p s t h e m o s t violent a n d d e s t r u c t i v e epi-
demic of this s o r t which h a s ever occurred. I t s evil lies in
the fact t h a t i n exact p r o p o r t i o n t o i t s virulence i t des-
t r o y s t h e confidence of the collective m i n d of h u m a n i t y
in the e n d u r i n g efficacy of the only t h i n g which t h e h i s t o r y
of mankind h a s demonstrated t o c o n t r i b u t e to t h e r e a l
advancement of his intellectual, physical, s p i r i t u a l a n d
m o r a l well being, namely t h a t o r d e r l y p r o g r e s s i o n of
ascertained knowledge which we n o w call science.
 THE PROBLEM 19
T h e r e a s o n w h y science finds itself helpless t o p r e -
vent s p i r i t u a l i s m ' s insidious s a p p i n g of t h e intellectual
fiber of the r a c e is because i t is asked to p r o v e a n e g a t i v e ,
u p o n t h e b a s i s of u n r e a l d a t a . H o w difficult such a t a s k
is is obvious a s it is p r o v e r b i a l . U n t i l science h a s demon-
s t r a t e d t h a t t h e r e is not a continuation of i n d i v i d u a l
s u p e r n a t u r a l existence a f t e r n a t u r a l d e a t h , t h e s p i r i t u a l -
ist can, a n d will, come f o r w a r d w i t h s u p p o s e d d e m o n s t r a -
tions t h a t t h e r e is such a continuation. B u t t h e m o s t
characteristic f e a t u r e of science is its actuality, its reality,
its n a t u r a l i t y . P e a r s o n h a s pointed out, i n c h a r a c t e r i s t i -
cally clear a n d v i g o r o u s l a n g u a g e , t h e r e a s o n why, in the
minds of u n i n f o r m e d p e r s o n s , science a p p e a r s helpless in
this situation. H e s a y s :
Scientific ignorance may either arise from an insufficient classification
of facts, or be due to the unreality of the facts with which science has been
called upon to deal. Let us take, for example, fields of thought which
were very prominent in medieval times, such as alchemy, astrology, witch-
craft. In the fifteenth century nobody doubted the "facts" of astrology
and witchcraft. Men were ignorant as to how the stars exerted their
influence for good or ill; they did not know the exact mechanical process
by which all the milk in a village was turned blue by a witch. But for
them it was nevertheless a fact that the stars did influence human lives,
and a fact that the witch had the power of turning the milk blue. Have
we solved the problems of astrology and witchcraft today?
Do we now know how the stars influence human lives, or how witches
turn milk blue? Not in the least. We have learnt to look upon the facts
themselves as unreal, as vain imaginings of the untrained human mind;
we have learnt that they could not be described scientifically because they
involved notions which were in themselves contradictory and absurd. With
alchemy the case was somewhat different. Here a false classification of
real facts was combined with inconsistent sequencesthat is, sequences
not deduced by a rational method. So soon as science entered the field
of alchemy with a true classification and a true method, alchemy was con-
verted into chemistry and became an important branch of human knowl-
edge. Now it will, I think, be found that the fields of inquiry, where
science has not yet penetrated and where the scientist still confesses
20 T H E BIOLOGY OF D E A T H
ignorance, are very like alchemy, astrology, and wJt^herafl nt fh# Middle
Ages Either they involve fact* which are in thwnaetiM* tmrrml-~*on.
ceptions which are self-contradictory and ahMird. ami thfrHorit in^puhlt
of analysis by the scientific or any other mi*><xi or, m th* <*!hi*r hum!,
our ignorance arises from an inadequate eJafiaiftcat itm tnd * nt*!*?! nf
scientific method.
This is the actual state of the atm with than m*nt*t imf npiriitmf
phenomena which are said to lifc outnide the proper m**p* of trtvnrt}, or
which appear to be disregarded by etontlfit; **. No 1*1 !*r #iamj>l#
can be taken than the range of phenomena which mm en litto!Hpfoitimttnm,
Here science is asked to analyse a mrim of !*<* whfch tir* i** n gr*>*i piii*nt
unreal, which arise from the vain imagining* vi tiufrainwt mindu Afwl
from atavistic tendencies to superstition. Bo far *# th* fpt# r# of this
character, no account can be givtn of them, tarfttiw, Uk thw wHeh'
supernatural capacity, their unreality will tm iaund t Imttom t mak#
them self-contradictory. Combined, howtv^r, with th* \wtm\ mrlm f
facts are probably others, conntctd with hypnotic ntl ofhrr ^nditiniii,
which are real and only incomprlinsibl WTJIWIM? thur^ U n y^l wrmrmty
any intelligent classification or truo AppiieHtlinn u\ mhnUftp mttJiudL Th#
former class of facts will, Uk antrology, i?r tm rdur4 to tiiw, fet will
one day be recognised as abnurd; thu tithtr, liki mkhmny, mnf %rtm iltp
by step into an important branch of wlem*, Whiiw^i>r4 tbi^rtfor*, w
are tempted to desert the scientific rotthud f mkln% it nth, wkmm^r llitt
silence of science suggests that om ui\mr pH#wny mit lit Q|fht to
knowledge, let us inquire first whithtr f h #lt*#l# nf fhn prubli*m, #f
whose solution we are ignorant, nmy not nfttr nit! Uk flit fi #1 wit<?fe
craft, arise from a superstition, and to mUmnUmAMmf fi4 iM^mpr<
hensible because they are unreal.
Let us recapitulate briefly our diteusnion tci thin pmut
Mankind has endeavored to prolong thu individual lif by
natural and by supernatural means. Thia latter plan
falls outside the present purview of th# acientifio mtthoci
The former is, in last analysis, roapoiuribla for a eonttd-
erable part^ at l e w t , of the development of the sekitCMi
of biology, pure and applied, and t k i artii wbieh foiled
their operations upon i t Biology can and haa eontrilmted
much to our knowledge of natural death and the eatifee
which determine the duration of life* I t ie the p w r p o i t
of this book to review some of th more important aaptcHte
 THE PROBLEM 21
of this phaHc of biological science, a n d tmdcuivor t o
forth in a n o r d e r l y a n d conniHUmt m a n n e r t h e pn*m*nt
s t a t e of k n o w l e d g e of t h e m i h j e e t
T h e p r o b l e m of n a t u r a l d e a t h han t w o mtpftttn, o n e
general, t h e o t h e r t*peeial. T h c n e m a y be HUIUHI in
this w a y :
1. W h y d o l i v i n g thingn dioT W h a t i s t h o m e a n i n g
of d e a t h in tho g e n e r a l pltilonophy of biology I
2. W h y d o l i v i n g thing*? d i e when t h e y ilof Whut
factors d e t e r m i n e tho d u r a t i o n of life* in g e n e r a ! a n d in
p a r t i c u l a r , a n d w h a t m t h e r e l a t i v e inflwmm of e a c h of
these f a c t o r s i n p r o d u c i n g t h e o b s e r v e d r e n u l t f
B o t h of t h e s e p r o b l e m s h a v e been t h e s u b j e c t of mtieh
speculation a n d d i s c u s s i o n . T h e r e him mummuhtiml,
especially in r e c e n t y e a r n , a considerable* a m o u n t of n e w
experimental a n d atatiHtieal d a t a bi^aritig u p o n t h e m . I
hope to be a b l e in w h a t follown t o nhow t h a t thi w*\v
material^ t o g e t h e r w i t h t h a t which ha for a Jong t i m e
l>een a p a r t of t h e c o m m o n Ktoro of biologtenl knowleiigr%
makes possible a c l e a r e r nml m o r e logiciilly eoimttr*r$t
picture t h a n w e h a v e hail of t h e m e a n i n g of d e a t h a n d
the d e t e r m i n a t i o n of l o n g e v i t y . L e t UH flmt eitamiite tfi
brief review t h e b r o a d generaltftaitonit nlmxt d e i i t h wlttf h
have grown u p in the* e o u r e of t h e d e v e l o p m e n t of b i o l o g y ,
and which m a y n o w b e r e g a r d e d an a g r e e d fr? by p r a c t i -
cally all biologists.

BIOLOCIICAX* aunmiAUZArmsH Aiiotit n^nntAh tmiru

T h e significant g e n e r a l (net which art* k n o w n nlnmi

natural death a r e thane:
( A ) . There i$ an enormowt variation in thr, duration
of Ufe, both intra and inter-racially. T a b l e I, w h i c h Is
a d a p t e d f r o m v a r i o u s a u t h o r i t i e s , is t o bu read w i t h t h e
22 T H E BIOLOGY O F D E A T H
u n d e r s t a n d i n g t h a t t h e figures a r e estimates, frequently
based u p o n somewhat general a n d inexact evidence, and
record extreme, t h o u g h i t is believed authentic instances.
While t h e figures, on t h e accounts which h a v e been men-
tioned, a r e subject to l a r g e p r o b a b l e e r r o r s , t h e table does
give a sufficiently reliable g e n e r a l picture of t h e t r u t h
to indicate t h e enormous differences which exist a m o n g
different f o r m s of animal life in respect of longevity.
TABLE 1
Longevity of Animals
Animal Approximate limits of maximum duration
of life in different species
Lower invertebrates Under 100 hours to ?
Insects Under 100 hours to 17 years
Fish ? to 267 years
Amphibia ? to 36 years
Reptiles ? to 175 years
Birds 9 years to 118 years
Mammals 1J^ years to over 100 years
W e see from this table t h a t life m a y endure i n differ-
ent forms from only the briefest period, m e a s u r e d i n
hours as in the case of Ephemeridae, to somewhere i n
the h u n d r e d s of y e a r s . The extremely l o n g d u r a t i o n s
are of course to be looked u p o n w i t h caution a n d r e s e r v a -
tion, but if we accept only extreme cases of known d u r a -
tion of life i n m a n , t h e r a n g e of v a r i a t i o n i n t h i s
characteristic of living things is sufficiently wide.
I t is probable t h a t m a n , i n exceptional instances, is
nearly the longest lived of all m a m m a l s . T h e common
idea t h a t whales a n d elephants a t t a i n g r e a t longevity
a p p e a r s t o be n o t well founded. T h e absolutely authentic
instances of h u m a n survival beyond a century a r e , con-
t r a r y to the p r e v a l e n t v^ew a n d c u s t o m a r y statistics,
extremely r a r e . T h e m o s t p a i n s t a k i n g and a c c u r a t e
 T H E PROBLEM 23
i n v e s t i g a t i o n of t h e f r e q u e n c y of occurrence* of c e n t e n -
a r i a n s which h a s e v e r boon m a d e in t h a t of T . R Y o u n g .
B e c a u s e of t h e c o n s i d e r a b l e intrinnic i n t e r e n t of (lie
m a t t e r , a n d t h e p o p u l a r m i s c o n c e p t i o n s which g e n e r a l l y
p r e v a i l a b o u t it, it will !>o w o r t h w h i t e to t a k e n little
time t o e x a m i n e YoungVi metlKKlK a n d r e s u l t s . H e p o i n t s
o u t in t h e b e g i n n i n g t h a t <h<* e v i d e n c e of g r e a t a g e which
is u s u a l l y a c c e p t e d by c e n s u s officials, by r e g i H t r a r s of
d e a t h , by n e w s p a p e r reporter**, a n d by the* g e n e r a l p u b l i c ,
is, g e n e r a l l y s p e a k i n g , of no v a l i d i t y o r trustworthincHM
w h a t e v e r . S t a t e m e n t s of the* p e r s o n coneeriKnl, o r of t h a t
p e r s o n ' s r e l a t i v e s o r f r i e n d s , an t o extreme* lottgevity f
can a l m o s t i n v a r i a b l y be nhown by e v e n a little investign*
tion t o be e x t r e m e l y unreliable*. T o be a c c e p t a b l e a s
scientific e v i d e n c e a n y s t a t e m e n t of g r e a t a g e m u s t be
s u p p o r t e d b y u n i m p e a c h a b l e documintary p r o o f of a t
l e a s t t h e following p o i n t s :
a. Th data of birth, ur f huptkm.
b. This datf* of death,
e. Th# identity of the ponton <lyitt|( Jtfc a iutffHHtil v#*ry 4ffi"wl agi*
with the* permm fur whom th Wrth tif Imjitisntitftl ri^f*ni
which thff rhutw at pjmit itgtt In \mbt*il, wan wm\*< unt
d. In th* imne |iartiruirJy t>f tmrrknl wtiiftifit th
t\w pprmm U> whom m&.rrM> tmtl ity tilh^r d*f* whh'H wlli
hi*lf to cMtAhliffh firimf *tt

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 identitytlmt
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)

BrMUftt nt&ttii U*t J.vujtff

(ingJr* or
jartarriml) Ytitm Mtmtto

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

living 30 September, 1005.

t Living 31 July, imn,

which one can find in a n y yniir f mortality *teUntiea, i t

striking. I n a n examination of thtt matter r t w m t l y , f o r
example, it w a s found that in tho registration nnm of tho
26 BIOLOGY OF D E A T H
United States there were recorder! in the yoar 1916, out
of a total of 1,001,921 deaths at all a g w tho following as
of ages 100 or over:
White males *T
Colored males IW
White females ,.,.,. t0
Colored female , . . . , . . . SW

Total . . . , . . , . . . . . > . lft

In this large total 4 pernon* wc*ri! rcjconlocl itt* having
died at the age of 120, and one, a c o l o m l fsmalts itt the*
preposterous age of 134!
B. There is no generally valid, orderly relationship
between the average duration of life of thr individuals
composing a species and any other broad fart now known
in their life history, or their structure, of thrJr physiology*
Many attempts have been made to not tip K<*n<rali2atioiw
establishing c o n n e c t i o n of thia n o r t W d u m a m t particu-
larly, has endeavored to ent&blt&h uch rclationn only to
have them overthrown, somotimcii by fuotn wliich hn him-
self presents. It has, for example, bet*tt mmUmdml that tht
larger ML animal the longer ita Ufa. Thin in obviounly
no general law* Again it hai* been held that n o animal
lives after reproducing, except itiah an care for their
young, but almost numberle** initancen mn be adduced
where no such relationship hoick. I t will not pay to ex-
amine ail the hypotheses of this general type whiph have*
at one time or another, been put forward- W i t h ona exetp
tion, to which we shall advert inunediately f they all suffer
from too many important exceptions to b considered
valid geueralizationfi.
0* Natural death as distinguished from aeddemtd
death is preceded by definite iimdwrd and fumtwnal
 THE PROBLEM 27
in the body. T h e s e changes in the s t r u c t u r e of
o r g a n s a n d p a r t s of the body, a n d i n t h e i r m a n -
functioning constitute t h e m a t e r i a l basis of w h a t is
senescence or g r o w i n g old. S o m e of the m o r p h o -
l o g i c a l a n d physiological changes which c h a r a c t e r i z e ex-
t senescence a r e a p p a r e n t and k n o w n t o all. S u c h
i n case of m a n the b e n t p o s t u r e which m e a n s a n a l t e r e d
p o s i t i o n and fusion of the elements of the v e r t e b r a l
c o l u m n , the w r i n k l e d visage, which denotes a p r o f o u n d al-
" t o r - a t i o n of t i s s u e elements, and the shuffling a n d uncer-
t a i n gait, which bespeaks a failing m o t o r coordination.
I n I F i g u r e 1 t h e s e senescent changes a r e all well indicated
i * x t h e case of a n old m a n who h a s received m u c h news-
J > a / p e r notice, " U n c l e " J o h n Shell of K e n t u c k y , who is
l x e x ~ e shown w i t h h i s l a s t wife and s u p p o s e d son. T h i s
p o o r old m a n h a s been exhibited about t h a t p a r t of t h e
a s " t h e oldest living h u m a n b e i n g , " a t a claimed
of 131 y e a r s . A s a m a t t e r of fact, N a s c h e r , who h a s
a careful i n v e s t i g a t i o n of t h e case, finds h i m to be
c
** a b o u t one h u n d r e d y e a r s old, possibly a y e a r y o u n g e r
o r o l d e r . " T h e p a t e r n i t y of t h e 4^/2 y e a r old boy, t h o u g h
o l a i x n e d b y Shell, is i n considerable doubt.
ZBesid-e t h e s e obvious senescent c h a n g e s t h e r e a r e
g ^ o l x x g on even m o r e significant changes in t h e cellular ele-
j a c x o x x t s which compose t h e body. C e r t a i n of t h e s e cellular
of a g e w e r e described i n a series of Lowell lec-
given a little m o r e t h a n a decade a g o b y t h e l a t e
J O o r . C h a r l e s Sedgwick Minot. Over a q u a r t e r of a cen-
a g o H o d g e m a d e a careful s t u d y of senile c h a n g e s in
cells. I n a m a n d y i n g n a t u r a l l y a t 92 y e a r s of a g e
f o x i n d m a r k e d c h a n g e s i n t h e cells of t h e s p i n a l gangKa
o o m p a r e d w i t h t h o s e of a n e w b o r n babe. T h e chief
iCJLidQ:orences a r e exhibited i n Table 3.
28 BIOLOGY O F D E A T H
TABLE 3
Shomng the Principal Differences Observed on Comparing thr Sfhtmt Cfanyltim
Cells (First Cervical Ganglion) from a Child at Birth With THQM?
from a Man Dying of Old Age at Ninety-two Year.
(From I lodge's data)
Baby at birth. Mate
Volume of nucleus 100 per cent. IH.'i j**r writ.
Nucleoli visible 53 per cent. ft j*?r vnt.
Deep pigmentation 0 per cent. 1)7 |*
Slight pigmentation 0 per cent. |r wnt.
Hodge found still m o r e m a r k e d c h a n g e s in t h e a n t e n -
n a r y lobe of the n e r v o u s s y s t e m of the honey bet*. T h e
n a t u r e of the changes is shown in F i g u r e 2.
I n the ganglion cells of both m a n a n d the honey bi*ct
the volume of the nucleus in p r o p o r t i o n to t h a t of the
r e s t of the cell body becomes reduced with a d v a n c i n g a g e .
Minot showed t h a t this w a s a v e r y g e n e r a l p h e n o m e n o n
in senescence, and was a eontitnioim proeeiu* front b i r t h to
death. H e gave to i t a n d r e l a t e d a n d associated c e l l u l a r
changes the n a m e " e y t o m o r p h o s i H , " and a t t r i b u t e d to it
t h e g r e a t e s t significance in b r i n g i n g a b o u t utmcmcemu! a n d
death. A s we shall p r e s e n t l y see, cytomorphoHttt m a y
p e r h a p s m o r e j u s t l y be r e g a r d e d an one of t h e m o r p h o -
logical results of senescence r a t h e r t h a n i t s o u t s e .
Eecently M r s . Pixell-Goodrich, a n Knglish w o r k e r , lia
re-studied the senescent changes in t h e cells of th honey
bee. H e r w o r k shows i n a s t r i k i n g way t h e Ions of proto
p l a s m in t h e aged c e l l I n t h e y o u n g bm i m m e d i a t e l y
after hatching, t h e cells a r e large a n d p l u m p t only s e p a r -
ated from each other by n a r r o w s t r a n d s of c o n n e c t i v e
tissue. I n t h e same region of the s a m e ganglion in a n old
bee which came f r o m a hive on a fins d a y in M a r c h , b u t
w a s too weak to effect a cleansing flight a n d soon b u e a m *
moribund, t h e n e r v e cells w e r e q u i t e w o r n o u t There
 THE PROBLEM 29
w a s left only a f r a m e w o r k of connecting tissue, w i t h a n
occasional nucleus of a n e r v e cell in a m o r e o r less
n e c r o t i c condition, w i t h only a little c y t o p l a s m a r o u n d it.

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.

D. Natural death (as distinguished from accident^0

occurs normally and necessarily only in animals coW1-
posed of many cells. Unicellular o r g a n i s m s are finally
known, to a considerable extent as the r e s u l t of t h e l
liant and p a i n s t a k i n g r e s e a r c h e s of Woodruff and
students, to be i m m o r t a l in esse a s well a s in posse.
the discovery by Woodruff and E r d m a n of the p r o c e s n
of nuclear reorganization, which t h e y call endomixis, thi-B
conclusion is as solidly g r o u n d e d if we r e g a r d a cycle o f
protozoan divisions a s t h e homologue of t h e m e t a z o a x i
body, as i t is if we consider each i n d i v i d u a l p r o t o z o a n a n
such homologue. Woodruff h a s been cultivating t h e c o m -
mon unicellular f o r m Paramecium, shown in F i g u r e 3 ,
for over 13 y e a r s .
D u r i n g all this time no conjugation or p a i r i n g of i n -
dividuals h a s occurred. I n a recent l e t t e r Dr. W o o d r u f f
s a y s : "After we h a d discovered a n d w o r k e d out e n d o -
mixis there seemed n o p a r t i c u l a r u s e of carefully r e c o r d -
ing the n u m b e r of g e n e r a t i o n s each day. B u t the c u l t u r e
is still going on as well as ever a n d i s a t a p p r o x i m a t e l y
the 8500th generation13y 2 y e a r s old! On M a y 1 ,
1915, (just 8 y e a r s old) it w a s a t t h e 5071st g e n e r a t i o n . 9 *
If in 8500 generationsa d u r a t i o n of h e a l t h y r e p r o d u c -
tive existence which, if the g e n e r a t i o n w e r e of the s a m e
length as in m a n would r e p r e s e n t r o u g h l y a q u a r t e r of a,
million y e a r s i n absolute t i m e n a t u r a l death, h a s n o t
 T H E PROBLEM 31
occurred, we m a y with reasonable UHHUranee cumeindu
that this animal is immortal.
Of even more probative value, in the opinion of
some workers, than the reaultn on Paramvcium are

the recent experimental of H a r t m a n n , who mil twitted

Eudorina elegant* for over (MM) %mprtitimw without con-
jugation or any ntiekmr rt*wifnnhntUm mmmpmuttng to
endomixis, and no ctoprwwion in tin* miltttr^ ormirrec).
T h e distinction between P r o t o z o a unil Motuxoa in
32 BIOLOGY OF D E A T H
respect of the incidence of natural death is HO important
that it requires a somewhat detailed explanation, together
with the reasons for it. Protozoa reproduce by a process
of simple division or fission. A particular individual
after growing to a certain size simply divides t ransvtsrady
into two like individuals, at first smaller in siao, hut ra-

FIG. 4Diagram showing the pree# ctf r@produ@tita8 by v* ft

fission in the unicellular orginism Pararmrwim. iV
pidly growing to full adult magnitude. The
gross features of this process are illustrated in F i g u r e 4.
One cannot say, after the act of fission i s accomplished,
which i s parent and which is offspring* One individual
simply becomes two and, in the process of becoming two,
loses totally its own identity as an individual. Upon occa-
sion another process known as conjugation may intervene,
In this process two individuals mate together. B y a
process of assortative mating, Hke sizes pair together,
 T H E PROBLEM 33
a s w a s first shown by the writer and later confivttuul by
Jennings. A f t e r p a i r i n g has occurred an intftrchaugo of
nuclear substance occurB by a mechanism described and
figured in m a n y elementary textbooks of zoology. Thin
process of conjugation new! not further concern tin hi^ro,
f o r the reason that Woodruff, in the work already rferred
to, has s h o w n that thin phenomenon in not onstmtial to
the continued life of the race, Its place* may bt*f and
normally very frequently is, takttn by the prwvnn cultml
endomixis. I n thin process then* occurs a nuclear break-
down and reorganization which appunrs to bo tlit* <i|uivn*
lent, functionally a t least, of that which takt** plain*
during conjugation.
There has been much discussion, particularly a m o n g
European workers, an for example* Dojtain, f tillon, Wi*dt*
kind, Slotopowaki, and o t h e r s about curtain phiUmophi-
eal, not to s a y zneUiphyHical, ap<*ctH of immortality in
the Protozoa. B u t all mich diHcuion hiw in no WIHII din
turbed o r altered the plain phynical fact that thi*r* in no
place for death in a Hcheme of reproduction by Mtrnplo
fission, such m is illufitratoil in F i g u r e 4. N o t h i n g in left
at any s t a g e to fulfill thci proverbial mhimm of M dtmt to
dust and ashes to aghtm," W h e n an individual i i t h r o u g h
its single individual existence it simply bcteom^H t w o imli*
viduals, which g o on p l a y i n g the fnnmnattng g a m o of
living here and now.
I n a few of tha s i m p b u t and m o s t lowly organizmi
groups of many-eelled animals or Motazoa thin powt*r of
mnltiplicatiou by simpki fission, o r buckling off a portion
of the body which reproduces the whole, i i riitiiintd an
a facultative a s s e t T h i s p r o c e s s of reproduction in which
the somatic or body ei*lls of one gunoration prottacn tin*
somatic cells of the n e x t generation hm Immi ciallitd
agamic reproduction. I t occurs ma the m o m uml but n o t
s
34 BIOLOGY OF DEATH

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
nirjcf 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.

Fig. HProgr<MHiofngftmirn^ timmtiwnmkith*-tmm&Umt?N#w

 THE PROBLEM 37
this m a n n e r i n t h e s e m u l t i c e l l u l a r f o r m s t h e r e is no place
f o r death. I n t h e p a s s a g e f r o m one g e n e r a t i o n to t h e
n e x t n o r e s i d u e is left behind. A g a m i c r e p r o d u c t i o n a n d
its associated a b s e n c e of d e a t h o c c u r s v e r y c o m m o n l y in
plants. Budding and p r o p a g a t i o n by cuttings are the
common f o r m s i n w h i c h i t is seen. T h e s o m a t i c cells h a v e
t h e c a p a c i t y of c o n t i n u i n g m u l t i p l i c a t i o n a n d life f o r a n
indefinite d u r a t i o n of t i m e , so l o n g a s t h e y a r e n o t acci-
dentally c a u g h t i n t h e b r e a k d o w n a n d d e a t h of t h e whole
individual in which they a r e a t the m o m e n t located. Thus
v i r t u a l l y e v e r y a p p l e t r e e i n e v e r y o r c h a r d i n t h i s coun-
t r y is simply a d e v e l o p e d b r a n c h o r b u d of s o m e original
a p p l e t r e e f r o m w h i c h i t w a s cut, i n m a n y c a s e s c e n t u r i e s
ago. A p p l e t r e e s c a n n o t of t h e i r o w n u n a i d e d efforts
p r o p a g a t e e i t h e r b u d s o r c u t t i n g s . So, u n t i l t h e i n t e r v e n -
tion of m a n , some a p p l e t r e e s d i e d n a t u r a l d e a t h s , s o m a t i -
cally s p e a k i n g , j u s t a s do t h e h i g h e r a n i m a l s of which
we shall s p e a k p r e s e n t l y . B u t t h e i r cells w e r e i n h e r e n t l y
capable of b e t t e r t h i n g s , a s w a s d e m o n s t r a t e d w h e n m a n
first cut off a s h o o t f r o m a n old a p p l e t r e e a n d p r o v i d e d
i t w i t h a r o o t b y g r a f t i n g . * T h e n i t w e n t on a n d m a d e a
n e w t r e e . F r o m i t i n t u r n c u t t i n g s w e r e t a k e n , a n d so t h e
p r o c e s s h a s c o n t i n u e d t o t h e p r e s e n t d a y . A p a r t of t h e
s o m a of one g e n e r a t i o n p r o d u c e s t h e s o m a of t h e n e x t
g e n e r a t i o n a n d goes on l i v i n g indefinitely.
A different m o d e of r e p r o d u c t i o n i s c h a r a c t e r i s t i c
of h i g h e r m u l t i c e l l u l a r a n i m a l s , a n d i n all b u t t h e lowest
g r o u p s is t h e exclusive m e t h o d . A n e w i n d i v i d u a l is
s t a r t e d b y the u n i o n of two p e c u l i a r cells of e x t r a o r d i n a r y
potentialities, called g e r m cells. T h e s e g e r m cells a r e of
two sorts, ova a n d spermatozoa. I n bisexual organisms
* This provision of roots was not essential, only practically convenient.
The cutting would, if enough pains were taken, grow its own roots.
38 BIOLOGY OF DEATH
the former are borne in the female, and the latter in the
male body. Both sorts undergo a complicated prepara-
tion for union, the result of which is that when union does
occur each party to it contributes either an exactly equal
or an approximately equal amount of hereditary mate-

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,

ision in eiegg of Cyclops, showing at one pole of spindle the granules

. FIG. 10.First division
which mark the germ path. (From Child, after Amma, by permission of University of
Chicago Press).
i t w a s shown b y W o o d s , m a n y y e a r s ago, t h a t t h e g e r m
cells a r e definitely localized a n d recognizable, a s shown
in F i g u r e 9.
I n some f o r m s , n o t a b l y t h e r o u n d - w o r m Ascaris, va-
rious Crustacea a n d insects, t h e cells w h i c h a r e t o become
g e r m cells a r e visibly set a p a r t f r o m t h e v e r y first o r one
of t h e first t h r e e or f o u r c l e a v a g e s of t h e fertilized ovum.
F o r example, i n t h e case of t h e c r u s t a c e a n Cyclops, A m m a
h a s shown t h a t t h e g r a n u l e s visible a t one pole i n the
v e r y first division m a r k t h e p r o s p e c t i v e g e r m p a t h , as
shown i n F i g u r e 10.
I n t h e g n a t Chironomus t h e s a m e t h i n g i s visible a t a
v e r y e a r l y c l e a v a g e , a c c o r d i n g t o t h e o b s e r v a t i o n s of
H a r p e r . F o r a c o m p r e h e n s i v e a n d critical r e v i e w of t h e
40 BIOLOGY O F D E A T H
e x t e n s i v e l i t e r a t u r e on t h e Ktimhahn one nhould v\
<m
the r e c e n t c o n t r i b u t i o n s of H e g n e r th> subject.
T o comlenBe a long a n d complicated m a t t e r we
s t a t e t h e s i t u a t i o n r e g a r d i n g r e p r o d u c t i o n a m i !*
t h e Metazoa in thin w a y . A higher, multicllular i
d u a l m a y f>e conceived, from t h e v i e w p o i n t of iht* pi
discussion, as composed of t w o eBmmtmlly t m i r p f
p o r t i o n s : t h e g e r m cells on the* one h a n d , whirl* m
m o r t a l in t h e s a m e sense* t h a t (ho P r o t o z o a tm* iinin
a n d the r e s t of the body, which it \H convpniftit ti
technically the Homa, on the* o t h e r h a n d . T h e norna ti
goes n a t u r a l d e a t h a f t e r an i n t e r v a l of t i m e which, \
h a v e seen, v a r i e s from species to 8pf*ri<8. T h e ^rtt)
which t h e individual h o a r s in its body nt t h e timr* i
d e a t h of c o u r s e die a!m>. B u t thin m p u r e l y nerid
d e a t h so f a r a s c o n c e r n s t h e g e r m celln. Htirfi of i\u
were, p r i o r t o t h e d e a t h of t h e Homii, nniiblei! t o
w i t h o t h e r g e r m cells w e n t on living jimt it* thm
dividing Paramecium. Reduced t o n f o r m u l a WP ma;
t h a t the fertilized ovum (united g e r m W I I N ) prodrii
soma f and m o r e gcrm w l l . T h e nomii eventiiiilly
Some of the g e r m cells, p r i o r to t h a t event f pn
s o m a t a a n d g e r m cells, a n d HO on in a ctmVmnmw
which h a s n e v e r y e t <?nrlcd Btrice t h e apptturanci* of n
cellular o r g a n i s m s on t h e e a r t h .
T h e c o n t r a s t between the* p r o t o z o a n a n d t h e inetii
m e t h o d of descent is s h o w n in F i g u r e 1 1 , which in n t
fication of a s i m i l a r d i a g r a m o r i g i n a l l y d u e t o my
league, D r . HL 8 . J e n n i n g s ,
T h e d i a g r a m r e p r e s e n t t h e d e s c e n t of g e n e r a l
T h e t i p p e r p o r t i o n of t h e d i a g r a m s h o w s t h e mod
d e s c e n t in f o r m s r e p r o d u c i n g from o r g a n i s m s ruprc
i n g f r o m a single p a r e n t . T h e lower, o r B p o r t i o n &
 THE PROBLEM 41

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 dtath, which the
writ*r though iliflforitiK (mm wmw of the e<mduionii, h& found useful
in tht 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 digroc*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
ourHclvHr 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
farli<tfi 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 tiMtu*x cutmtnw or <Iwtr<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* rcmtttute & 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

CONDITIONS OF CELLULAE IMMORTALITY

I N the preceding chapter it was pointed out t h a t the

g e r m cells of h i g h e r o r g a n i s m s a r e p o t e n t i a l l y , a n d u n d e r
certain conditions i n fact, i m m o r t a l . W h a t a r e t h e con-
ditions of i m m o r t a l i t y i n this c a s e ? A r e t h e y s u c h a s
to s u p p o r t t h e t h e s i s t h a t t h e p r o c e s s e s of m o r t a l i t y a r e
essentially physico-chemical i n n a t u r e , a n d follow
physico-chemical l a w s ?

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 mg-
im?ii tut i cm
n/2(K)0 KCN F<w Kggj* go through th< <*nrly HtugttH of

n/1000 KCN Many eggn wgiucnt nnci dcvi^loji in to Hwitn-

ming Inrvfi*
n/750 KCN Many ogg mfiptwnl firiti cii*vi4inp into xwtrn-
ming inrvif!
n/400 KCN A fuw eggs develop into ftwiiimiing larva*
n/300 KCN No (gg nfgrncmtn
n/250 KCN No egg D($gm(*titN
n/200 KCN No <?gg Ms%nu*ntn
n/100 KCN No i w wgimnu

F r o m thin t a b l e i t in seen t h a t in concctntratioiiH of

K C N f r o m n / 7 5 0 t o n / 1 0 0 0 t h e eggn d e v e l o p e d p e r f e c t l y
i n t o s w i m m i n g larva*. I n o t h e r w o r d a , by t h e a d d i t i o n
of t h i s v e r y small a m o u n t of K C N , t h e life p e r i o d h a s
been p r o l o n g e d to t h r e e t i m e s w h a t i t w o u l d n o r m a l l y
be u n d e r t h e s a m e e n v i r o n m e n t a l condition**. Concen-
t r a t i o n s of K C N w e a k e r t h a n n / 1 0 0 0 w e r e i n c a p a b l e of
p r o d u c i n g t h i s r e s u l t , o r a t best* if d e v e l o p m e n t s t a r t e d ,
t h e p r o c e s s c a m e v e r y quickly t o a n end. I n a l r o n g e r
concentrations than n/400 the eggs were evidently poi-
soned, a n d n o d e v e l o p m e n t o c c u r r e d .
O t h e r e x p e r i m e n t s of L o e b ' s ahow t h a t the* l e t h a l
effects of v a r i o u s toxic a g e n t s u p o n t h e e g g ec*ll m a y be
inhibited or p o s t p o n e d for a r e l a t i v e l y l o n g time, b y
 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 55
suitable chemical t r e a t m e n t , s u c h a s lack of oxygen, K C N ,
or chloral h y d r a t e . A t y p i c a l e x p e r i m e n t of t h i s kind
m a d e u p o n t h e sea-urchin, Strongylocentrotus purpuratus
m a y be q u o t e d :
Eggs were fertilized with sperm and put eleven minutes later into
three flasks, each of which contained 100 c. o. of sea-water + 16 c. c. 2-%
m CaCl2. One flask was in contact with air, while the other two flasks
were connected with a hydrogen generator. The air was driven out from
these two flasks before the beginning of the experiment. The eggs were
transferred from one of these flasks after four hours and fourteen minutes,
from the second flask after five hours and twenty-nine minutes, into normal
(aerated) sea-water. The eggs that had been in the hypertonic sea-water
exposed to air were transferred simultaneously with the others into
separate dishes with aerated normal sea-water. The result was most
striking. Those eggs that had been in the hypertonic sea-water with air
were all completely disintegrated by "black cytolysis." Ten per cent,
of the eggs had been transformed into <fshadows" (white cytolysis). It
goes without saying that all the eggs that had been in the aerated hyper-
tonic sea-water five and a half hours were also dead. The eggs that had
been in the same solution in the absence of oxygen appeared all normal
when they were taken out of the solution, and three hours laterthe
temperature was only 15C.they were all, without exception in a per-
fectly normal two- or four-cell stage. The further development was also
in most cases normal. They swam as larv at the surface of the vessel
and went on the third day (at the right time) into a perfectly normal
pluteus stage, after which their observation was discontinued. Of the
eggs that had been five and a half hours in the hypertonic sea-water
deprived of oxygen, about 90 per cent, segmented.
L e t u s c o n s i d e r one m o r e i l l u s t r a t i o n f r o m L o e b ' s
work i n t h i s field. N o r m a l l y , i n t h e f o r m s w i t h which
he chiefly w o r k e d , sea-urchin, starfish, a n d c e r t a i n mol-
luscs, an absolutely e s s e n t i a l condition f o r t h e c o n t i n u a -
tion of life of t h e germ-cells a f t e r t h e y a r e d i s c h a r g e d
f r o m t h e b o d y is t h a t t w o cells, t h e o v u m a n d t h e s p e r -
matozoon, s h a l l u n i t e i n n o r m a l f e r t i l i z a t i o n . P u t in
a n o t h e r w a y , p a r t h e n o g e n e s i s does n o t n o r m a l l y occur i n
these forms- F e r t i l i z a t i o n is a n e s s e n t i a l c o n d i t i o n f o r
the c o n t i n u a t i o n of life a n d d e v e l o p m e n t . B u t L o e b ' s
56 BIOLOGY OF D E A T H
p a i n s t a k i n g a n d b r i l l i a n t r e s e a r c h e s , e x t e n d i n g over a
n u m b e r of y e a r s , show t h a t w h e n we s a y t h a t fertilization
is a n essential condition for the continued life of the
germ-cells o u t s i d e t h e body, o u r l a n g u a g e t e n d s to ob-
scure the m o s t i m p o r t a n t fact, w h i c h i s simply t h a t for
the continuation of life i n t h e s e cells only c e r t a i n i n t e r n a l
physico-chemical conditions a n d adjustments* m u s t be
realized. I t m a t e s n o essential difference to t h e r e s u l t
w h e t h e r t h e s e conditions a r e r e a l i z e d t h r o u g h the
i n t e r v e n t i o n of t h e s p e r m , a s i n n o r m a l fertiliza-
tion, o r by p u r e l y artificial chemical m e t h o d s initiated,
controlled a n d d i r e c t e d a t e v e r y s t e p b y h u m a n agency.
"We can, i n o t h e r w o r d s , r e g a r d all c a s e s of suc-
cessful artificial p a r t h e n o g e n e s i s a s f u n d a m e n t a l l y a con-
t r i b u t i o n to t h e p h y s i o l o g y of n a t u r a l d e a t h , a n d a demon-
s t r a t i o n of i t s e s s e n t i a l l y m e c h a n i s t i c b a s i s . T h e condi-
tions of continued existence a r e p h y s i c a l a n d chemical,
and controllable a s such. T h e m e t h o d s finally w o r k e d out
as o p t i m u m afford a complete d e m o n s t r a t i o n of t h e t h e s i s
we h a v e j u s t s t a t e d . T h u s , f o r e x a m p l e , t h e unfertilized
egg of t h e sea-urchin, Strongylocentrotus purpuratus,
will continue i n life a n d d e v e l o p p e r f e c t l y n o r m a l l y if i t
is subjected t o t h e following t r e a t m e n t : T h e eggs a r e
first placed i n s e a - w a t e r t o w h i c h a definite a m o u n t of
weak solution of b u t y r i c acid h a s b e e n a d d e d (50 cc. of
sea-water + 2.8 c c . n / 1 0 b u t y r i c a c i d ) . I n t h i s solution
a t 15 C. t h e e g g s a r e allowed t o r e m a i n f r o m 1 % t o 3 or
4 m i n u t e s . T h e y a r e t h e n t r a n s f e r r e d t o n o r m a l sea-water,
in which t h e y r e m a i n f r o m 15 t o 20 m i n u t e s . T h e y a r e
then t r a n s f e r r e d f o r 30 to 60 m i n u t e s a t 15 C. t o s e a - w a t e r
which h a s h a d i t s osmotic p r e s s u r e r a i s e d b y t h e a d d i t i o n
of some s a l t s (50 c c . of s e a - w a t e r + 8 cc. of 2 % m NaCl, or
2y2 m N a C l + K C L + C a C l 2 in t h e p r o p o r t i o n i n which t h e s e
 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 57
s a l t s exist i n s e a - w a t e r ) . A f t e r t h e s t a y of f r o m 30 t o 60
m i n u t e s i n t h i s solution, t h e eggs a r e t r a n s f e r r e d b a c k t o
normal sea-water, the t r a n s f e r r i n g being in batches a t
i n t e r v a l s of 3 t o 5 m i n u t e s b e t w e e n each b a t c h t r a n s f e r r e d .
I t is t h e n f o u n d t h a t t h o s e eggs which h a v e b e e n j u s t t h e
r i g h t l e n g t h of t i m e i n t h e h y p e r t o n i c s e a - w a t e r d e v e l o p
i n t o p e r f e c t l y n o r m a l s e a - u r c h i n larvse. I n o t h e r w o r d s ,
w e h a v e h e r e a definite a n d k n o w n physico-chemical p r o -
cess completely r e p l a c i n g w h a t w a s , b e f o r e t h i s w o r k ,
u n i v e r s a l l y r e g a r d e d a s a p e c u l i a r l y v i t a l p r o c e s s of
e x t r a o r d i n a r y complexity,. Iprobably b e y o n d p o w e r of
h u m a n control.
T h e s e t h r e e e x a m p l e s f r o m L o e b ' s w o r k on t h e s u b -
j e c t of p r o l o n g a t i o n of life i n t h e e g g cell will suffice f o r
o u r p r e s e n t p u r p o s e s . T h e lesson w h i c h t h e y t e a c h i s
plain, a n d is one w h i c h h a s , a s will be r e a d i l y p e r c e i v e d ,
a m o s t i m p o r t a n t b e a r i n g u p o n t h e g e n e r a l c o n c e p t of
life a n d d e a t h o u t l i n e d i n t h e p r e c e d i n g c h a p t e r . The
experiments demonstrate t h a t the conditions essential to
c o n t i n u e d life of t h e germ-cells outside t h e b o d y a r e p h y -
sico-chemical c o n d i t i o n s , a n d t h a t w h e n t h e s e cells d i e i t
i s b e c a u s e t h e n o r m a l physico-chemical m a c h i n e r y f o r t h e
c o n t i n u a t i o n of life h a s e i t h e r b r o k e n down, o r h a s n o t
been g i v e n t h e p r o p e r a c t i v a t i n g chemical c o n d i t i o n s .
L a c k of s p a c e alone p r e v e n t s g o i n g i n d e t a i l i n t o a n -
other extremely interesting and important development
of t h i s subject, d u e t o D r . P r a n k R. Lillie of t h e U n i v e r -
s i t y of Chicago. H e h a s , i n r e c e n t y e a r s m a d e a t h o r o u g h
a n a l y s i s of t h e biological f a c t o r s o p e r a t i n g w h e n t h e egg
of t h e s e a - u r e h i n is n o r m a l l y fertilized b y a s p e r m a t o -
zoon. T h e c o n c e p t i o n of t h e p r o c e s s of f e r t i l i z a t i o n t o
which Lillie c o m e s i s " t h a t a s u b s t a n c e b o r n e b y t h e e g g
(fertilizin) e x e r t s t w o k i n d s of a c t i o n s : (1) a n a g g l u t i n -
58 BIOLOGY OF D E A T H
a t i n g action on t h e s p e r m a t o z o o n a n d (2) a n activating
action on t h e egg. I n o t h e r w o r d s , t h e s p e r m a t o z o o n is
conceived, by m e a n s of a s u b s t a n c e which i t b e a r s and
which e n t e r s i n t o u n i o n w i t h the f ertilizin of the eggy to
release t h e a c t i v i t y of this s u b s t a n c e w i t h i n t h e egg.jy
F r o m the s t a n d p o i n t of t h e p r e s e n t discussion i t is ob-
vious t h a t L i l l i e ' s r e s u l t s so f a r p r e s e n t n o t h i n g which
in a n y w a y d i s t u r b s t h e conclusion w e h a v e r e a c h e d as
t o the essentially physico-chemical n a t u r e of t h e processes
which condition t h e c o n t i n u a t i o n of life and development
of the egg.

TISSUE CULTXJEE IN VITRO

L e t u s t u r n n o w t o a n o t h e r question. A r e t h e germ-
cells t h e only cells of t h e m e t a z o a n b o d y which possess
the c h a r a c t e r i s t i c of p o t e n t i a l i m m o r t a l i t y ? T h e r e is
now a n a b u n d a n c e of evidence t h a t such is n o t the case,
b u t t h a t , on t h e c o n t r a r y , t h e r e a r e a n u m b e r of cells a n d
tissues of t h e body, which, u n d e r a p p r o p r i a t e conditions,
m a y continue living indefinitely, except for t h e p u r e l y
accidental i n t e r v e n t i o n of l e t h a l circumstances. E v e r y
child knows t h a t all t h e t i s s u e s d o n o t d i e a t t h e s a m e time.
I t is p r o v e r b i a l t h a t t h e tail of t h e s n a k e , whose h e a d a n d
body h a v e been b a t t e r e d a n d c r u s h e d u n t i l even the small
boy is willing t o a d m i t t h a t t h e j o b of killing is complete,
16
will n o t die u n t i l t h e s u n goes d o w n . ' ' G a l v a n i 'si f a m o u s
e x p e r i m e n t w i t h t h e f r o g ' s l e g s only succeeded because
some p a r t s s u r v i v e a f t e r t h e d e a t h of t h e o r g a n i s m a s
a whole. A s H a r r i s o n p o i n t s o u t " A l m o s t t h e whole of
our knowledge of muscle-nerve physiology, a n d m u c h of
t h a t of t h e action of t h e h e a r t , is b a s e d u p o n e x p e r i m e n t s
with s u r v i v i n g o r g a n s ; a n d i n s u r g e r y , w h e r e we h a v e t o
do w i t h c h a n g e s involved i n t h e r e p a i r of i n j u r e d p a r t s ,
 Vm l.'f. V'vw of tHHtU' from frog cinlMyo <MiItivutc<I in lymph, two dnyH oI<J.
The (i.'trk j>jtior .nlnmw oriKinitl hit of tmum*. Li^Iitcr portioti.s uic new growth.
(I'Vorn HurriNon. i
h'la If, *",? o l , , | ; : i
 C O N D I T I O N S O P C E L L U L A R I M M O R T A L I T Y 59
i n c l u d i n g p r o c e s s e s of g r o w t h a n d differentiation, t h e
p o w e r of s u r v i v a l of t i s s u e s a n d o r g a n s a n d t h e i r t r a n s -
p l a n t a b i l i t y t o s t r a n g e r e g i o n s , even to o t h e r i n d i v i d u a l s ,
h a s l o n g f o r m e d t h e b a s i s of p r a c t i c a l p r o c e d u r e s . "
T h e first successful c u l t u r e s of s o m a t i c cells a n d t i s -
s u e s o u t s i d e t h e b o d y w e r e t h o s e of L e o L o e b , d e s c r i b e d
i n 1897. H i s first m e t h o d consisted i n c u l t i v a t i n g t h e
tissues in a p p r o p r i a t e media in test tubes. L a t e r he used
also a n o t h e r m e t h o d , which involved t h e t r a n s p l a n t a t i o n
of t h e solid m e d i u m a n d t h e t i s s u e i n t o t h e b o d y of a n -
o t h e r a n i m a l . W h a t h a s b e e n r e g a r d e d a s a d e f e c t of
b o t h t h e s e m e t h o d s is t h a t t h e y do n o t p e r m i t t h e contin-
u e d o b s e r v a t i o n of t h e cells of t h e g r o w i n g c u l t u r e d t i s s u e .
T o H a r r i s o n is d u e t h e d e v e l o p m e n t of a m e t h o d w h i c h
does p e r m i t s u c h s t u d y . I n 1907 h e a n n o u n c e d t h e d i s -
c o v e r y t h a t if pieces of t h e d e v e l o p i n g n e r v o u s s y s t e m of
a f r o g e m b r y o w e r e r e m o v e d f r o m t h e b o d y w i t h fine
needles, u n d e r s t r i c t l y a s e p t i c p r e c a u t i o n s , p l a c e d on a
s t e r i l e cover slip in a d r o p of f r o g l y m p h , a n d t h e c o v e r
slip t h e n i n v e r t e d o v e r a hollow g l a s s slide, t h a t t h e t i s -
s u e s w o u l d r e m a i n alive for m a n y d a y s , g r o w a n d e x h i b i t
remarkable transformations. B y this technique it was
possible to study the changes with a high power micro-
scope a n d p h o t o g r a p h t h e m .
F i g u r e 13 is a g e n e r a l view of one of t h e s e t i s s u e cul-
t u r e s t w o d a y s old. I t s h o w s a piece of n e r v o u s t i s s u e
f r o m t h e f r o g e m b r y o , w i t h cells g r o w i n g o u t f r o m i t
i n t o t h e l y m p h . T h e l i g h t e r p o r t i o n s a r e t h e n e w cells.
I n his r e m a r k a b l e m o n o g r a p h H a r r i s o n s h o w s n e r v e
cells d e v e l o p i n g fibers a t first thickened, b u t p r e s e n t l y
b e c o m i n g of n o r m a l c h a r a c t e r a n d size. A t t h e fcnds a r e
p s e u d o p o d i a l p r o c e s s e s , b y which t h e g r o w i n g fiber a t -
t a c h e s itself t o t h e c o v e r slip o r o t h e r solid bodies* F i g -
60 BIOLOGY OF D E A T H
u r e 14 shows a p a r t i c u l a r l y beautiful n e r v e iibvr p r e p a r
a t i o n m a d e by B u r r o w s .
T h e fibers g r e w from a p r e p a r a t i o n of t h e embryonic
n e r v o u s s y s t e m of t h e chick. T h e r e c a n lx n o d o u b t , m
t h e s e figures HO clearly show, of the* life of theme celh
outside t h e body, o r of t h e n o r m a l i t y of t h e i r develop
mental and growth processes.
U n d e r t h e g u i d a n c e of H a r m o n , a n o t h e r worker
B u r r o w s , i m p r o v e d t h e technique of t h e c u l t i v a t i o n oi
t i s s u e s outside the body, first by u s i n g p l a s m a f r o m th<
blood i n s t e a d of l y m p h a n d l a t e r in v a r i o u s o t h e r way$
H e devised a n a p p a r a t u s f o r a f f o r d i n g t h e tissue* eultur*
a continuous s u p p l y of fresh n u t r i e n t m e d i u m . T h e r e it
in HUB a p p a r a t u s a l a r g e c u l t u r e c h a m b e r which t&km
t h e place of t h e p l a i n h a n g i n g d r o p in a n hermeticallj
sealed cell. O n t h e t o p of t h i s c u l t u r e churabur t h e r e U
a wick, which c a r r i e s t h e c u l t u r e fluid f r o m a supplying
c h a m b e r a n d d i s c h a r g e s i t i n t o a r e c e i v i n g c h a m b e r . TlM
t i s s u e is p l a n t e d a m o n g t h e fibers of t h e wick, which ar<
pulled a p a r t w h e r e i t c r o s s e s t h e t o p of this c h a m b e r
T h e whole s y s t e m in k e p t s t e r i l e a n d no a r r a n g e d thai
t h e g r o w i n g t i s s u e can be k e p t u n d e r o b s e r v a t i o n wlti
high p o w e r s of t h e m i c r o s c o p e . T h e n u t r i e n t medium
m a y bo modified a t will, a n d t h e effects of k n o w n s u b
s t a n c e s u p o n t h e cellular a c t i v i t i e s of e v e r y s o r t m&j
be s t u d i e d . j
B u r r o w s b e g a n h i s i n v e s t i g a t i o n s in t h i s field on thj
t i s s u e s of t h e e m b r y o chick. W i t h t h e ueee of ihes*
c u l t u r e s w a s established t h e f a c t t h a t t h e t i t i U i i of i
w a r m blooded a n i m a l w e r e a s c a p a b l e of life, d e v e l o p
m e n t , a n d g r o w t h outside t h e b o d y an w e r e t h o s e of coldj
blooded a n i m a l s , such a s t h e f r o g . B u r r o w s s u c c e e d ^
i n c u l t i v a t i n g o u t s i d e t h e b o d y , cells of t h e c e n t r a l uervoui
 %ll
lift \'f IlllJI*;itir|jj|rrt!Vi'f|:ni|i (f\\ U \l t\ ;Ull, r'f IL SI > * *< I Wl || (M li MHH Ml Jlil. TIlC IMl I lU'l1
IIJHII^ ls -M?i \<:%f tnv. Hi* ji;i jHjfj"a c,l i nlfujc LJs". it il'i i fVl It p.'t'.taK'1, witKhinp; (lie t'ciiiui
juttUun "t 1 J<i'ii|}iii<' with Itimtt't --utliit ii>i\ \\\\ \tmtt rvmitvittir. it \tinn tin- covct-i/lnm, ntt<
IiiM '! Ut h j!;t m :>n<i -tJ:irt. !'} |>f j;ti:if JM -.huw.t (In* i'Xfcut of irrcwlh ulifuinftl in IK II
frln |i7jjJn JJI! H-]b M-tiiuuinrs-' ;{' i*xhi pa I mrt ! f he ha^rinchl. i Afh-f Loiu-c tttul I'JIM'U
 C O N D I T I O N S OF CELLULAR I M M O R T A L I T Y 61
system, the heart, and mesenchymatous tissue of the
chick embryo. A t the same time Carrel w a s carrying
on studies in this same direction at the Rockefeller In-
stitute. I n his laboratory were made the first successful
cultures in vitro of the adult tissues of mammals. H e
developed a method of culture on a plate which permitted
the growing of l a r g e quantities of material. H e found
that almost all the adult and embryonic tissues of dog,
cat, chicken, rat, guinea pig, and man could be cultivated
in vitro. F i g u r e 15 shqws a culture of human tissue,
made at the Rockefeller Institute. I a m indebted to
Doctor Carrel and Doctor E b e l i n g f o r p e r m i s s i o n to pre-
sent this photograph here.
According to the nature of the t i s s u e s cultivated, con-
nective or epithelial cells were generated, which g r e w out
into the plasma medium in continuous layers or radiating
chains. Not only could normal tissues be cultivated but
also the cells of pathological growths (cancer c e l l s ) .
It has been repeatedly demonstrated that normal cell
division takes place i n these tissues cultivated outside the
body. The complex process of cell division, which is
technically called m i t o s i s , h a s been rightly regarded as
one of the m o s t characteristic, because complicated and
unique, phenomena of normal life processes. Y e t this
process occurs w i t h p e r f e c t normality i n cells cultivated
outside the body. T i s s u e s f r o m various organs of the
body have been successfully cultivated* including the
Mdney, the spleen, the thyroid gland, etc. B u r r o w s w a s
even able to demonstrate that the isolated heart muscle
cells of the chick embryo can divide a s well a s differen-
tiate, and "beat rhythmically in the culture medium.
P e r h a p s e v e n m o r e remarkable than the occurrence
of such physiological activity as that of the heart muscle
62 BIOLOGY O F D E A T H
cells in vitro is t h e fact t h a t in c e r t a i n l o w e r f o r m s of life
a s m a l l b i t of t i s s u e o r even a single cell, m a y d e v e l o p in
c u l t u r e i n t o a whole o r g a n i s m , d e m o n s t r a t i n g t h a t the
c a p a c i t y of m o r p h o g e n e s i s is r e t a i n e d in t h e s e isolated
s o m a t i c cells. H . V. W i l s o n h a s shown t h a t in coelenter-
a t e s a n d s p o n g e s complete new i n d i v i d u a l s m a y develop
in vitro f r o m isolated cells t a k e n f r o m a d u l t animals*
B y s q u e e z i n g small b i t s of t h e s e a n i m a l s t h r o u g h bolting
cloth he w a s able to s e p a r a t e small g r o u p s of cells o r
even single cells. I n c u l t u r e t h e s e would g r o w into s m a l l
m a s s e s of cells which would t h e n d i f f e r e n t i a l slowly into
t h e n o r m a l f o r m of t h e c o m p l e t e o r g a n i s m . F i g u r e 16
shows a n e x a m p l e of this t a k e n f r o m W i l s o n ' s work.
I t w a s e a r l y d e m o n s t r a t e d by C a r r e l a n d B u r r o w s
t h a t t h e life of t h e t i s s u e s in vitro, which v a r i e d in differ-
ent e x p e r i m e n t s f r o m 5 t o 20 day**, could be p r o l o n g e d by
a p r o c e s s of successive t r a n s f e r s of t h e culture* io a n
indefinite p e r i o d . Cells which w e n t H e a r i n g the* end of
t h e i r life a n d g r o w t h in one c u l t u r e need only be t r a n n -
f e r r e d t o a n e w c u l t u r e m e d i u m t o keep on g r o w i n g a n d
m u l t i p l y i n g , I)r- a n d M r s , W a r r e n I L L e w i s m a d e t h e
i m p o r t a n t d i s c o v e r y t h a t t i s s u e s of tho chick e m b r y o
could be c u l t i v a t e d o u t s i d e tho b o d y in p u r e l y i n o r g a n i c
solutions, such a s s o d i u m e h l o r i d e , B i n g e r ' a s o l u t i o n ,
L o c k e ' s solution, etc. N o g r o w t h in t h e s e i n o r g a n i c cul-
t u r e s took p l a c e w i t h o u t s o d i u m ehloricla. G r o w t h w a s
p r o l o n g e d a n d i n c r e a s e d by a d d i n g c a l c i u m a n d p o t a s -
sium. If m a l t o s e o r d e x t r o s e , o r p r o t e i n c l o a v a g o p r o -
ducts w e r e a d d e d p r o l i f e r a t i o n of t h e cells incrcniscid.
B y t h e m e t h o d of t r a n s f e r t o f r e s h n u t r i e n t m e d i a ,
C a r r e l h a s been able t o keep e u l t u r n s of tinfsue from t h e
h e a r t of t h e chick e m b r y o a l i v e for a l o n g p e r i o d of
y e a r s . I n a l e t t e r , r e c e n t l y received, h e s a y s : "The
 Flu. I<. lYiih.'iriii. HfHtitiiium UUIHA H\X duyn old, roinph-tfly iitcfurnorjihoMcd, with
tU*vf\o\u't\ hydranthi, Op. j>i>rin;tr'oi orUcinul IHHH; X, pd-ritiarc of outgrowth iulh<tr<'n<, to
^IJIHH. (From Wilson.)
 C O N D I T I O N S OF C E L L U L A R I M M O R T A L I T Y 63
strain of connective tissue obtained f r o m a piece of chick
heart is still alive, and will be nine years old the seven-
teenth of January, 1 9 2 1 . " F i g u r e 17 i s a photograph
showing the p r e s e n t condition of this culture. I t should
be understood that this long continued culture has gone
o;n at body temperature i n an incubator, and n o t by keep-
ing the culture at a low temperature and m e r e l y slowing
down the vital p r o c e s s e s .
This i s indeed a remarkable result. I t completes the
demonstration of the potential immortality of somatic
cells, w h e n r e m o v e d f r o m the body t o conditions which
permit of their continued existence. Somatic cells h a v e
lived and are still living outside the body f o r a far longer
time than the normal duration of life of the species f r o m
which they came. I think the present e x t e n t of Carrel's
cultures i n time fully disposes of H a r r i s o n ' s criticism
to the effect that we are " n o t justified in referring to
the cells as potentially immortal or e v e n i n speaking
of the prolongation of life by artificial means, at l e a s t
not until we are able to( keep the cellular elements alive
in cultures f o r a period exceeding the duration of life
of the organism f r o m which, they are taken. T h e r e i s
at present no r e a s o n to suppose this cannot be* done, but
it simply h a s n o t been do;ne as y e t . " I h a v e h a d m a n y
y e a r s ' experience with the domestic fowl, and have par-
ticularly studied its normal duration of life, and discus-
sed the m a t t e r w i t h competent observers of poultry. I
a m quite sure that f o r most breeds of domestic poultry
the normal a v e r a g e expectation of life at birth i s not
substantially m o r e t h a n t w o y e a r s . F o r the longest
lived races w e know this normal a v e r a g e expectation
of life cannot be over four y e a r s . I h a v e n e v e r been able
to keep a B a r r e d P l y m o u t h Rock alive m o r e than seven
64 BIOLOGY O F D E A T H
y e a r s . T h e r e a r e on r e c o r d i n s t a n c e s of fowls living to
a s m a n y as 20 y e a r s of a g e . B u t t h e s e a r e wholly excep-
t i o n a l i n s t a n c e s , u n q u e s t i o n a b l y f a r r a r e r t h a n t h e occur-
rence of c e n t e n a r i a n s a m o n g h u m a n beingn. T h e r e can
be n o q u e s t i o n t h a t t h e nine y e a r s of life of C a r r e l ' s
c u l t u r e h a s r e m o v e d w h a t e v e r v a l i d i t y m a y h a v e origin-
ally i n h e r e d i n H a r r i s o n ' s p o i n t A n d f u r t h e r t h e cul-
t u r e is j u s t a s v i g o r o u s in i t s g r o w t h t o d a y a s i t ever
w a s , a n d gives e v e r y i n d i c a t i o n of b e i n g a b l e t o g o on
indefinitely, f o r 20 o r 40, o r a n y d e s i r e d n u m b e r of y e a r s .
T h e p o t e n t i a l i m m o r t a l i t y of s o m a t i c cells h a s been
logically j u s t a s fully d e m o n s t r a t e d in a n o t h e r way a s
it h a s by t h e s e t i s s u e c u l t u r e s . S o m e n i n e t e e n y e a r s ago,
Leo L o e b first a n n o u n c e d t h e i m p o r t a n t d i s c o v e r y t h a t
p o t e n t i a l i m m o r t a l i t y of s o m a t i c calls could IKS demon-
s t r a t e d t h r o u g h t u m o r t r a n s p l a n t a t i o n s . H i s l a t e s t sum-
m a r y of t h i s w o r k m a y well be q u o t e d h e r e :

"We must remember that common, transpi&nt&bk tumor* art the

descendants of ordinary Umxw m\H, uch an w# normally find in tit
Individuals of the particular uptdt** which wn um, Ttii* ttirtttim may bo
derived from a variety of normal ti*wijum and, in guttural, the trasiifor-
mation from normal cell* into tumor call* takftti plaat tindtr th Inftutiiet
of a long continual action of various factor** ttibatidng growth. Tumor cell*
are, therefore, merely nomatic eel to which havu gaitsgd tit iiwrftftiMMl growth
energy and at the tame time somthow gained, in mmw aum, tb power to
escape the detractive eon*equcm,Trt of hoxxtoioU*iit;t. Thin ability of mr*
tain tumor* to grow in other individuals of t\w imm |>ecb* ha, rimbled
u to prove, through apparently intdlttui propagation of th*sn tumor mil*
in other individualH, that ordinary mwmiUs mVm pmsmm polllal im-
mortality in the same ens* in which protozoa mmd ^tm vn\U pmtmm
immortality' Thus tumor transplantation made ptmafble the t*UhlUhmeflt
of a fact of great biological lat*rmtf which, becauiws of the hi)iw>iomn#itlv$-
nets of normal tiue, could not be ahown in the latter,
"We wish, however, especially to emphfctti&ft tb fact that our experi-
ments did not merely prove tho immortality of tumor at!]*, \ml $4 the
ordinary tissue cells as well, the large majority or all of which can be
transformed into tumor cells. At an early stage of our
 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 65
we drew, therefore, on the basis of these experiments, the conclusion that
ordinary tissue cells are potentially immortal; notwithstanding the fact
that, especially under Weismann's influence, the opposite view had been
generally accepted, and as it seems to us, with full justiflcation, inasmuch
as no facts were known at that time which suggested the immortality
of somatic cells. It was the apparently endless transplantation of tumor
cells which proved the contrary view.
"To recapitulate what we stated al>ove: tumors are merely transformed
tissue cells. All or the large majority of adult tissues are potential tumor
cells. Tumor cells have been shown experimentally to be potentially im-
mortal, therefore tissue cells are potentially immortal.
" This wider conclusion I expressed nineteen years ago, Quite recently,
the immortality of certain connective tissue cella has been demonstrated
by Carrel through in vitro culture of these cells. Under those conditions
the tissue cells escape the mechanisms of attack to whiuh the homoiotoxins
expose the ordinary tissue cells in other individuals of the same species.
Under these conditions the reactions of the host tissue against homoiotoxins
which would have taken place in vivo, are eliminated. We rmist, however,
keep in raind that this method of proving the immortality of somatic cells
applies only to one particular, very favorable kind of cells; and it is very
doubtful, if, by cultivation in vitro, the same proof could be equally well
supplied in the case of other tissues. On the basis of tumor transplanta-
tions, on the contrary, we were able to show that a considerable variety,
perhaps the large majority of all tissue cells possess potential immortality."
T o L o c b u n q u e s t i o n a b l y belongs t h e c r e d i t f o r first
p e r c e i v i n g t h a t d e a t h w a s n o t a n e c e s s a r y i n h e r e n t con-
sequence of life i n t h e s o m a t i c cell, a n d d e m o n s t r a t i n g by
a c t u a l e x p e r i m e n t s t h a t s o m a t i c cells could, u n d e r cer-
t a i n c o n d i t i o n s , go on living indefinitely.
B e f o r e t u r n i n g t o t h e n e x t p h a s e of o u r d i s c u s s i o n ,
let u s s u m m a r i z e t h e g r o u n d we h a v e covered u p t o t h i s
p o i n t W e h a v e s e e n t h a t by a p p r o p r i a t e c o n t r o l of
conditions, i t is possible to p r o l o n g t h e life of cells a n d
t i s s u e s f a r b e y o n d t h e l i m i t s of longevity to w h i c h t h e y
would a t t a i n if t h e y r e m a i n e d i n t h e m u i t i c e l l u l a r b o d y
f r o m w h i c h t h e y came- T h i s is t r u e of a w i d e v a r i e t y
of cells a n d t i s s u e s d i f f e r e n t i a t e d in v a r i o u s w a y s . I n -
deed, t h e r a n g e of f a c t s which h a v e b e e n a s c e r t a i n e d
5
 00 I'.IOLCGY O F D H A ' i H
by e x p e r i m e n t a l w o r k In this field, p r o b a b l y w a r r a n t s
t h e conclusion t h a t thin p o t e n t i a l l o n g e v i t y i n h e r e s in
m o s t of t h e different kinds of cells of t h e m e t n z o a u body,
except thoHe which a r e e x t r e m e l y d i f f e r e n t i a t e d for p a r -
t i c u l a r functions. T o b r i n g this p o t e n t i a l i m m o r t a l i t y
to a c t u a l i t y r e q u i r e s , of c o u r s e , special c o n d i t i o n s in
each p a r t i c u l a r case. M a n y of t h e s e special conditions
have a l r e a d y been discovered for p a r t i c u l a r t i s s u e s and
p a r t i c u l a r a n i m a l s . D o u b t l e s s , in the future! m a n y m o r e
will be worked out. W e have* f u r t h e r m o r e seen t h a t in
c e r t a i n cases the physico-chemical n a t u r e of t h e condi-
tions n e c e s s a r y to i n s u r e the c o n t i n u a n c e of life h a s been
definitely w o r k e d o u t a n d is well u n d e r s t o o d . Again
this w a r r a n t s t h e e x p e c t a t i o n t h a t , with m o r e e x t e n d e d
and p e n e t r a t i n g i n v e s t i g a t i o n s in a field of r e s e a r c h
wliich is really j u s t a t its beginning, we shall u n d e r s t a n d
the p h y s i c s a n d c h e m i s t r y of p r o l o n g a t i o n of life* of cells
and t i s s u e s in a g r e a t m a n y c a s e s where* now we* know
n o t h i n g a b o u t it.
One f u r t h e r p o i n t a n d we shall h a v e clone with this
p h a s e of o u r discussion. T h e e x p e r i m e n t a l c u l t u r e of
cells a n d t i s s u e s in vitro h a s now covered p r a c t i c a l l y all
the essential tissue e l e m e n t s of the* mtAnmnn botly, even
including t h e m o s t highly differentiated of t h o s e t i s s u e s .
N e r v e cells, muscle cells, h e a r t muscle cells, spleen cells,
connective t i s s u e cells, epithelial cells f r o m v a r i o u s loca-
tions in the body, kidney cells, a n d o t h e r s h a v a all been
successfully c u l t i v a t e d in vitro. W e m a y f a i r l y s a y , I be-
lieve, t h a t t h e p o t e n t i a l i m m o r t a l i t y of a l l e s s e n t i a l cel-
l u l a r e l e m e n t s of t h e b o d y e i t h e r hits been fully
d e m o n s t r a t e d , or e k e h a s been c a r r i e d f a r e n o u g h t o
make the probability very g r e a t t h a t p r o p e r l y conducted
e x p e r i m e n t s would d e m o n s t r a t e t h e continuance* of t h e
 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 67
life of t h e s e cells in c u l t u r e to a n y definite e x t e n t . I t
is n o t to b e expected, of c o u r s e , t h a t s u c h t i s s u e s a s h a i r ,
or nails, w o u l d be c a p a b l e of i n d e p e n d e n t life, b u t t h e s e
a r e e s s e n t i a l l y u n i m p o r t a n t t i s s u e s in t h e a n i m a l econ-
omy a s c o m p a r e d w i t h t h o s e of t h e h e a r t , t h e n e r v o u s
s y s t e m , t h e k i d n e y s , etc. W h a t I a m l e a d i n g t o is t h e
b r o a d g e n e r a l i z a t i o n , p e r h a p s not c o m p l e t e l y d e m o n -
s t r a t e d yet, b u t h a v i n g r e g a r d to L e o L o e b ' s w o r k , so
near it as to m a k e little risk inhere in predicting the
final outcome, that all the essential tissues of the meta-
zoan body are potentially immortal. The reason that
they are not actually immortal, and t h a t multicellular
a n i m a l s do n o t live f o r e v e r , is t h a t i n t h e d i f f e r e n t i a t i o n
a n d s p e c i a l i z a t i o n of function of cells a n d t i s s u e s i n t h e
b o d y a s a whole, a n y i n d i v i d u a l p a r t d o e s n o t find t h e
conditions n e c e s s a r y f o r i t s c o n t i n u e d e x i s t e n c e . I n
t h e b o d y a n y p a r t is d e p e n d e n t for t h e n e c e s s i t i e s of i t s
existence, a s f o r e x a m p l e n u t r i t i v e m a t e r i a l , u p o n o t h e r
p a r t s , or p u t i n a n o t h e r w a y , u p o n t h e o r g a n i z a t i o n of
t h e b o d y as a whole. It is the differentiation and spe-
cialization of function of the mutually dependent aggre-
gate of cells and tissues which constitute the meiazoan
body which brings about death, and not any inherent or
inevitable mortal process in the individual cells them-
selves.
A n e x a m i n a t i o n of different lines of evidence h a s
led u s t o t w o g e n e r a l conclusions, viz:
a. T h a t t h e i n d i v i d u a l cells a n d t i s s u e s of t h e b o d y ,
in a n d b y t h e m s e l v e s , a r e p o t e n t i a l l y i m m o r t a l .
b . T h a t d e a t h of t h e m e t a z o a n b o d y o c c u r s , f u n d a -
m e n t a l l y , b e c a u s e of t h e w a y i n w h i c h t h e cells a n d t i s -
sues a r e o r g a n i z e d i n t o a m u t u a l l y d e p e n d e n t s y s t e m .
I s t h e r e a n y f u r t h e r a n d d i r e c t e v i d e n c e t o be h a d
68 BIOLOGY OF D E A T H
u p o n t h e second of t h e s e conclusions? S o f a r o u r evi-
dence in ita f a v o r h a s been i n d i r e c t a n d i n f e r e n t i a l ,
t h o u g h cogent BO f a r a s it goon. In t h i s connection, a
p a p e r of F r i e d e n t h a P s is of c o n s i d e r a b l e i n t e r e s t . H e
shows t h a t t h e r e is a m a r k e d c o r r e s p o n d e n c e between t h e
longevity of v a r i o u s species of a n i m a l s a m i a c o n s t a n t
of o r g a n i z a t i o n which he calls the " c e p h a l i s a t i o n f a c t o r . "
T h i s c e p h a l i s a t i o n f a c t o r in p u r e f o r m , in his s e n s e , is
given by t h e e q u a t i o n .
/ill.,,
CttphauHfition e *
factor Braini weightrT" v i
,., v
Iotal Riatuf of body prottipmntn.
Now " t o t a l m a s s of body protoplasm," an d i s t i n c t f r o m
s u p p o r t i n g s t r u c t u r e s , such a s bone etc., is obviously
difficult to d e t e r m i n e directly. B u t F r i e d e n t h a l is well
convinced t h a t , t o a first a p p r o x i m a t i o n , t h e capitalisa-
tion f a c t o r m a y bo w r i t t e n in t h i s w a y :
m % %> .. factor
Gephahwition ,-- />-;
Brain wtiight
r--t y%'3 ,
1
(Body woight)
(/omputed u p o n t h e l a t t e r basin h e etft u p t a b l e s of tin*
r e l a t i o n between cephalination f a c t o r a n d longevity f o r
m a m m a l n a n d for btrdn. I t in n o t necoHKary to r e p r o -
duce h e r e t h e l o n g tables, b u t t o s h o w t h e g e n e r a l p o i n t ,
the following table for five* selected specicm of m a m m a l s
will suffice:
TABLE 5
Relation hitmen the cephalisation fndm and inngmUy (Frkdetiikal)
Duration at lift
Munm 0.045 Hyettm
Ilnhhit .066 H ye&ri
Marniom t (VaUithriz) .216 12 ymm
Dcor If} ytar
Man 2.7 100 warn

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 conHwjiiiMUfi? cf 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 grtat 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 reemblim 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."

I t in doubtful if t h i s position in w a r r a n t e d . Binco

J e n n i n g s w r o t e t h e s t a t e m e n t quotcul, Homo new a n d
p e r t i n e n t d a t a havci a p p e a r e d in. rtigurd to a m i e r o m i -
cleate i n f u s o r i a . W o o d r u f f nml hin coAvorkers h a v e
Bhown t h a t such r a c e s m a y occur r a t h e r commonly* T h u s
Woodruff, in 1921, s a y s :
"During th pat ymr, t\w mAnt'um for wrtain f?x|H*Hm^nt* of 14
"wild" linen ni>rmsnting 6 nptcii? of hypntrichoiii* dilate* ru^^tlwi 7 Unm
(4 upecieM) with mieronucld and 7 Una* (2pi*i'kn| withtntt morpholc^iml
micromu'let. Ten of the llnm witrn all hnlmlml from a "wilcl" mnm rttitur
of thr Marnf* pwiH U runty to grandi** found in a laboratory m\%mrhim
Six of thtmci llnm wt'r amirroDiicli^ti^. A!! tf th** Unm of all of t\w
n\m>im have? brod true* with rpttpecrt to th^ rtmmM^r in qm^titin, mtd om*
amicronudeata Hnc at jrwf*ni in at th^ 1024 generation,
Similarly a eulturi* of Pammetdum mudatum, whii'h fh# pr#jn*rtt wr$lr
HuppIicKl a ytmr ago to a eotami in proUmutUwy for ttm study of tft#
failed to reviml a micronurleuii, although in othtr rmm the
wan readily d@siontrat<xl."

Now, Binco i t in t h e microttueleus which furnifthew for

the proccfls of endomixis t h e " r e s e r v e Htock of a m a t e r i a l
essential to l i f e " which J e n n i n g s <lt8eufuu?g9 i t in p l a i n
t h a t t h e existence of amicronueleate r a c e s of Protozoa.
 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 73
a t once p u t s a n e w face u p o n the whole m a t t e r . D a w s o n
h a s s t u d i e d in c o n t i n u e d c u l t u r e one of t h e s e a m i c r o n u -
cleate r a c e s of Oxytricha hymenostoma S t o k e s . H i s con-
clusion is a s follows :
"The existence of a form which not only apparently may live indefi-
nitely without conjugation, autogamy, or endomixis (assuming the possi-
bility of the latter phenomenon in an hypotrichous form), but also
apparently docs not possess the ability to undergo any of these phenomena,
brings to light an entirely new possibility in the life history of ciliates.
It has been proved quite conclusively, (Woodruff, '14), that in forms
which ordinarily conjugate, the continued prevention of this process brings
about no loss of viability if a favorable environment be provided. How-
ever, in the organism under consideration there is apparently no possi-
bility not only of conjugation or endomixis, but also of autogamy; and
thus we have from another source crucial evidence that none of these
phenomena is an indispensable factor in the life-history of this hypo-
trichous form."

I n the light of t h e s e clean c u t a n d definite r e s u l t s

one is m o r e d i s p o s e d t h a n w a s f o r m e r l y t h e case to
a c c e p t a t t h e i r face v a l u e t h e r e s u l t s of E n r i q u e s w i t h
Glaucoma pyriformis, a n d t h o s e of H a r t m a n n w i t h
Eudorina elegans, in which r e p r o d u c t i o n w e n t on indef-
initely w i t h i m d i m i n i s h e d v i g o r a n d n o evidence of a n y
p r o c e s s c o m p a r a b l e t o endomixis.
A l t o g e t h e r , it Beems to m e t h a t the w e i g h t of t h e evi-
dence n o w is t h a t in t h e P r o t o z o a , senescence ( o r d e a t h ) ,
is n o t a necessary or inevitable consequence of life.
Given t h e a p p r o p r i a t e a n d n e c e s s a r y c o n d i t i o n s of envi-
r o n m e n t , t r u e i m m o r t a l i t y t h e absence of b o t h senes-
cence a n d n a t u r a l d e a t h , each defined i n t h e m o s t critical
m a n n e r i s in f a c t t h e r e a l i t y f o r a n u m b e r of f o r m s .
T u r n i n g t o t h e m e t a z o a n side of t h e c a s e , t h e evidence
r e g a r d i n g senescence i s equally cogent. I n t h e first place,
in t h e l o n g e s t c o n t i n u e d in vitro t i s s u e c u l t u r e s k n o w n
( t h o s e of C a r r e l ) t h e r e is, a s a l r e a d y s t a t e d , n o a p p e a r -
74 BIOLOGY OF D E A T H
ance of senescence i n t h e cells. B u t i t m a y be objected
t h a t a n e l e m e n t of u n c e r t a i n t y is injected into t h e case 3
b y t h e f a c t t h a t , as C a r r e l a n d E b e l i n g h a v e lately dis-
cussed i n some detail, it h a s been n e c e s s a r y i n c a r r y i n g
a l o n g this long-continued c u l t u r e to a d d r e g u l a r l y t o th
c u l t u r e m e d i u m a small a m o u n t of iiembryonic juice.'
One m i g h t u r g e t h a t , b u t f o r t h e ' ' embryonic j u i c e , ' ' cellu-
l a r senescence a n d d e a t h would have a p p e a r e d . Bui
s u p p o s e t h i s t o b e g r a n t e d fully. I t does n o t m e a i
t h a t senescence is a n e c e s s a r y a n d inevitable consequence
of life, b u t only t h a t to realize a p o t e n t i a l immortality
t h e cells m u s t h a v e an a p p r o p r i a t e environment, on<
element of which, is p r e s u m a b l y some chemical combinatioi
which, so f a r , one h a s supplied only t h r o u g h " e m b r y
onic j u i c e . "
A n e n t i r e l y different s o r t of evidence a n d one o
g r e a t significance is found i n the facts of clonal p r o p a g a
tion of p l a n t s , well known t o h o r t i c u l t u r i s t s . A n individ
u a l a p p l e t r e e g r o w s old, a n d eventually dies, a s a tre*
B u t a t all p e r i o d s of its life, including all s t a g e s o
senescence u p t o t h e t e r m i n a l one, death, i t p r o d u c e
shoots each s p r i n g . If one of t h e s e shoots is g r a f t e d t
a n o t h e r r o o t , i t will, in t h e p a s s a g e of time, m a k e firs
a y o u n g t r e e , t h e n a middle a g e d t r e e , a n d finally a n ole
senescent t r e e ; which, i n t u r n , will m a k e n e w shooti
which m a y , i n t u r n , be g r a f t e d t o n e w r o o t s , a n d so o
ad infinitum. I t is n o t even absolutely n e c e s s a r y the
the shoot be g r a f t e d to a n e w r o o t ; though, of cours<
this i s t h e m a n n e r i n which t h e g r e a t m a j o r i t y of on
o r c h a r d s a r e , i n fact, p r o p a g a t e d , a n d h a v e been sine
the b e g i n n i n g of h o r t i c u l t u r a l h i s t o r y . A n y o n e w h o :
f a m i l i a r w i t h t h e woods of N e w E n g l a n d , n o t too f a r f roi
s e t t l e m e n t s , h a s seen a p p l e t r e e s in the woods w h e r e
 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 75
shoot, w h o s e c o n t i n u i t y w i t h the b a s e of i t s p a r e n t t r e e
h a s n e v e r b e e n b r o k e n , m a k e s a n e w t r e e a f t e r t h e old one
h a s d i e d i n d e e d i n s o m e cases t h e s h o o t h a s h e l p e d the
m o r t i f e r o u s p r o c e s s by t h e v i g o r o u s c r o w d i n g of youth.
I n t h i s w h o l e p i c t u r e h o w f a r e s a n y i d e a of t h e necessity
o r inevitableness of cellular ( s o m a t i c ) senescence? S u c h
a n i d e a p l a i n l y h a s n o place in t h e r e a l i t i e s of t h e con-
t i n u e d existence of a p p l e t r e e s .
F r o m t h e s e facts i t is a logically c o g e n t i n d u c t i o n t o
i n f e r t h a t w h e n cells show t h e c h a r a c t e r i s t i c s e n e s c e n t
c h a n g e s w h i c h w e r e discussed i n t h e p r e c e d i n g c h a p t e r ,
it is b e c a u s e t h e y a r e reflecting in t h e i r m o r p h o l o g y a n d
p h y s i o l o g y a consequence of t h e i r m u t u a l l y d e p e n d e n t
a s s o c i a t i o n i n t h e b o d y a s a whole, a n d n o t a n y n e c e s s a r y
progressive process inherent in themselves. I n other
w o r d s , m a y w e n o t t e n t a t i v e l y , in t h e l i g h t of o u r p r e s e n t
knowledge, r e g a r d senescence as a phenomenon appear-
ing in the multicellular body as a whole, a s a r e s u l t of
t h e fact t h a t it is a differentiated a n d c o n f e r e n t i a t e d (to
employ a useful t e r m lately i n t r o d u c e d b y B i t t e r ) mor-
phologic and dynamic organization. This phenomenon
is reflected m o r p h o l o g i c a l l y in the c o m p o n e n t cells. B u t
it does n o t p r i m a r i l y o r i g i n a t e in a n y p a r t i c u l a r cell
because of t h e f a c t t h a t t h a t cell is old in t i m e , o r b e c a u s e
t h a t cell in a n d of itself h a s been a l i v e ; n o r does i t occur
in t h e cells w h e n t h e y a r e r e m o v e d f r o m t h e m u t u a l l y
d e p e n d e n t r e l a t i o n s h i p of t h e o r g a n i z e d b o d y a s a
whole a n d given a p p r o p r i a t e physico-chemical condi-
tions. I n s h o r t , senescence a p p e a r s n o t t o be a p r i m a r y
a t t r i b u t e of t h e physiological economy of cells a s such.
If t h i s conception of t h e p h e n o m e n o n of senescence
is c o r r e c t i n i t s m a i n f e a t u r e s , it s u g g e s t s t h e essential
futility of a t t e m p t i n g t o i n v e s t i g a t e its c a u s e s b y p u r e l y
76 BIOLOGY O F D E A T H
cytological m e t h o d s On t h e o t h e r h a n d , by clearinj
a w a y t h e u n e s s e n t i a l e l e m e n t s , It i n d i c a t e s w h e r e researel
i n t o t h e p r o b l e m of cauHation of m m e m m e e m a y b<
profitable.
A n e x t r e m e l y i n t e r c n t i n g c o n t r i b u t i o n t o the problcn
of senescence h a s been m a d e by C a r r e l a n d E l u d i n g ii
t h e i r m o a t r e c e n t p a p e r , in which t h e y s h o w t h a t t h e r a t
of m u l t i p l i c a t i o n of fibroblantn hi vitro, ami t h e d u r a t i o .
of life of such culture*), in i n v e r s e l y p r o p o r t i o n a l to th
a g e of t h e a n i m a l from which t h e wernm for the? e n l t n r
m e d i u m is taken. T h e s e r e s u l t s a r e of uch cmiKidcrabl
i n t e r e s t t h a t it will be well t o q u o t e in full thr* Humznar
of t h e m given by t h e ant h o r n :
"Pure culture of fibrohU*U <ii|layt*fl mark**! flitft*ri*nr*s In thu
activity in this jplanma of young, middle ftgwJ, ami old c?liirktftit. Tint ra
of cell multiplication varifd in invent* ratio t*> th* %#$ <*f th? futimtti frn
which the plttHimi wa t&k?tt. TItrr^ wan a iliffliiil^ r*laUt*n l*tw^i*tt I
ag^ of the animal and the nmtmnt nt tiw ttemv prttltwrA in i\* \thim
in a givim time. Th< elmrt obtain***! l*y |>f*4fin$f tb* tntt* *f n4l jr4ifi*y
tlon in orditmUm, mid the i*g t>f tht? anlma! In JtWiwit*, nhfiwit! tlmt t
rate of growth dtacwaaed mart quickly tlmn th@ iigt iiirrm4. The l
crmm in th r&U of growth wa BO iwt 4?nt, dtiritig thi flrnt 3 ymrn
life, while in th following 0 ynrn it wait witly n jw*r rt*nt Wh* t
duration of the life of the liilturw lit thu four jiliistmi* mm mmpMmU
curve wan obtalncid which nhowiil about tha muw h&rmy*thUm T
duration of life of th fihrib!ai*t in xitm v&t'mi In iiivir^ ratiti tUi t
aga of th animal, and dmriand mor* c|iiltkly than t!it ag* mrnrnmnh
"As the difftrfnrtu in tht ummmt of ntnv tinniit prmhuml In I
plHMrna of young, middlt KH1, anil old rhfclctinft wi*r largr* thu
of a pure culture of fthrMmnU miM im ampUnymH m n rragimt for
ing eert&in ehangen oecwrring in th ptsum* uncter thu ifH*nr<* of
41
A comparative study of the growth of fittntbtimu U% mmttt*
no erum, and nerum umkr low attdi high mnmntruiluim w*m mml, In mi
to ascertain whether the decrmiing rate of i*#ll mtiltiptitmtfnti wan due
the IOMS of an neenlerating im-U>rt or td th# li!rmiwi of an ihibitiic t*
In high and low conetntrationit of th mtum of ytitmg anittlf no
in the rate of multiplication of fibrobkwi* wm ohrrv<*d, 7'hb
that the serum of an actively growing animal did not crmUin any
 T H E CHANCES OF DEATH 77
erating agent. The same experiments were repeated with the serum of
a 3 year old and a 9 year old chicken. The medium made of a high
concentration of serum had a markedly depressing effect on the growth,
and this effect was greater in the serum of the older animal.
"The results of the experiments showed in a very definite manner that
certain changes occurring in the serum during the course of life can be
detected by modifications in the rate of growth of pure cultures of fibro-
blasts and that these changes are characterized by the increase of an
inhibiting factor, and not by the loss of an accelerating one. It appeared,
therefore, that the substances which greatly accelerate the multiplication
of fibroblasts and are found in the tissues do not exist in the blood serum,
or are constantly shielded by more active inhibiting factors. The curve
which expresses the variations of the inhibiting factor in function of the
age was compared with that showing the variations of the rate of healing
of a wound according to the age of the subject. For wounds of equal size,
the index of cicatrization, which expresses the rate of healing, varies in
inverse ratio to the age. The different values of the index of cicatrization
of a wound 40 sq. cm. in area, taken from measurements made by du Notiy,
were plotted in ordinates, and the age of the subject in abscissas. The
curve showed a decrease in the activity of cicatrization, which resembled
the decrease in the rate of growth of fibroblasts in function of the age
of the janimal. This suggested the existence of a relation between the
factors determining both phenomena."

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 physieochemiea
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

THE LIFE TABLE

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 ) .

CHAN0K8 INT EXPECTATION OF MKB

I wish t o pasH in g r a p h i c review sonn* of t h e s e life
tables in o r d e r t o call a t t e n t i o n in vivid f o r m t o a n i m p o r -
t a n t f a c t a b o u t t h e d u r a t i o n of h u m a n lifu. I n o r d e r to
b r i n g o u t t h e p o i n t w i t h which we a r e here? c o n c e r n e d i t
will be n e c e s s a r y to m a k e use of a n o t h e r function of t h e
m o r t a l i t y t a b l e t h a n e i t h e r t h e / o r dM lines which a r e
shown in F i g u r e 18. I w i s h t o dimmnn e x p e c t a t i o n of
life a t each a g e . T h e e x p e c t a t i o n of life a t a n y a g e i s
defined in a c t u a r i a l science a s t h e m e a n o r a v e r a g e n u m b e r
of y e a r s of s u r v i v a l of p e r s o n s alive a t t h e s t a t e d age*.
I t is g o t by d i v i d i n g t h e t o t a l m i r v i v o r - y e a r s of a f t e r life
by t h e n u m b e r s u r v i v i n g a t t h e s t a t e d a g e . O r , if we lc*t
e* d e n o t e w h a t h called t h e c u r t a t e e x p e c t a t i o n of life
l^^^^^-^^^^^^^^....

To a first a p p r o x i m a t i o n , sufficiently accurate* for our

p r e s e n t p u r p o s e s , t h e total e x p e c t a t i o n of life, called e9 *
may b eo b t a i n e d from the curtate expectation b y the
simple relation
 T H E CHANCES OF D E A T H
TABLE 6
Changes in expectation of life from the seventeenth century to
the present time
Average length of life remaining Average length of life remaining
to each one alive at beginning to each one alive at beginning
of age interval of age interval
Age Age
Breslau, Carlisle, Breslau, Carlisle,
17th 18th U.S.1910 17th 18th U.S.1910
century century century century
0- 1 33.50 38.72 51.49 50- 51 16.81 21.11 20.98
1- 2 38.10 44.67 57.11 51- 52 16.36 20.39 20.28
2- 3 39.78 47.55 57.72 52- 53 15.92 19.68 19.58
3- 4 40.75 49.81 57.44 53- 54 15.48 18.97 18.89
4- 5 41.25 50.76 56.89 54- 55 14.99 18.27 18.21
5- 6 41.55 51.24 56.21 55- 56 14.51 17.58 17.55
6- 7 41.62 51.16 55.47 56- 57 14.02 16.89 16.90
7- 8 41.16 50.79 54.69 57- 58 13.54 16.21 16.26
8- 9 40.95 50.24 53.87 58- 59 13.06 15.55 15.64
9-10 40.50 49.57 53.02 59- 60 12.57 14.92 15.03
10-11 39.99 48.82 52.15 60- 61 12.09 14.34 14.42
11-12 39.43 48.04 51.26 61- 62 11.62 13.82 13.83
12-13 38.79 47.27 50.37 62- 63 11.14 13.31 13.26
13-14 38.16 46.50 49.49 63- 64 10.67 12.81 12.69
14-15 37.51 45.74 48.60 64- 65 10.20 12.30 12.14
15-16 36.86 44.99 47.73 65- 66 9.73 11.79 11.60
16-17 36.22 44.27 46.86 66- 67 9.27 11.27 11.08
17-18 35.57 43.57 46.01 67- 68 8.81 10.75 10.57
18-19 34.92 42.87 45.17 68- 69 8.36 10.23 10.07
19-20 34.26 42.16 44.34 69- 70 7.91 9.70 9.58
20-21 33.61 41.46 43.53 70- 71 7.53 9.17 9.11
21-22 32.95 40.75 42.73 71- 72 7.17 8.65 8.66
22-23 32.34 40.03 41.94 72- 73 6.85 8.16 8.22
23-24 31.67 39.31 41.16 73- 74 6.56 7.72 7.79
24-25 31.00 38.58 40.38 74- 75 6.25 7.33 7.38
25-26 30.38 37.86 39.60 75- 76 5.99 7.00 6.99
26-27 29.76 37.13 38.81 76- 77 5.79 6.69 6.61
27-28 29.14 36.40 38.03 77- 78 5.71 6.40 6.25
28-29 28.51 35.68 37.25 78- 79 5.66 6.11 5.90
29-30 27.93 34.99 36.48 79- 80 5.67 5.80 5.56
30-31 27.35 34.34 35.70 80- 81 5.74 5.51 5.25
31-32 26.76 33.68 34.93 81- 82 5.86 5.20 4.96
32-33 26.18 33.02 34.17 82- 83 6.02 4.93 4.70
33-34 25.59 32.36 33.41 83- 84 5.85 4.65 4.45
34-35 25.05 31.68 32.66 84- 85 4.39 4.22
35-36 24.51 31.00 31.90 85- 86 4.12 4.00
36-37 23.97 30.32 31.16 86- 87 3.90 3.79
37-38 23.43 29.63 30.42 87- 88 3.71 3.58
38-39 22.88 28.95 29.68 88- 89 3.59 3.39
39-40 22.33 28.27 28.94 89- 90 3.47 3.20
40-41 21.78 27.61 28.20 90- 91 3.28 3.03
41-42 21.23 26.97 27.46 91- 92 3.26 2.87
42-43 20.73 26.33 26.73 92- 93 3.37 2.73
43-44 20.23 25.71 25.99 93- 94 3.48 2.59
44-45 19.72 25.08 25.26 94- 95 3.53 2.47
45-46 19.22 24.45 24.54 95- 96 3.53 2.35
46-47 18.72 23.81 23.82 96- 97 3.46 2.24
47-48 18.21 23.16 23.10 97- 98 3.28 2.14
48-49 17.71 22.50 22.39 98- 99 3.07 2.04
49-50 17.25 21.81 21.69 99-100 2.77 1.95
l a each of t h e s e r i e s of d i a g r a m s w h i c h follow t h e r e
is p l o t t e d t h e a p p r o x i m a t e v a l u e of t h e e x p e c t a t i o n of
84 BIOLOGY O F DEATH
life f o r s o m e g r o u p of people at mint* p r r i o d In \\w mow
or less r e m o t e pant, and for c o m p a r i s o n th ixp'Mnti<n
of life, e i t h e r from ( J l m e r ' n inhh\ for lit** p o p u l a t i o n of
the U n i t e d KtntoH Rt^'int r a t i o n A r e a in HMo tlu- >xpi*r-
tation of life of o u r people now, in 8 h o r t - - o r i'i|uivah*nt
figure** for a m o d e r n Hn^linh o r F r r i i r h pptla1inn.
B e c a u s e of t h e eonHiderahle i n t e r t ^ t of t h e tuatU*r v
and the fact t h a t the? d a t a a r e u o t eanily availalilf* t*

n, T a b l e fi in i n s e r t e d , Kivinn; t h e f*xpt*rtntitn# 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 iwcrp nttfitcti?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 lw*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 45middle lifeis 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

t u r y a s i t is now. A t age 47 t h e eighteenth c e n t u r y 1

c r o s s e s t h a t f o r the twentieth c e n t u r y , a n d w i t h a i
t r i f l i n g exceptions, n o t a b l y in t h e y e a r s f r o m 56 to
t h e expectation of life f o r all h i g h e r ages w a s gre^
t h e n t h a n i t i s now. W e see i n the e i g h t e e n t h cent
t h e s a m e kind of r e s u l t as w a s indicated in the se^
t e e n t h , only differing i n d e g r e e .
MILNE'S CARLISLE 1760 - 1767' UFZ TABLE

S5

50

45 1 ^\
N
v
<
30
s
6 0 s
/S

10 si
5

0 =
YEAR'S OF LIFE
Fia. 20Comparing 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
\ , \
" ! * * t ..:...
X . ,
b

"\ <. __
t s
"**'
\
"1 i i i ;""*?** ;:
":>^ '. !
. 1

th rjii #f lift* #4 tli*f *4

day Atti JU V

F o r comparison, the expectation of lift* from (Hover's

1910 United State?** life table* is inserted.
It will be mm a t once* that the general sweep of the
line in of the same sort that we haves already obtiervcMl
in the case of the seventeenth century table*. In the
early years of life the expectation was far below that of
the present time, but somewhere between a#em 65 and 70
the Kgyptian line crosses the modern American line, and
from that period on the individuals living in Kgypt at
about the time of the birth of Christ could apparently look
forward to a longer remaimngduratioii of lift%on the aver-
 T H E CHANCES OF D E A T H 89
age, t h a n can the A m e r i c a n of t h e p r e s e n t d a y . P e a r s o n ' s
comment on this f a c t i s w o r t h q u o t i n g . H e s a y s : " I n
the course of t h o s e c e n t u r i e s m a n m u s t h a v e g r o w n r e -
m a r k a b l y fitter t q his e n v i r o n m e n t , o r else h e m u s t h a v e
fitted his e n v i r o n m e n t i m m e a s u r a b l y b e t t e r t o himself.
N o civilized c o m m u n i t y of t o - d a y could show such, a c u r v e
a s t h e civilized R o m a n o - E g y p t i a n s of 2,000 y e a r s a g o
exhibit. "We h a v e h e r e e i t h e r a s t r o n g a r g u m e n t f o r t h e
survival of t h e p h y s i c a l l y fitter m a n o r f o r t h e s u r v i v a l
of t h e civilly fitter society. E i t h e r m a n i s c o n s t i t u t i o n -
ally fitter to s u r v i v e to-day, o r h e is m e n t a l l y fitter, i.e.,
b e t t e r able t o o r g a n i z e h i s civic s u r r o u n d i n g s . B o t h con-
clusions p o i n t p e r f e c t l y definitely t o a n e v o l u t i o n a r y
p r o g r e s s . . . . T h a t t h e e x p e c t a t i o n of life f o r a
E o m a n o - E g y p t i a n o y e r 68 w a s g r e a t e r t h a n f o r a m o d e r n
E n g l i s h m a n or w o m a n i s w h a t w e m i g h t expect, f o r w i t h
the m o r t a l i t y of y o u t h a n d of m i d d l e a g e e n o r m o u s l y
emphasized only t h e v e r y s t r o n g e s t would s u r v i v e t o
this age. O u t of 100 E n g l i s h alive a t 10 y e a r s of a g e 39
survive t o be 6 8 ; o u t of 100 R o m a n o - E g y p t i a n s n o t 9
survived. L o o k i n g a t t h e s e t w o c u r v e s w e r e a l i z e a t a
glance either t h e g r e a t p h y s i c a l p r o g r e s s of m a n , which
enables h i m f a r m o r e effectually t o w i t h s t a n d a hostile
environment, o r t h e g r e a t social a n d s a n i t a r y p r o g r e s s
he h a s m a d e which enables h i m t o m o d i f y t h e e n v i r o n -
ment. I n e i t h e r c a s e w e c a n definitely a s s e r t t h a t 2,000
y e a r s h a s m a d e h i m a m u c h ' fitterJ b e i n g . I n t h i s com-
p a r i s o n i t m u s t be r e m e m b e r e d t h a t w e a r e n o t p l a c i n g
a civilized r a c e a g a i n s t a b a r b a r i c t r i b e , b u t c o m p a r i n g
a m o d e r n civilization w i t h o n e of t h e h i g h e s t t y p e s of
ancient civilization.''
Macdonell w a s a b l e t o continue t h i s i n v e s t i g a t i o n on
much m o r e extensive m a t e r i a l e x t r a c t e d f r o m t h e Corpus
90 BIOLOGY OF D E A T H
Inscriptionum Latinarum of t h e B e r l i n A c a d e m y , which
gives r e c o r d s as to a g e of d e a t h for m a n y t h o u s a n d
R o m a n citizens d y i n g , for t h e m o s t p a r t , within the first
t h r e e o r f o u r c e n t u r i e s of t h e C h r i s t i a n e r a . H i s mate-
rial m a y , t h e r e f o r e , be taken to r e p r e s e n t the conditions
a few c e n t u r i e s l a t e r than those* of P e a r s o n ' s R o m a n o -
E g y p t i a n p o p u l a t i o n . Mncdonell wan abl<* t o calculate

'"
NJ 1 ...1 . i 1
. . . I , I
J.. i
,
*
-

. . .

4., ....

. * t * i
t \

j TvV j i i . ^

. w .i~- "KSc * i
i
i '*^v i ! I
] |.. -.i i

.. _ f
|
j "ill, I
J *
i
|
H--j j *-- -j ]
1
-j-p- ! P'-rLLLL i mm
2

i, 2 2 . - " - < " ' c o m p a r i n g th* i

i with thiit nf

t h r e e tables of e x p e c t a t i o n of lifo-the first f o r R o m a n

citizens living in t h e city of Homes itacslf; nnconii for
those living in t h e p r o v i n c e s of l l t s p a n i a nntl LtiHitnnia;
a n d t h i r d , for those living in Africa- T h e r e s u l t s a r e
plotted a g a i n s t t h e U n i t e d St&teB 1010 d a t a , us before,
in F i g u r e s 22, 23, a n d 24.
F i g u r e 22 r e l a t e s t o i n h a b i t a n t s of t h e city of Home
itself. T h e d e a t h s from which t h e e x p e c t a t i o n * a r e
calculated r u n i n t o t h e t h o u s a n d s , a n d f o r t u n a t e l y one
is able to s e p a r a t e m a l e s a n d females* A s in PearBon**
case, which we h a v e j u s t e x a m i n e d , m o d e r n A m e r i c a n
 T H E CHANCES OF DEATH 91
d a t a a r e e n t e r e d for c o m p a r i s o n . I t will be n o t e d a t
once t h a t j u s t a s i n t h e R o m a n o - E g y p t i a n p o p u l a t i o n the
expectation of life of i n h a b i t a n t s of a n c i e n t E o m e w a s ,
in the early y e a r s of life, a p p a r e n t l y i m m e n s e l y i n f e r i o r to
t h a t of the m o d e r n p o p u l a t i o n . F r o m a b o u t t h e a g e of 60
on, however, t h e e x p e c t a t i o n of life a p p e a r s t o h a v e been
b e t t e r t h e n t h a n now. C u r i o u s l y enough, t h e expectation

55 P\ UNITED i. TA7CS
SO
\

45
s.
40

35

JO

\
\
20
rtMA
15

MSP/W/A AND LUSI TANIA

10
f \ f\
5 1
.'
i JO 15 20 25 30 35 4<0 45 50 56 60 65 T) 7L5 60 85 SO 05 JOO

YEARS OT A6C

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.

of life of f e m a l e s w a s p o o r e r a t p r a c t i c a l l y all a g e s of life

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

state of affairs. M a c d o n e l l believes this difference t o b e

real a n d t a i n d i c a t e t h a t t h e r e w e r e special influences

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

AS

.J-M

AF <?ICA

^ ^
"*
% *

10

- .
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

table. T h e striking thing, h o w e v e r , i s t h a t a t a b o u t a g e

 T H E CHANCES OF D E A T H 93
40 t h e lines c r o s s , a n d f r o m t h e n on t h e e x p e c t a t i o n of
life w a s definitely s u p e r i o r i n t h e e a r l y y e a r s of t h e
C h r i s t i a n e r a t o w h a t i t is now.
I t s h o u l d b e said t h a t t h e c u r i o u s z i g z a g g i n g of t h e
lines i n all of t h e s e R o m a n tables of M a c d o n e l l i s due t o
t h e t e n d e n c y , which a n c i e n t R o m a n s a p p a r e n t l y h a d i n
common with present day American negroes, towards
h e a v y g r o u p i n g on t h e even m u l t i p l e s of 5 in t h e s t a t e -
m e n t of t h e i r a g e s .
S u m m a r i z i n g t h e whole m a t t e r we see t h a t d u r i n g a
p e r i o d of a p p r o x i m a t e l y 2,000 y e a r s m a n ' s e x p e c t a t i o n
of life a t b i r t h a n d s u b s e q u e n t e a r l y a g e s h a s a p p a r e n t l y
been s t e a d i l y i m p r o v i n g , while a t t h e s a m e t i m e h i s expec-
t a t i o n of life a t a d v a n c e d ages h a s b e e n s t e a d i l y w o r s e n i n g .
T h e f o r m e r p h e n o m e n o n m a y p r o b a b l y be a t t r i b u t e d essen-
tially t o e v e r i n c r e a s i n g k n o w l e d g e of h o w best t o cope
w i t h t h e lethal forces of n a t u r e . * P r o g r e s s i v e l y b e t t e r
s a n i t a t i o n , in t h e b r o a d e s t sense, d o w n t h r o u g h t h e c e n t u r -
ies h a s s a v e d f o r a time the lives of e v e r m o r e a n d m o r e
babies a n d y o u n g people who f o r m e r l y could n o t with-
s t a n d t h e u n f a v o r a b l e conditions t h e y m e t , a n d died in
consequence r a t h e r p r o m p t l y . B u t j u s t b e c a u s e t h i s p r o -
cess t e n d s t o p r e s e r v e t h e weaklings, w h o w e r e speedily
eliminated u n d e r t h e r i g o r o u s action of u n m i t i g a t e d n a t -

* No absolute reliance can, of course, he put upon Macdonell's or

Pearson's curves. Besides laboring under the serious actuarial difficulty
of being expectations calculated from a knowledge of deaths alone, the
randomness of the sampling, even on that basis, is extremely doubtful.
The only real evidence that the*e Roman curves represent a rough pic-
ture of the truth as to expectation of life in those days, arises from the
consideration that they show a difference from present-day expectations
which is of the same kind as that which is found between populations of
one and two centuries ago and the present, and of a greater amount, as
would be expected from the longer time interval, and from what we know
ha occurred in the material development of civilization in the meantime.
94 BIOLOGY OF DEATH
u r a l selection, t h e r e a p p e a r now in t h e h i g h e r age g r o u p s
of t h e p o p u l a t i o n m a n y w e a k e r i n d i v i d u a l s t h a n f o r m e r l y
ever g o t t h e r e . Consequently t h e a v e r a g e expectation
of life a t a g e s b e y o n d s a y 60 to 70 is n o t n e a r l y so good
now a s i t w a s u n d e r t h e m o r e r i g o r o u s regime of ancient
times. T h e n , a n y i n d i v i d u a l w h o a t t a i n e d a g e 70 w a s
the s u r v i v i n g r e s u l t a n t of a "bitterly d e s t r u c t i v e p r o c e s s
of selection. T o r u n successfully t h e g a u n t l e t of e a r l y
and m i d d l e life, he n e c e s s a r i l y h a d t o h a v e a n e x t r a o r -
d i n a r i l y v i g o r o u s a n d r e s i s t a n t constitution. H a v i n g
come t h r o u g h successfully to 70 y e a r s of age i t is no m a t -
t e r of w o n d e r t h a t his p r o s p e c t s w e r e for a longer old
age t h a n his d e s c e n d a n t s of the same age to-day can look
f o r w a r d to. Biologically, these expectation of life c u r v e s
give u s t h e first i n t r o d u c t i o n to a p r i n c i p l e which w e
shall find a s w e go on t o be of the v e r y f o r e m o s t i m p o r -
tance in fixing t h e s p a n of h u m a n longevity, n a m e l y t h a t
inherited constitution fundamentally and primarily de-
termines how long an individual will live.

ANALYSIS OF THE LIFE TABLE

I shall n o t develop this p o i n t f u r t h e r now, b u t i n s t e a d
will t u r n b a c k t o consider briefly c e r t a i n f e a t u r e s of t h e
dx line of a life t a b l e . F i g u r e 18 shows t h a t this line,
which gives t h e n u m b e r of d e a t h s o c c u r r i n g a t each a g e ,
h a s t h e f o r m of a v e r y much s t r e t c h e d l e t t e r S r e s t i n g
on i t s back. S o m e y e a r s ago, P e a r s o n u n d e r t o o k t h e
a n a l y s i s of t h i s complex curve, a n d d r e w c e r t a i n i n t e r -
esting conclusions a s t o the f u n d a m e n t a l biological c a u s e s
lying b e h i n d i t s c u r i o u s sinuosity. H i s r e s u l t s a r e s h o w n
in F i g u r e 25.
H e r e g a r d e d t h e dm line of t h e life table as a c o m p o u n d
curve, a n d b y s u i t a b l e m a t h e m a t i c a l a n a l y s i s b r o k e i t u p
 T H E C H A N C E S OF D E A T H 95
into five c o m p o n e n t f r e q u e n c y c u r v e s . T h e d a t a w h i c h h e

u s e d w e r e f u r n i s h e d b y t h e dx line of O g l e ' s life table,

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

...
..... :

- -

1
-

i
"OX t -
\

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.

This component, according to Pearson's graduation,

accounted for 484.1 deaths out of the total of 1,000, or
nearly one-half of the whole. Its range extends from
u n d e r 20 y e a r s of a g e to the u p p e r limit of life, at approx-
imately 106 years. The second component includes the
deaths of middle life. This is the smooth curve having
its modal point between 40 and 45 years at the point on
the base marked O2. Its range extends from about 5
years of age to about 65. It accounts for 175,2 deaths
out of the total of 1,000. It is a long, m u c h spread out
 06 BIOLOGY O F D E A T H
c u r v e , e x h i b i t i n g g r e a t v a r i a b i l i t y . T h e t h i r d compi
n e n t is m a d e u p by t h e d e a t h s of y o u t h . T h i s accoum
f o r 50.8 d e a t h s o u t of t h e t o t a l of a t h o u s a n d , a n d ii
r a n g e e x t e n d s f r o m a b o u t t h e t i m e of b i r t h to n e a r l y 4
y e a r s . I t s m i d - p o i n t is b e t w e e n 20 a n d 25 y e a r s , a n d
exhibits less v a r i a b i l i t y t h a n e i t h e r t h e m i d d l e life o r t l
old a g e c u r v e s . T h e f o u r t h c o m p o n e n t , t h e m o d a l poii
of which is a t t h e p o i n t on t h e b a s e of t h e d i a g r a m m a r k e
04 c o v e r s t h e childhood m o r t a l i t y . I t account8 for 46
d e a t h s o u t of t h e t o t a l of 1,000. K B r a n g e a n d variabilii
a r e obviously less t h a n t h o s e of a n y of t h e o t h e r thrc
c o m p o n e n t s so f a r c o n s i d e r e d . T h e last, excessively ske
c o m p o n e n t , is t h a t wliich d e s c r i b e s t h e m o r t a l i t y of h
fancy. I t in given by a J s h a p e d c u r v e a c c o u n t i n g fc
245.7 d e a t h s a f t e r b i r t h , a n d a n a n t e n a t a l m o r t a l i t y c
605. I n o r d e r t o g e t a n y fit a t all f o r t h i s p o r t i o n of ti:
m o r t a l i t y c u r v e i t is n e c e s s a r y t o a s s u m e t h a t the* cleati
in utero a n d t h o s e of t h e first m o n t h s a f t e r b i r t h a r e
homogeneous connected group.
S u m m i n g all t h e s e c o m p o n e n t s t o g e t h e r it is sec
t h a t the r e s u l t i n g s m o o t h c u r v e v e r y closely fits the s e n *
of small circles which a r e t h e o r i g i n a l o b s e r v a t i o n
F r o m t h e s t a n d p o i n t m e r e l y of c u r v e fitting no bettc
r e s u l t t h a n t h i s could be h o p e d for, B u t a b o u t i t s h\<
logical significance t h e e a s e is n o t q u i t e s o clear, a s ^
shall p r e s e n t l y set*.
P e a r s o n himself t h i n k s of t h e s e five c o m p o n e n t s <
t h e m o r t a l i t y c u r v e a s t y p i f y i n g five D e a t h s , ghootin
with different w e a p o n s , a t different s p e e d s a n d w i t h di:
f a r i n g d e g r e e s of p r e c i s i o n a t t h e p r o c e s s i o n of h u m a
b e i n g s c r o s s i n g t h e B r i d g e of Life, T h e first D e a t h i
a c c o r d i n g t o P e a r s o n , a m a r k s m a n of d e a d l y a i m , coi
c e n t r a t e d fire, a n d u n r e m i t t i n g d e s t r u e t i v e n e s s . H e kill
 T H E CHANCES OF D E A T H 97
before b i r t h a s well a s a f t e r a n d m a y b e conceived a s
b e a t i n g d o w n y o u n g lives w i t h t h e bones of t h e i r ances-
t o r s . T h e second m a r k s m a n who a i m s a t childhood h a s
a n e x t r e m e l y c o n c e n t r a t e d fire, which m a y be typified
b y t h e m a c h i n e gun. Only because of t h e c o n c e n t r a t i o n
of t h i s fire a r e w e able t o p a s s t h r o u g h i t w i t h o u t a p p a l -
l i n g loss. T h e t h i r d m a r k s m a n D e a t h , w h o s h o o t s a t
youth has not a very deadly or accurate weapon, perhaps
a bow a n d a r r o w . T h e fire of t h e f o u r t h m a r k s m a n is
slow, s c a t t e r e d a n d n o t v e r y d e s t r u c t i v e , s u c h a s m i g h t
r e s u l t front, a n old f a s h i o n e d b l u n d e r b u s s . T h e l a s t D e a t h
plies a rifle. N o n e escapes his s h o t s . H e a i m s a t old age
but sometimes hits youth. H i s unremitting activity
m a k e s his toll l a r g e .
W e m a y l e t P e a r s o n s u m t h e whole m a t t e r u p i n his
own w o r d s : i ' O u r i n v e s t i g a t i o n s on t h e m o r t a l i t y s t a t i s -
tics h a v e t h u s led u s t o some v e r y definite conclusions
w i t h r e g a r d t o t h e chances of d e a t h . I n s t e a d of seven
w e h a v e five a g e s of m a n , c o r r e s p o n d i n g t o t h e p e r i o d s
of infancy, of childhood, of y o u t h , of m a t u r i t y o r m i d d l e
a g e , a n d of senility o r old age. I n t h e case of e a c h of
t h e s e p e r i o d s w e s e e a p e r f e c t l y r e g u l a r chance d i s t r i -
b u t i o n , c e n t e r i n g a t a g i v e n age, a n d t a i l i n g off on e i t h e r
side a c c o r d i n g t o a p e r f e c t l y c l e a r m a t h e m a t i c a l l a w . . .
" A r t i s t i c a l l y , w e n o l o n g e r t h i n k of D e a t h as s t r i k i n g
c h a o t i c a l l y ; w e r e g a r d h i s a i m a s p e r f e c t l y r e g u l a r in
t h e m a s s , if u n p r e d i c t a b l e in the i n d i v i d u a l i n s t a n c e . I t
is n o l o n g e r t h e D a n c e of D e a t h w h i c h p i c t u r e s f o r us
D e a t h c a r r y i n g off i n d i s c r i m i n a t e l y t h e old a n d y o u n g ,
t h e r i c h a n d t h e p o o r , t h e toiler a n d t h e idler, t h e babe
a n d i t s g r a n d s i r e . W e see s o m e t h i n g q u i t e different,
the c o h o r t of a t h o u s a n d t i n y m i t e s s t a r t i n g a c r o s s the
B r i d g e of Life, a n d g r o w i n g i n s t a t u r e a s t h e y a d v a n c e ,
7
98 BIOLOGY OF D E A T H
till at the f a r end of the bridge w e see only the gray-
beard and the 'lean and slippered pantaloon.' A s they
p a s s along the causeway the throng 1 is more and more
thinned; five D e a t h s are posted at different stages of the
route longside the bridge, and with different skewness
of aim and different weapons of precision they fire at
the human t a r g e t till none remains to reach the end of the
causewaythe limit of l i f e . "
This whoje, somewhat fanciful, conception of Pear-
son's needs a little critical examination. W h a t actually
he has done is to g e t a good empirical fit of the d% line
by the u s e of equations involving all told some 17 con-
stants. B e c a u s e the combined curve fits well, and funda-
mentally for no other reason, he implicitly concludes
that the fact that the fit is got by the use of five compo-
nents means biologically that the dx line i s a compound
curve, and indicates a five-fold biological heterogeneity in
the material. B u t i t is a very hazardous proceeding to
draw biological conclusions of this type from the mere
fact that a theoretical mathematical function or functions
fits well a series of observational data. I fully discussed
this point several y e a r s ago and pointed out:
" T h e Mnd of evidence under discussion can at best
have but inferential significance; i t can never be of de-
monstrative worth. I t is based on a process of reasoning
which assumes a fundamental or necessary relationship
to exist between two sets of phenomena because the same
curve describes the quantitative relations of both sets.
A little consideration indicates that this method of rea-
soning certainly cannot be of general application, even
though w e assume i t t o be correct i n particular cases.
The difficulty arises from the fact that the mathematical
functions commonly used with adequate results in physi-
 T H E C H A N C E S OF D E A T H 99
cal, chemical, biological, and mathematical investigations
are comparatively f e w in number. T h e literature) of
science shows nothing clearer than that the same type
of curve frequently serves to describe w i t h complete
accuracy the quantitative relations of widely different
natural phenomena. A s a consequence, a n y proposition
to conclude that two sets of phenomena are causally or
in any other w a y fundamentally related solely because
they are described b y the same t y p e of curve i s of a very
doubtful v a l i d i t y . "
Henderson has p u t P e a r s o n ' s five components together
in a single equation, as f o l l o w s :
7.7525
( 1
_71 A 0.2215 (x71.5) [.05524 (a-41.5)p

(.09092 (s 22.5)]'
4- 2.6 e + 8.5 I x2 J .3271 (x 3)

Henderson says regarding this method of P e a r s o n ' s

f o r analyzing the l i f e table: " . . . i t i s difficult t o l a y a
firm foundation for it, because no analysis of the deaths
into natural divisions by causes or otherwise has yet been
made such that the totals in the various groups would
conform to those frequency curves. *' T h e italics in this
quotation are the present writer's for the purpose of em-
phasizing the crucial point of the whole matter.
N o w it is altogether probable that one could get just
a s good a fit to the observed dx line a s is obtained by
P e a r s o n ' s five components by u s i n g a 17 constant equa-
tion of the t y p e
 100 BIOLOGY OF D E A T H
a n d in t h a t e v e n t one w o u l d be q u i t e a s fully justific
( o r r e a l l y unjustified) in c o n c l u d i n g t h a t t h e <lx line wj
a h o m o g e n e o u s c u r v e a s P e a r s o n in in c o n c l u d i n g froi
his five-component fit t h a t it is c o m p o u n d . I n d e e d W i t
stein's formula involving but four constants
n n
(M - J) i (mi)
& a j ^ a

gives a s u b s t a n t i a l l y good fit o v e r t h e whole* r a n g e of lif

I t is, of c o u r s e , a p p a r e n t t h a t t h e f o r m u l a an hero give
is in t e r m s of a n o t h e r function, qx, of t h e life table, rath*
t h a n the dx which we h a v e h i t h e r t o been d i s c u s s i n g . Bi
no difference in in fact involved, q* v a l u e s m a y be* imm
diat<?]y c o n v e r t e d into th v a l u e s by a s i m p l e a r i t h m e t
cal t r a n s f o r m a t i o n .
B u t in n e i t h e r P e a r s o n ' s , W i t t s t o m X n o r a n y othi
case is t h e curve-fitting evidence, by a n d of itwlf, in an
sense a d e m o n s t r a t i o n of t h e biological h o m o g e n e i t y <
h e t e r o g e n e i t y of t h e m a t e r i a l . Of fur g r e a t e r ixnpo:
tance, a n d indeed conclusive significance, in t h e fact, 1
be b r o u g h t out in a l a t e r c h a p t e r , t h a t in m a t e r i a l rxper
mentally known to be biologically homogeneous, a popi
lat ion m a d e u p of full b r o t h e r s a n d BtHtero o u t of a broth*
x s i s t e r m a t i n g a n d k e p t t h r o u g h o u t life* in a u n i f o n
e n v i r o n m e n t identical for all individual**, one grift a a
line in all Us essential feature, s a v e f o r t h e a b s e n c e <
excessive i n f a n t m o r t a l i t y a r i s i n g f r o m p e r f e c t l y c k i
biological c a u s e s , identical with the human d* line. \
h a s l o n g been a p p a r e n t t o t h e t h o u g h t f u l biologist thi
t h e r e w a s n o t t h e s l i g h t e s t biological r e a s o n to suppoi
t h a t t h e p e c u l i a r s i n u o s i t y of t h e h u m a n d* liim owed i
o r i g i n t o a n y f u n d a m e n t a l h e t e r o g e n e i t y in t h e m a t e r i a
o r differentiation in r e s p e c t of t h e forces of m o r t a l i t ;
 T H E CHANCES OF D E A T H 101
Now w e h a v e e x p e r i m e n t a l proof, t o b e discussed i n a
l a t e r c h a p t e r , t h a t w i t h complete h o m o g e n e i t y of the
m a t e r i a l , b o t h genetic a n d e n v i r o n m e n t a l , one g e t s j u s t
t h e s a m e k i n d of dx line as i n n o r m a l h u m a n m a t e r i a l .
W e m u s t t h e n , I think, come to t h e conclusion t h a t bril-
l i a n t a n d p i c t u r e s q u e a s is P e a r s o n ' s conception of the
five D e a t h s , a c t u a l l y t h e r e i s no s l i g h t e s t r e a s o n t o sup-
p o s e t h a t i t r e p r e s e n t s a n y biological r e a l i t y , s a v e in the
one r e s p e c t t h a t h i s c u r v e fitting d e m o n s t r a t e s , a s a n y
o t h e r equally successful would, t h a t d e a t h s do n o t occur
chaotically i n r e s p e c t of age, b u t i n s t e a d i n a r e g u l a r
m a n n e r c a p a b l e of r e p r e s e n t a t i o n b y a m a t h e m a t i c a l
function of a g e .
A n i n t e r e s t i n g a n d s u g g e s t i v e a n a l y s i s of t h e dx line,
r e s t i n g u p o n a s o u n d e r biological b a s i s t h a n P e a r s o n ' s ,
h a s lately b e e n g i v e n b y A r n e F i s h e r . H e b r e a k s the
c u r v e u p i n t o 8 o r 9 components, b a s e d u p o n t h e c o m p a r -
atively s t a b l e v a l u e s of t h e d e a t h r a t i o s f o r different
g r o u p s of d i s e a s e s c h a r a c t e r i s t i c of different a g e s . T h e
r e s u l t i n g t o t a l c u r v e fits t h e f a c t s f r o m a g e 10 on, v e r y
well, a n d m a k e s p o s s i b l e t h e calculation of a complete
life t a b l e f r o m a k n o w l e d g e of d e a t h s only.
 CHAPTER IV

THE CAUSES OF DEATH

IT han beenfluggetc?<lin earlier chapters that natura
death of the mctazoan body may come about fundamen
tally because of the differentiation and consequent mu
tual dependence of structure and function of tha
body. It is a complex aggregate of cells and tissues
all mutually dependent upon each other and in i
delicate state of adjustment and balance. If one orgai
for any accidental reason, whether internal or external
fails to function normally it upsets this delicate balance
and if normal functioning of the part is not promptl;
restored, death of the whole organism eventually results
Furthermore, it is apparent that diath does not strike i)
a haphazard or random manner, but instead in a moa
orderly way. There are certain periods of lifenotabl;
youthwhere only an insignificant fraction of those ex
posed to risk ever die. At other ages, as, for example
extreme old age and early infancy, death strikes wit]
appalling precision and frequency. Further we recal
with Seneca that nascimuB uno mod multin mor
Truly there are many ways of dying. The fnet m obviou
enough. But what is the biological meaning of this mul
tiplicity of pathways to the rivar Styx! There in \m
one pathway into the world. Why so many to go out
To the consideration of some phases of this problen
attention is directed In this chapter.
By international agreement among statisticians tin
causes of human mortality are, for ntatistical purposes
 T H E CAUSES OF D E A T H 103
rather rigidly defined and separated into something over
180 distinct units. I t should be clearly understood that
this convention is distinctly and essentially statistical in
its nature. I n recording the statistics of death the regis-
trar is confronted with the absolute necessity of putting
every demise into some category or other in respect of
its causation. H o w e v e r complex biologically m a y have
been the train of events leading up to a particular end,
the statistician m u s t record the t e r m i n a l " cause of death "
as some particular thing. The International Classifica-
tion of the Causes of Death is a code which is the result
of m a n y y e a r s 7 experience and thought. Great as are
its defects in certain particulars, it nevertheless has cer-
tain marked advantages, the most conspicuous of which
is that by its use the vital statistics of different countries
of the world are p u t upon a uniform basis.
The several separate causes of death are grouped in
the International Classification into fourteen general
classes. These a r e :

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.

P e r h a p s the m o s t outstanding feature which strikes

one about the International List i s that i t i s not primarily
104 BIOLOGY OF DEATH
a biological clasHification. Tt firBt # r o u p , for c x a m p l
called " G e n e r a l DiHcasea/ 7 which canned in 1916 in tl
Her^Btralion A r e a of t h e U n i t e d S l a t e s a p p r o x i m a t e ]
o n e - f o u r t h of all t h e d e a t h s , in a xnonf c u r i o u s biologic;
a n d clinical m e l a n g e . I t includes nuch d i v e r g e e n t i t i e s t
moasleH a n d m a l a r i a , tetamiB a n d tubereulomn, cancc
and gonococcuH infection, alcoholinm a n d g o i t e r , an
m a n y o t h e r unlike cauncH of d e a t h . F o r t h e purponen <
the fttnt-JNtical r e g i s t r a r it p e r h a p s h a s useful p o i n t s t
makc> thin " ( k m e r a l DincaHen" g r o u p i n g , hut it elearl
c o r r e s p o n d s to n o t h i n g n a t u r a l in t h e biological \vorl<
A g a i n in micli p a r t s of t h e neheme a s d o have* noun
biological f o u n d a t i o n t h e basin in different in difFerer
r u b r i c s . Home* h a v e an o r g a n o l o g i e a l bsuuH, while other
have a causationaL
F o r p u r p o s e s of biological a n a l y s i s , I developed som
t i m e a g o a n e n t i r e l y different classification of the minm
of d e a t h , on w h a t a p p e a r s to be a r e a s o n a b l y consisted
barns.* T h e u n d e r l y i n g i d e a of t h i s n e w classiiiciitio
w a s t o g r o u p all c a u s e s of d e a t h u n d e r t h e h e a d s of th
s e v e r a l o r g a n BystemB of t h e body, t h e functional breali
d o w n of which is t h e i m m e d i a t e o r p r e d o m i n a n t cansit n
t h e cessation of life. AH e x c e p t a few of t h e statistical!;
recognized caunes of d e a t h in t h e I n t e r n a t i o n a l C'btssifl
cation can he a s s i g n e d p l a c e s in Hursh a biologicall;
#
It should 1MS ckmrly timtarnttiodi that 1 srn t ndvamlinit a mr
i>f th mimtm &i d^iith fur nUitintimt ttm I nhr*n!4 ttppm
any attempt in mi\mlltuiM & m*w <*\nmlfimtltm lmUu> m
othrr) for thf intvrimlUnml Lint now In UHC. Uniformity in Htmtiiilei
in i*MMi*nt$ml i<> unitbit*, pruvi'tnil vital ^fiiHwfirw, Bttrh tin
Imn mm hft*omn wtll f*fwbH*h*d throtigh th Intertmiiiinttl Ciarnt
ikation. It would Im mont udi?iiral!ii to mukf any rjutiriU r\mng*>* in ti
ClaMNiiicfttian now, I Jmvii madi* rmrtmng^mmt of thi* nttiKiw of distil, k
tin* pttrpcmni ot a Hpwifk liiohigitmi pr<iti!c*m, IMI n<i othi-r. I am itc
"proponing a mw dmulflmUtm of vital nUlintirn99 fc>r vf&cM or any othi
UHC exempt the one to whirh it in htm put.
 T H E CAUSES O F D E A T H 105
g r o u p e d list. I t h a s a s o u n d logical f o u n d a t i o n i n t h e
f a c t t h a t , biologically considered, d e a t h r e s u l t s because
some o r g a n s y s t e m , o r g r o u p of o r g a n s y s t e m s , fails to
continue i t s functions.
T h e h e a d i n g s finally decided u p o n i n t h e n e w classi-
fication w e r e a s f o l l o w s :
I. . Circulatory system, blood and blood-forming organs.
II. Respiratory system.
III. Primary and secondary sex organs.
IV. Kidneys and related excretory organs.
V. Skeletal and muscular systems.
VL Alimentary tract and associated organs concerned in metabolism.
VII. Nervous system and sense organs.
VIII. Skin.
IX. Endocrinal system.
X. All other causes of death.

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.*

GENEBAL, BESULTS OF BIOLOGICALLY CLASSIFIED DEATH BATES

Here I should like to present first some general statis-
tical results of this classification. The data which we
shall first discuss are in the form of death rates, from
various causes, per hundred thousand living at all
ages, arranged by organ systems primarily concerned
in death from specified diseases. The statistics came
from three widely separated localities and times, viz.,
(a) from the Eegistration A r e a of the United S t a t e s ;
(b) from England and W a l e s ; and (c) from the City of
Sao Paulo, Brazil,
The summarized results are shown in Table 7, and in
graphic form in F i g u r e 26.
The rates are arranged i n descending order of magni-
tude for the United States Eegistration Area, with the
exception of those of group X , all other causes of death.
W e note in passing that this biologically unclassifiable
group includes roughly 10 to 15 per cent of the total
mortality. I t m a y be well to digress a moment to con-
sider why these deaths cannot be put into our general
scheme. Table 8 exhibits the rates included in class X .
This residue comprises i n general three categories
(a) accidental and homicidal d e a t h s ; (&) senility; and
C/. particularly Pearl, R. " On the embryological basis of human
mortality." (Proc. Natl. Acad. Sci. VoL 5, pp. 593-598, 1919) and "Cer-
tain evolutionary aspects of human mortality rates." (Amer. Natl. VoL
LIV. pp. 5-44, 1920). The following section as well as Chapter V are
largely based upon the second of the two papers.
 T H E CAUSES OF D E A T H 107
(c) d e a t h s f r o m a v a r i e t y of causes which a r e s t a t i s t i -
cally l u m p e d t o g e t h e r a n d c a n n o t be d i s e n t a n g l e d . Ac-
cidental a n d homicidal d e a t h s find n o place in a biologi-
TABLE 7
Showing the Relative Importance of Different Organ Systems in
Human Mortality
Death Rates per 100,000
Group Registration Area, England Sao
No. Organ System U.S.A. and Paulo
Wales 1917
1906-10 1901-05 1914

II Respiratory system 395.7 460.5 420.2 417.5

VI Alimentary tract and associated
organs 334.9 340.4 274.1 613.8
I Circulatory system, blood 209.8 196.8 208.6 254.8
VII Nervous system and sense organs . 175.6 192.9 151.9 124.3
IV Kidneys and related excretory
organs 107.2 107.4 19.4 83.4
III Primary and secondary sex organs. 88.1 77.4 95.4 103.2
V Skeletal and muscular system 12.6 13.7 18.2 6.8
VIII Skin 10.1 13.3 12.0 7.9
IX Endocrinal system 1.5 1.2 1.9 1.1
Total death rate classifiable on a
biological basis 1,335.5 1,403.6 1,201.7 1,612.8
All other causes of death 171.3 211.8 141.4 109.8

c a l classification of m o r t a l i t y . A m a n organically sound

i n every respect m a y be instantly killed b y being struck
by a railroad train or an automobile. The best possible
case that could be m a d e out for a biological factor in such
deaths would be that contributory carelessness or negli-
gence, which is a factor in some portion of accidental
d e a t h s , b e s p e a k s a s m a l l but definite organic m e n t a l in-
feriority or weakness, and that, therefore, accidental
deaths should be charged against the nervous system.
This, however, is obviously not sound. F o r , in the first
p l a c e , in m a n y accidents t h e r e is n o f a c t o r of c o n t r i b u t o r y
 RESPIRATORY
SYSTEM

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

negligence in fact, a n d , in t h e nccnnd places in t h o n e CUHGH

w h e r e s u d i n e g l i g e n c e c a n f a i r l y be a l l e g e d itB d e g r e e o r
significance i u n d e t e r m i n a b l e a n d in m a n y caH8 s u r e l y
slight
S e n i l i t y an a c a u s e of d e a t h in n o t f u r t h e r duHBifiahic*
TABLK 8
Alt Otkrr Vnmrn

IUitrli.in Arm,
Pmth** V. H. A.
No. taf ft licit lmi
1017
...mow

All extftrnn! <?fttH4*f* (vxwpt fiiitcttit* 87.B mi IMA

187,
188 &
189 29.4 47.H 7.3 36.3
154 Hrntiity , 2tt.C) 41.0 81.5 11.1
45 Cancer tti olhrr firgnrw <ir of n
not 10.1 lfl.fl
152* (Mlwr muwm t rnrly $ii-
fnncy. ZA 2 r> i 3.3
34 of thi 2.1 2,0 i.o 0,2
Other ttirnom (f<*malt! or
1.0 1.5 o.
65 1.0 0.5 i.fl
153 Lark f on VIM on 0
03 0.2 on 0.2
i 171.3 211 H Hl.V 100.8

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 rfprcmc*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 bot 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-
tionthe 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 goodIndeed 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.

SPECIFIC DEATH BATES BIOLOGICALLY CLASSIFIED

U p to this p o i n t in o u r discussions we have been deal-
i n g w i t h c r u d e d e a t h r a t e s , u n c o r r e c t e d for the age a n d
sex d i s t r i b u t i o n s of t h e p o p u l a t i o n s concerned. I t i s ,
of course, a well k n o w n fact t h a t differences i n a g e a n d
 T H E C A U S E S OF D E A T H 113
sex constitution of populations m a y make ccmmderable
differences in crude death rates, in c a s e s where no real
differences in the true force of mortality exist. What
is essential for the further prosecution of the a n a l y s i s of
the causes of d e a t h in to got specific death rates for the
several causes. B y an ago and BOX specific d e a t h rate
is meant the rate g o t by dividing the number of persons,
of particular Hprcifitd age and sex, d y i n g from a particu-
lar cause, by the total number of persons living in the
same population of the same age and . I n other
words, we nttmi to g*t an the divisor of the rate fraction
the number of persons who can be regarded a s truly ex-
posed to rink. T h i s cxpoHttd-to-risk portion of the popu-
lation is never correctly stated in a crude death rate.
F o r example, a parson now 75 y e a r s old cannot be re-
garded a s e x p o s e d to rink of death a t a g e 45, H o w a s
once exposed to that risk but passed it safely. Y e t in a
crude death rate he is counted with those* of a g e 45.
A g e and sex specific death rates have* hitherto bean
available for thi* American people*, in any gumsral or com-
prehensive form* only from thu extensive* m e m o i r by
Dublin* K o p f a n d V a n Buren, based upon the mortality
experience of the Metropolitan Lifa Insurance C o m p a n y
with ita industrial policy holders, In a broad w a y , i t
may be said t h a t the d a t a o n which th# following discus-
sion is baned, derived from the g<m<nd population of
the Registration A r e a , a m essentially in accord with those
of Dublin on a m o r e restricted group. O w i n g to limita-
tions of space, i t i s not possible to present all the detailed
rates here.
With the aid of D r . William I I D a v i s , director of
vital statistics in the C e n s u s Bureau, w h o vnry kindly
provided me with thu n t e a i s a r y unpublished d a t a , i t h a s
8
114 BIOLOGY OF D E A T H
been possible t o calculate the specific d e a t h r a t e s <
of the 189 causes of d e a t h of t h e I n t e r n a t i o n a l L
each sex s e p a r a t e l y , a n d for each a g e i n 5 y e a r |
for the U n i t e d S t a t e s E e g i s t r a t i o n A r e a , exclua
N o r t h Carolina, i n 1910. T h e s e r e s u l t s h a v e b e
t o g e t h e r i n t h e biological scheme of classification a i
be p r e s e n t e d briefly i n t h e f o r m of d i a g r a m s .
T h e s u m m a r y table f r o m which t h e s e curves a i
t e d is given as T a b l e 9.
L e t u s first c o n s i d e r d e a t h s f r o m all c a u s e a
together, in o r d e r t o recall to m i n d t h e g e n e r a l <
a d e a t h r a t e c u r v e . I t will b e n o t e d , a t once, t l
r a t e s a r e plotted along t h e v e r t i c a l axis on w h a t
one a t first as a p e c u l i a r scale. T h e scale is l o g a r .
T h e h o r i z o n t a l lines a r e s p a c e d i n p r o p o r t i o n
l o g a r i t h m s of t h e n u m b e r s at t h e i r left, i n s t e a d of :
p o r t i o n t o the n u m b e r s themselves. T h e a d v a n t ^
this m e t h o d of p l o t t i n g in the p r e s e n t case a r e t w
F i r s t , i t is possible to g e t a m u c h w i d e r r a n g e of
on the d i a g r a m ; a n d second the l o g a r i t h m i c scale p
direct a n d a c c u r a t e estimation of t h e r a t e of c h a n ^
variable. A s t r a i g h t line f o r m i n g an angle with t l
izontal on a l o g a r i t h m i c scale m e a n s t h a t the v ^
is increasing o r d e c r e a s i n g , as t h e case m a y be, at
stant rate of change.
F i g u r e 27 gives the specific d e a t h r a t e s for thu
bined t o t a l of all causes. T h e curve i n general Ix
f o r m of a V, w i t h one limb m u c h extended a n d p u l l e
to the r i g h t . E x a m i n i n g i t m o r e i n detail, we n o t
in t h e first y e a r of life, the specific d e a t h r a t e , o r ,
m a y r o u g h l y call it, t h e force of m o r t a l i t y , b e a m
i e r on female i n f a n t s t h a n on t h e m a l e s . Out of a
s a n d exposed t o risk, 124 m a l e babies die in t h a t
 TABLE 9
Showing the Specific Death Rates, per 1000 Living of the same Age and Sext for each Biological Grou
in the U. S. Registration Area, exclusive of North Carolina, in 1910
Group I Group II Group III Group IV GroupV Group VI GroupVII Group VIII Group IX Group X All Causa
Ages
9 <? 9 9 d1 9 cT 9 d1 9 d1 9 d1 9 d1 9 d1 9 d1 9 i
n Under 1 6.87 5.1330.1524.681.74 43.16 0.93 0.630.400.34 66.3254.25 5.46 4.58 1.01 0.97 0.01 0.005 11.50 9.69 124.38143.43
1-4 1.00 1.00 6.89 6.53 .03 .04 .18 .18 .14 .12 4.04 3.56 1.43 1.25 .04 .04 .0006 .0009 1.34 1.12 15.09 13.84
5-9 .57 .63 1.26 1.33 .006 .006 .10 .10 .11 .11 .56 .55 .45 .41 .006 .008 .0001 .64 .34 3.71 3.49
ft 10-14 .37 .45 .51 .74 .004 .008 .08 .11 .12 .11 .51 .54 .30 .25 .009 .005 .002 .003 .60 .16 2.50 2.39
15-19 .38 .40 1.38 1.52 .008 .24 .12 .21 .10 .08 .79 .70 .36 .30 .02 .01 .004 .018 .97 .22 4.14 3.69
20-24 .37 .43 2.30 2.29 .03 .69 .17 .37 .07 .06 .93 .77 .48 .33 .02 .02 .003 .027 1.59 .24 5.97 6.22
25-29 .44 .52 2.68 2.59 .04 .96 .30 .50 .06 .06 .95 .81 .60 .38 .03 .02 .004 .031 1.67 .25 6.77 6.12
30-34 .67 .68 3.15 2.57 .06 1.12 .42 .62 .07 .06 1.05 .91 .87 .49 .05 .03 .007 .04 1.65 .28 8.00 6.79
35-39 .95 .92 3.59 2.46 .07 1.25 .65 .78 .10 .07 1.27 1.18 1.19 .64 .05 .03 .009 .05 1.86 .38 9.75 7.77
40-44 1.42 1.31 3.88 2.34 .08 1.31 .98 1.00 .11 .08 1.62 1.50 1.55 .85 .09 .04 .01 .07 1.87 .43 11.61 8.93
1 45-49 2.06 1.91 4.15 2.32 .10 1.41 1.48 1.22 .12 .10 2.26 2.12 2.14 1.27 .13 .67 .01 .07 2.09 .53 14.64 11.03
50-54 3.13 2.71 4.57 2.67 .12 1.46 2.14 1.73 .16 .14 3.21 3.00 2.81 2.03 .16 .09 .02 .07 2.19 .73 18.51 14.63
55-59 5.11 4.24 5.46 3.54 .19 1.64 3.28 2.51 .19 .21 4.50 4.46 4.17 2.91 .23 .10 .02 .10 2.52 .90 25.66 20.62
60-64 8.25 6.79 6.59 5.07 .32 1.75 4.98 3.40 .24 .22 6.26 5.81 5.98 4.53 .34 .18 .02 .08 3.10 1.45 36.07 29.30
65-69 13.4710.71 8.12 7.75 .73 2.00 7.05 4.94 .38 .36 8.11 8.24 8.99 7.38 .55 .28 .02 .10 4.04 2.53 51.44 44.29
70-74 21.2117.27 10.96 11.981.33 2.08 10.41 7.24 .48 .57 10.2710.7113.1911.21 .80 .50 .02 .09 6.44 5.17 75.10 66.82
75-79 32.7325.4115.94 18.212.24 2.53 14.87 9.68 .67 .8513.0913.9120.0017.971.17 .73 .02 .09 11.51 11.51112.24100.90
80-84 46.4239.2424 2628.533.71 2.56 19.7512.51 .79 1.25 17.26 17.6127.1825.911.82 1.38 .01 .06 26.94 26.84168.14155.87
85-89 61.7951.3334.6640.704.38 2.83 26.7615.781.07 .8720.4721.4733.7132.152.91 1.63 .02 52.16 55.95237.91222.74
90-94 69.0058.8349.6558.134.45 2.89 26.2116.021.68 1.3125.6024.3433.7837.913.61 2.01 .18 99.05108.12313.02309.73
95-99 79.2173.2670.0160.909.19 1.77 26.1723.392.83 .4428.2922.5131.1237.514.24 2.65 159.03146.51410.18368.93
100 and
over 81.0868.8361.7872.87 6.07 38.6110.12 15.4418.2238.6138.46 6.07 258.69251.01494.21471.66
116 BIOLOGY OF D E A T H
a n d 143 female. T h i s is t h e only y e a r of life in which t h e
t o t a l force of m o r t a l i t y is h e a v i e r a m o n g females t h a n
males. F r o m t h a t t i m e on t o t h e end of the s p a n of life,
IJOOO\

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

the male curve. After the heavy mortality ;of early

infancy, the curve drops in almost a straight line to the

 TIIK CAU8KS O F D E A T H 117
age p e r i o d of 10-15, w h e r e it reachtm itn lowunt pointy a n d
only a p p r o x i m a t e l y 2-1,2 pen-urns o u t of a t h o u n a n d ex-
posed to rink die. Th<* specific m o r t a l i t y c u r v e them be-
gins to ri*u% mid contiim*n to da no a t a u a p p r o x i m a t e l y
constant a n d r a p i d r a t e for t e a y e a r n t h a t in t o t h e
age? p e r i o d 20-25. F r o m then on to the* age* p e r i o d 50-55
it rmm a t a s l o w e r b u t comttant r a t e . Thin in the p e r i o d
of middle lif\ a n d h*r<* the* female mrw dropH f a r t h e r
below the m a l e c u r v e t h a n a t any o t h e r place in t h e wpan
of life. A f t e r Urn age* p e r i o d 50-55 w i t h t h e o n - c o m i n g of
old ago, both malt? a n d femali* ctirvcm begin a g a i n t o r i s e
more r a p i d l y . T h e y continue) this r i s e , a t a {practically
eoriBtnnt r a t e of snrrwuti*, to t h e cud of lifc% which IB h e r e
taken m fitlliiig in t h e a g e p e r i o d 9 0 1 0 0 . In thin lant
class t h e r a t e hat* bt&omo v e r y high. O u t of 1,000 p e r -
sons living n t tin* #*gi*H f 95 a n d 100 f a n d i h m t f o r t * ex-
poses! t o risk of d o n t h w i t h i n t h a t p e r i o d , 41)4 maleH a n d
473 females d$c*f t n k t n g nn n v e r a g i ! f o r tho whoht ftv-
y c a r p e r i o d . Of r o u n d ? , boforo the <u>mphttion of t h e
period, priicticaliy all of the* t h o u s a n d will h a v e p a s s e d
away.
T h e i m p o r t a n t t h i n g * t o n o t e a b o u t thin curvet a r e
thc*e: F i r t f thti htghoni npimifm fortmn of m o r t a l i t y oe-
eur a t t h e t?xtrc*mtt emln of Hfo, a n d art* highcir a t t h e
final end thatt a t thti iMigiiining. lit t h o mtcond pluce 9
t h e r e is a s h a r p a n d utogtly d r o p , i n a l m o s t a n t r a i g l t t
line, f r o m t h e h i g h upteifie foreii of m o r t a l i t y in i n f a n c y
to t h e low p o i n t a t About t h e t i m e of p t t b o r t y . From
then on to t h u u n d of tho u p a n of Hfe f t h e f o r m of m o r t a l -
ity become* g r e a t e r e v e r y y e a r a t a n e a r l y e o n n t a n t r a t e
of inerfjaat, w i t h o n l y mieh s l i g h t d e v i a t i o n s f r o m t h i s
constancy of r a t e a s hmvt a l r e a d y \num p o i n t e d o u t
T u r n i n g n e x t t o t h a m o r t a l i t y of o u r I r s t biological
118 BIOLOGY OF D E A T H
g r o u p n a m e l y d e a t h s caused by b r e a k d o w n of t h e cir-
c u l a t o r y system, blood a n d b l o o d - f o r m i n g o r g a n s w e
n o t e in F i g u r e 28 a m a r k e d difference in t h e f o r m of t h e
100

CIRCULATORY SYSTEM, BL.OQD AND

BLOOD-FORMING ORGANS

aoi
AOL
FIQ. 28.Diagram showing tho ipecific death rat t r.rb u^ Umu brmkdumu i>i lh#
circulatory yttm, 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 *ux 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 itliHrlntely 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 ^ytc*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

The lowest point of the respiratory curve falls in the

a g e group 10-15. Between the a g e s 25-70 there is a v e r y
striking difference in the two s e x e s in respect o f specific
 T H K CAUSES O F D E A T H 121
m o r t a l i t y f r o m b r e a k d o w n of the* r e s p i r a t o r y s y n t e m .
The male crvi risen in n e a r l y a Htraight line, while t h e
female c u r v e lien f a r below if, ancl a c t u a l l y shown a p o i n t
of inflection a t a b o u t #* 45, b e c o m i n g f o r a n h o r t p e r i o d
convex t o t h e bane. T h e e x p l a n a t i o n f o r t h e g r e a t nep-
aration of the* t w o c u r v e s in thin p e r i o d in p r o b a b l y fun-
damentally o c c u p a t i o n a l . F r o m t h e n a t u r e of t h e i r
activity m a l e s , d u r i n g thin p e r i o d of life, a n * p r o b a b l y
subject in a grrenter rink of b r e a k d o w n of t h e r e s p i r a t o r y
system t h a n a r e t h e m o r e p r o t e c t e d femalo liven. F r o m
age 70 on, b o t h a t r v e n aHtwnd with i n c r e a s e d r a p i d i t y t
the female c u r v n r i s i n g iilmve t h e m a l a , p r e s u m a b l y in
compensation f o r t h e m a r k e d d i p which it exhibitB in m i d -
dle life. I t is, of c o u r s e , well known t h a t r e s p i r a t o r y
m o r t a l i t y bear** h e a v i l y u p o n t h e a g e d .
T h e n e x t ffrotip which %w nltal! coimider ban t o d o
with d e a t h s f r o m breitkthnvn of t h e p r i m a r y a n d necond-
iiry aex orKarw, Thin ciiiiHe ^ r o t t p furntHhen a n ex-
tremely interf*Htinic p n i r of ciirvim nhown in F i g u r e 30.
Before dincitHimiK in detiiil thrnr form f a w o r d of e x p l a n -
ation at* t o t h e i r m a k e u p should bo given. Thin m a y
beat \m dmw b y e x h i b i t i n g rind c]inctiHitig f o r a m o m e n t
the etmmm of d e a t h w h i c h arc* included in thin g r o u p .
Table 10 nhow* thu d u t a .
In thin r i i b r i a a m itielitded ' ' P r e m a t u r e b i r t h 1 * a n d
u
I n j u r i e s a t b i r t h / ' T h u q u e s t i o n a t onco liriHiiB, w h y
should them* t w o itt*mitf ftlPremature birth 1 * a n d " I n j u r i e s
a t b i r t h 1 f be i n d u d e d w i t h tlui p r i m a r y a n d 8<&midary sex
o r g a n s inen i t in o b v i o t i i e n o u g h t h a t t h e i n f a n t i w h o s e
d e a t h s a r e reccirded u n d e r t h m e hciadu in thu v a s t m a j o r -
ity of e a s t s , if n u t nil, h a v e n o t h i n g w h a t e v e r t h e m a t t e r
with e i t h e r their p r i m a r y o r icicionclmry o r g a n s ! T h e
a n s w e r mf in g e n e r a l t e r m n , t h a t o n my p r o p e r biological
122 BIOLOGY OF D E A T H
basis, d e a t h coming u n d e r e i t h e r of t h e s e two categories is
n o t p r o p e r l y c h a r g e a b l e , organically, a g a i n s t t h e i n f a n t a t
TABLE 10
Primary and secondary sex organs
Registration Area, England
" Cause of Death " as per International U. S. A. and Sao
No. Classification Wales Paulo
1914 1917
1906-10 1901-05
151* Premature birth 35.7 30.8 46.9 66.8
42 Cancer of the female genital organs 10.8 10.0 12.9 6.5
137 Puerperal septicemia 6.8 6.3 3.7 6.5
152* Injuries at birth 6.6 5.0 2.8 2.1
43 Cancer of the breast 6.5 5.6 10.4 1.5
37 Syphilis 5.4 4.1 5.8 15.0
126 Diseases of the prostate 3.4 2.6 4.2 0.7
132 Salpingitis and other diseases of
9 genital organs 2.2 2.1 0.5 0.2
129 Uterine tumor (non-cancerous) . .. 1.8 1.8 0.8 0
134 Accidents of pregnancy 1.7 1.1 0.2
130 Other diseases of the uterus 1.6 0.4 0.4
136 Other accidents of labor 1.3 0.9 1.1 0.7
140 Following childbirth 1.1 1.5 0.1
131 Cysts and other tumors of ovary.. 1.0 1.3 0.8 0.2
135 Puerperal hemorrhage 1.0 1.0 1.3 1.7
125 Diseases of the urethra, urinary
abscesses, etc 0.4 0.4 1.2 0.7
Gonococcus infection 0.3 0.1 0.2 0
128 Uterine hemorrhage (non-puerpe-
ral) 0.2 0.3 0 0
127 Non-venereal diseases of d" genital
organs. 0.1 0.1 0.2 0
133 Non-puerperal diseases of breast
(except cancer) 0.1 0.1 0.1 0
139 Puerperal phlegmasia, etc 0.1 0.9 0
Totals 88.1 77.4 95.4 103.2
* In part.

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 ,

n rliiithn urt* a r g i n t i r a i l y rtiitrg^itltk 1 t o b r e a k d o w n o r

failure? to fiinittjott p r o p e r l y of tin* rc|ircKJiieti^ o r g a n i
of thct mothor* Tlwm* i k ^ t b n , thcMrt*foref g o Into this
126 BIOLOGY O F D E A T H
female g r o u p . B y t h e fifth y e a r of life, t h e specific r a t e s
of m o r t a l i t y c h a r g e a b l e to r e p r o d u c t i v e organs* have
d r o p p e d in b o t h sexes p r a c t i c a l l y t o z e r o , a m o u n t i n g to
less t h a n 0.01 p e r t h o u s a n d e x p o s e d t o rink. A t about
the time of p u b e r t y t h e female c u r v e begin** t o r i s e and
goes u p v e r y steeply. B y a g e 30 it h a s r e a c h e d a value
of 1 p e r t h o u s a n d exposed to risk. F r o m t h a t p o i n t the
force of this specific m o r t a l i t y r i s e s slowly, b u t 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 , to e x t r e m e old ago. T h e male
curve is in s t r i k i n g c o n t r a s t t o t h e female. F r o m a b o u t
age 20 i t r i s e s steadily, a t a n a l m o s t c o n s t a n t r a t e of
increase, b u t a m u c h slower one t h a n t h e female, until
the end of t h e life s p a n , Tt c r o s s e s the female* c u r v e i n -
d i c a t i n g a h i g h e r specific r a t e of m o r t a l i t y f r o m break-
down of t h e r e p r o d u c t i v e o r g a n s in men t h a n in women
f o r t h e first t i m e a t a b o u t a g e 78. Thin in, of c o u r s e , the
time of life w h e n d i s t u r b e d f u n c t i o n i n g of t h e p r o s t a t e
gland in t h e m a l e b e g i n s to t a k e a r e l a t i v e l y h e a v y toll.
F i g u r e 31 shows specific r a t e s of m o r t a l i t y from
b r e a k d o w n of t h e k i d n e y s a n d r e l a t e d e x c r e t o r y o r g a n s .
D e a t h f r o m t h e s e c a u s e s is r e l a t i v e l y i n f r e q u e n t in in-
fancy a n d e a r l y childhood. T h e low p o i n t is r e a c h e d ,
a s in so m a n y of t h e o t h e r c a s e s , a t a b o u t t h e t i m e of
p u b e r t y . F r o m t h e n on p r a c t i c a l l y t o t h e m& of t h e
s p a n of life t h e specific force of m o r t a l i t y f r o m e x c r e t o r y
failure increases a t a n almost constant rate. During
t h e r e p r o d u c t i v e p e r i o d , f r o m a b o u t 15 t o 45 y e a r s of a g e ,
specific r a t e s of m o r t a l i t y f r o m t h e s e c a u s e s a r e h i g h e r
in t h e f e m a l e t h a n in t h e m a l e . A f t e r t h a t p o i n t thct m a l e
c u r v e i s h i g h e r . T h e r e l a t i v e l y h e a v y specific m o r t a l i t y
of the female in e a r l y life i s u n d o u b t e d l y d u e t o t h e h e a v y
s t r a i n p u t u p o n h e r e x c r e t o r y o r g a n s by c h i l d - b e a r i n g .
T h e specific f o r c e of 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 CAUSES OP D E A T H 127
the skeletal and muKcular s y s t e m s , shown in F i g u r o 32,
presents an i n t e r e s t i n g pair of curves. T h r o u g h o u t the
span of life there in practically no difference between

loorz

the femail* ami malv in th* ittt*hU*nmt of thin mirtality t

the eiirvc**! wimiitiK w f l ut about eft<?h other. T h e
striking chararttrtntirMi of the <fttnfi* iiri: flrnt, that the
ipcciflc fcirwn f mortalify art* nl^ohttoly low for
128 BIOLOGY OF D E A T H
organ s y s t e m s ; and second, that the minimum point is
reached not, as in most of the other cases, around the time
of puberty, but at a much later periodnamely in the
100

SKELETAL AND MUSCUl-AR SYSTEMS

K> k B J 3d JB Jb S 36 h 60 k
Of.
Fio. 32.Diagram ahowingpcifio<Ifmth r&am t#n@H *m (mm brmk*tvmn of thm
ttd muaoular uyutvam (Group V),

late twenties. The whole curve shows a very gradual

change in the rates.

The next diagram, Figure 33, shows one of the most

 T H E CAUSES OF D E A T H 120
significant o r g a n g r o u p s i n t h e f o r c e of i t s specific m o r -
tality. B r e a k d o w n a n d f a i l u r e t o f u n c t i o n p r o p e r l y of
the p r i m a r y o r g a n s of m e t a b o l i s m t h e o r g a n s w h i c h

too

TPAC7 AND ASSOClATCD

ORGANS COKCtPNO) W M&TABQUSM

ABL
WIQ. 33.Diagram * how Jog th r*dfk rat<* of < frum breakdown
alimentary tract and aiwociated orgs

t r a n s f o r m the fuel of the h u m a n m a c h i n e into vital e n e r g y

occur with relatively heavy frequency at all periods
of life. These curves are a m o n g the few which show aa
130 BIOLOGY OF D E A T H
absolutely higher specific force of mortality in infancy
than i n extreme old age. There is practically no signif-
icant difference between the male and female curve at

NERVOUS SYSTEM AND SENSE ORGANS

OOt _L
io is io JO 4$ so h do
ABC
Fio. 34.Diagram showing the upcciflc death ttXm Rt ^rh *m from tr#*ltbwit o( th
nervous tystom nod enc orgmwt (Group VIIh

any portion of life. During early adult life the female

curve lies l>elow t h e m a l e , but by only a small amount.

Out of every thousand infants under one, about sixty

 T H E C A U S E S OF D E A T H 131
die in the first year of life from breakdown of the ali-
mentary tract and its associated organs. A f t e r the low
point, which falls i n the relatively early period of 7 to 12
years of age, there is a rapid rise f o r about ten y e a r s
in the specific rates of mortality, followed by a slowing
off in the rate of increase for the next ten or fifteen y e a r s ,
after which point the curve ascends at a practically uni-
form rate until the end of the span of life.
Figure 34 shows the trend of the specific mortality
from breakdown of the nervous system and sense organs.
This organ group, on the whole, functions v e r y well, giv-
i n g a relatively low rate of mortality until towards the
end of middle life. Then the specific rates get f a i r l y
large. The low point i n this curve is, as in most of the
others, at about the time of puberty. F r o m then on to
the end of the life span the specific rates increase at a
practically uniform rate. The female curve everywhere
lies below the male curve except at the extreme upper
end of the life span. Before that time, and particularly
between the ages of 20 and 50, the business of living
evidently either imposes n o such heavy demand on the
nervous system of the female as i t does on that of the
male, or else the nervous system of the female is organi-
cally sounder than that of the male. The former sug-
gestion seems the more probable.
That breakdown and failure to function properly, of
the skin as an organ system, i s a relatively insignificant
factor in human mortality, is demonstrated by F i g u r e 35.
F r o m a specific death rate of about 1 per thousand i n
the first y e a r of life it drops abruptly, practically to zero,
i n early childhood. A t about the time of puberty it be-
gins to rise again, and ascends at a steady rate during
all the remainder of life. The final high point reached
132 BIOLOGY OF D E A T H
i s absolutely low, however, amounting to a specific death
rate among those exposed to risk of only a little more than
4 per thousand at the extreme end of life. The female

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

FZG. 3 6 . D i a g r a m s h o w i n g t h e specific d e a t h rates a t e a c h a g e f r o m b r e a k d o w n of the

endocrinal system (Group I X ) .

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

ALL OTHER CAUSES OF DEATH

100

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 1014 Respiratory 33.3
43.6 Respiratory 1519 Respiratory 43.8
52.6 Respiratory 2024 Respiratory 46.0
49.7 Respiratory 2529 Respiratory 44.2
45.6 Respiratory 3034 Respiratory 39.5
39.9 Respiratory 3539 Respiratory 33.2
33.3 Respiratory- 4044 Respiratory 27.5
28.0 Respiratory 4549 Respiratory 22.1
23.6 Respiratory 5054 Alimentary tract 21.6
25.0 Circulatory 5559 Alimentary tract 22.6
28.4 Circulatory 6064 Circulatory 24.4
30.9 Circulatory 6569 Circulatory 25.6
32.5 Circulatory 7074 Circulatory 28.0
32.9 Circulatory 7579 Circulatory 28.4
33.3 Circulatory 8084 Circulatory 30.4
8589 Circulatory 30.8

If o u r l u n g s w e r e a s organically good relatively a s

our h e a r t s , h a v i n g r e g a r d i n each case f o r t h e w o r k t h e
o r g a n is called u p o n t o do a n d t h e conditions u n d e r which
it m u s t do it, we should live a considerable n u m b e r of
y e a r s l o n g e r on t h e a v e r a g e t h a n we do now. One c a n n o t
but feel t h a t t h e w o r k i n g out of a r a t i o n a l and scientifi-
cally g r o u n d e d s y s t e m of p e r s o n a l h y g i e n e of t h e r e s p i r -
 T H E CAUSES O F D E A T H 137
a t o r y o r g a n s , on t h e b r o a d e s t b a s i s , t o i n c l u d e all such,
m a t t e r s as v e n t i l a t i o n of b u i l d i n g s , etc., a n d t h e p u t t i n g
of such a p e r s o n a l h y g i e n e i n t o g e n e r a l u s e t h r o u g h
education, would p a y a b o u t a s l a r g e d i v i d e n d s a s could
be hoped f o r f r o m a n y i n v e s t m e n t i n p u b l i c h e a l t h secu-
rities. I a m a w a r e t h a t m u c h h a s a l r e a d y b e e n d o n e i n
this direction, b u t i n o r d e r to r e a p a n y s u c h d i v i d e n d s
a s I a m t h i n k i n g of, a v a s t a m o u n t m u s t be a d d e d t o o u r
p r e s e n t knowledge of t h e physiology, p a t h o l o g y , e p i d e m i -
ology, a n d e v e r y o t h e r a s p e c t of t h e f u n c t i o n s a n d s t r u c -
t u r e s of r e s p i r a t i o n .
 CHAPTER V

EMBRYOLOGY AND HUMAN MORTALITY

Iir the preceding chapter attention wa confined
strictly to the organological incidence of death. It is
possible to, push the matter of human mortality still
farther back. In the embryological development of the
vertebrate body, there are laid down at an early stage,
in fact immediately following the process of gnat rotation,
three morphologically definite primitive tisniic! elements,
called respectively the ectoderm, the menoderm and the
endoderm. These are termed the germ-lay erg, and em-
bryological science has, for a great many forms, auceeeded
in a broad way in tracing back to the primitive germ
layer from which it originally started Ha development,
substantially every one of the adult organs and organ
systems of the body. Itjmakes no difference to the validity
or significance of thecliseussion which we are about to enter
upon, in what degree of esteem or contempt in biological
philosophy the germ layer theory or doctrine, which oc-
cupied so large a place in morphological speculation 50
years ago, may be held. We are here concerned only with
the well-established broad descriptive fact, that in general
all adult organ systems may he traced back over the path
of their embryological development to the germ layer, or
combination of germ layers, from which they origin-
ally started.
Having arranged, so far aa possible, all causes of death
on an organological basis, it occurred to me to go one
 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 139
step f u r t h e r back a n d combine t h e m u n d e r the h e a d i n g s
of the p r i m a r y g e r m l a y e r s f r o m which t h e s e v e r a l o r g a n s
developed embryologically. T o do t h i s w a s a t a s k of
considerable difficulty. I t r a i s e d i n t r i c a t e , a n d i n some
TABLE 12
Showing the relative influence of the primary germ layers in human mortality
(Items 64 and 65 charged to ectoderm)
Death rate per 100,000 due to functional breakdown
of organs embryologically developing from
Locality
Ecto- Per Meso- Per Endo- Per
derm oent. derm cent. derm cent.
United States Registration
Area, 1906-10 191.1 14.3 425.2 31.8 719.6 53.9
United States Registration
Area, 1901-05 210.6 15.0 407.1 29.0 786.2 56.0
England and Wales, 1914... 177.1 14.4 374.0 30.3 681.5 55.3
Sao Paulo, 1917 134.9 8.4 468.0 29.0 1009.9 62.6

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

VS. REGISTRATION AREA 1306'JO

ENGLAND AND WALES

m m
m
M M . i s ?

SAO PAULO J9J7

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*sint 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

Fci.i 39.Bhow nai speccfi deah

t rate* In m & k
* ^wrn cig to b t g^
othriniggin.likTen
is10patim
rtesofatshemd eyathderaathtes aasccoeuith
a
n ntesrfomresosd
ectoderm at this peroid of life* Prom about age 12
 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 145
the case of o r g a n s of e c t o d e r m i c o r i g i n , a n d f r o m a b o u t
a g e 22 on in canes of m e s o d e r m i c origin, t h e d e a t h r a t e
c u r v e s rine 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 e x t r e m e old
146 BIOLOGY OF D E A T H
be claimed f r o m t h e details r e v i e w e d in t h i s a n d t h e p r e -
ceding c h a p t e r ? I h o p e t h a t t h e s e f a c t s will h a v e s e r v e d
in some m e a s u r e to c o m p l e t e a n d r o u n d out in c l e a r e i r

, ._
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 viw, 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

W E have seen that in the case of man, where alone

quantitative data are available, the breakdown of partic-
ular organ systems, and consequent death of the whole,
occurs in a highly orderly manner in respect of time or
age. Each organ system h a s a characteristic time curve
for its breakdown, differing from the curve of any other
system. The problem which now confronts us is to find
out what lies back of these characteristic time curves and
determines their form. I n v i e w of the biological facts
about death which w e have learned, what determines that
John Smith shall die at 58, while H e n r y Jones lives to
the obviously more respectable a g e of 85? W e have
seen that there is every reason to believe that all the
essential cells of both their bodies are inherently capable
under proper conditions of living indefinitely. I t fur-
ther appears probable that i t is the differentiated and
specialized structure of their bodies which prevents the
realization of these favorable conditions. B u t all this
helps u s no,t at all to understand w h y in fact one lives
nearly 30 years longer than the other.
It may help to visualize this problem of the determina-
tion of longevity t o consider an illustrative analogy.
Men behave i n respect of their duration of life not unlike
a lot of eight-day clocks cared for by an unsystematic
person, who does not wind them all to an equal degree
and is not careful about guarding them from accident.
Some he winds up fully, and they run their full eight days.
 T H E I N H E R I T A N C E OF DURATION 151
O t h e r s h e w i n d s only h a l f w a y , a n d t h e y s t o p a f t e r f o u r
d a y s . A g a i n t h e clock which h a s been w o u n d u p f o r t h e
full e i g h t d a y s m a y fall off t h e Rhelf a n d be b r o u g h t to a
stop a t t h e t h i r d d a y . O r s o m e o n e m a y t h r o w s o m e s a n d
in the w o r k s whon t h e c a r e t a k e r is off his g u a r d . S o ,
similarly, s o m e m e n b e h a v e a s t h o u g h t h e y h a d b e e n
wound u p for a full 9 0 - y e a r r i m , while o t h e r s a r e b u t
p a r t i a l l y w o u n d u p a n d s t o p a t 40 o r 65, o r s o m e o t h e r
point. Or, a g a i n , t h o m a n w o u n d u p f o r 80 y e a r s m a y ,
like t h e clock, be b r o u g h t u p m u c h s h o r t of t h a t b y a n
accidental invasion of m i c r o b e s , p l a y i n g t h e role of t h e
s a n d in t h e w o r k s of t h e clock. I t is of no a v a i l f o r
either t h e clock o r the* m a n to s a y t h a t t h e e l e m e n t s of t h o
m e c h a n i s m a r e in whole o r in m a j o r p a r t c a p a b l e of f u r -
t h e r service. T h e e s s e n t i a l p r o b l e m i s : w h a t d e t e r m i n e s
the g o o d n e s s of t h e o r i g i n a l w i n d i n g ? A n d w h a t r e l a -
tive p a r t do e x t e r n a l thingH p l a y in b r i n g i n g t h e r u n n i n g
to a n end b e f o r e t h e t i m e which t h e o r i g i n a l w i n d i n g w a s
good f o r f I t is w i t h t h i s p r o b l e m of t h e w i n d i n g u p a n d
r u n n i n g of t h e h u m a n m e c h a n i s m t h a t t h e p r e s e n t c h a p -
t e r will d e a l .
T h e r e a r e t w o g e n e r a l c l a s s e s of f a c t o r s w h i c h m a y
\m involved h e r e . T h e s e a r e , on t h e one h a n d , h e r e d i t y
and, on t h e o t h e r h a n d , e n v i r o n m e n t , u s i n g t h e l a t t e r t e r m
in the b r o a d e s t s e n s e . I n a s m u c h a w e c a n b e r e a s o n -
ably s u r e on a priori g r o u n d s t h a t longevity, like m o s t
o t h e r biological p h e n o m e n a , is influenced b y b o t h h e r e d -
i t y a n d e n v i r o n m e n t tho p r o b l e m p r a c t i c a l l y r e d u c e s itself
to t h e m e a s u r i n g of t h e r e l a t i v e i m p o r t a n c e of e a c h of
t h e s e t w o f a c t o r g r o u p s in d e t e r m i n i n g t h e r e s u l t s we see.
B u t before we s t a r t t h e d i s c u s s i o n of e x a c t m e a s u r e m e n t s
in t h i s field l e t u s first e x a m i n e s o m e of t h e g e n e r a l evi-
1 5 2
BIOLOGY OF D E A T H
d e n c e t h a t h e r e d i t y p l a y s a n y p a r t a t all in the deter-
m i n a t i o n o f longevity.

THE HYDE FAMILY

T h e first m a t e r i a l which we shall discuss is t h a t p r o -
v i d e d b y t h e d i s t i n g u i s h e d eugenist, D r . A l e x a n d e r
G r a h a x n B e l l , in h i s s t u d y of t h e H y d e family. E v e r y
g e n e a l o g i s t i s f a m i l i a r w i t h t h e " G e n e a l o g y of the H y d e
F a m i l y , " b y E e n b e n H . W a l w o r t h . I t is one of the fin-
e s t e x a m p l e s i n existence of 'careful a n d p a i n s t a k i n g
g e n e a l o g i c a l r e s e a r c h . U p o n t h e d a t a included in this
b o o k , B e l l h a s made a most interesting and penetrating
a n a l y s i s o f t h e f a c t o r s influencing longevity. A t first
t h o u g h t o n e m i g h t conclude t h a t highly b i a s e d r e s u l t s
w o u l d p r o b a b l y flow f r o m t h e c o n s i d e r a t i o n of only one
family. B e l l m e e t s t h i s p o i n t v e r y well, however, in the
following words:
A little consideration will show that the descendants did not constitute
a single family at all, and indeed had very little of the Hyde blood in them.
Kven the children of William Hyde owed only half of their blood to
him, and one-half to his wife. The grandchildren owed only one-quarter of
their blood to William Hyde, and three-quarters to other people, etc. The
descendants of the seventh generation, and there are hundreds of them, owed
only one sixty-fourth of their blood to William Hyde, and sixty-three
sixty-fourths to the new blood introduced through successive generations of
marriages "with persons not of the Hyde blood at all.
I t will thus be seen that the thousands of descendants noted in the
Hyde Genealogy constitute rather a sample of the general population of
the country than a sample of a particular family in which family traits
might be expected to make their appearance.
T h e s u b s t a n t i a l n o r m a l i t y of t h e m a t e r i a l is shown
i n F i g u r e 4 1 , which gives t h e lx line, t h a t is, t h e n u m b e r
o f s u r v i v o r s a t each age, of t h e 1,606 m a l e s a n d 1,352
f e m a l e s f o r whojn d a t a w e r e available. T h e solid line
i s t h e m a l e lx line a n d t h e d o t t e d line t h e female ls. It
i s a t o n c e a p p a r e n t that the curves have the same general
 THE I N H E R I T A N C E OF DURATION 153
sweep in t h e i r p a s s a g e o v e r t h e s p a n of life a s h a s t h e
g e n e r a l p o p u l a t i o n life c u r v e d i s c u s s e d i n t h e p r e c e d i n g
c h a p t e r . T h e d e s c e n t is a little s t e e p e r i n e a r l y a d u l t
life. T h e female c u r v e differs i n two r e s p e c t s f r o m t h e
n o r m a l g e n e r a l p o p u l a t i o n c u r v e s . I n t h e first place,

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

Mother's age at death

Person's age at death
Stated -40 40-60 60-80 80 +
Stated. 1,805 191 435 713 466
Under 20 511 88 129 199 95
20 and under 40 . 407 42 104 176 85
40 and under 60 . 379 27 92 159 101
60 and under 80 . 360 26 80 129 125
80 and over 148 8 30 50 60
Percentages
Stated 100.0 10.6 24.1 39.5 25.8
Under 20 100.0 17.2 25.2 39.0 18.6
20 and under 40 100.0 10.3 25.6 43.2 20.9
40 and under 60 100.0 7.1 24.3 42.0 26.6
60 and under 80 100.0 7.2 22.2 35.9 34.7
80 and over 100.0 6.4 20.3 33.8 40.5
 156 BIOLOGY OF D E A T H
is t o t h e s e l a s t d o t s t h a t a t t e n t i o n should be p a r t i c u l a r l y
directed. I t will be n o t e d t h a t t h e d o t t e d line connecting
t h e l a s t d o t s of e a c h of t h e 5 p o l y g o n s in g e n e r a l rises
a s we p a s s f r o m the left-hand side of t h e d i a g r a m to the
r i g h t - h a n d side. I n t h e case of offspring d y i n g u n d e r 20,
24 p e r cent, of t h e i r f a t h e r s died a t a g e s o v e r 80. A b o u t
$38 467
PLRSQNS
DIED OHO Mb
-2O ?Q +0 40 "00
30

40 to so
Via. 42.Influence
42.Influence of fntWa a af*AtAt<i^ftth
<i^ftth uptm
uptm bngevit
bngevity of tiiUprtua Iirst <itt in
each di
diagram how
h th th n<*r<!imtaglli&vtna;f fftthTs
h who
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d tntMi mmml < h gttwai
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nge having fathr who dwd from 4t>0J;4t>-0fJ; third dot thu purmmUkWn
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h dm!d!
f 00-U;
from 00U futh
fourth ddot
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theppreiUg@
U hhaving i fth father* who <Hd 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

fill 4O7 3'tO SAO

PCk'SONS PR?10N$ PtRSONS PCtSOMS
DffP DlLD OiCD
ZQ-+O 40-60 60-SO oa+

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-
0ntiftf 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)

Number of Number of Per oent. of

Age at death of parents persons persons lived persons lived
80 + 80 +
Stated 1,594 139 8.7

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
overthe 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).

Mothor'fi ag# at death

Father's ago
at death Under 60 00-80 OwrHO

Under 60 32.8 years 33.4 years 36 3 yeans

(128) (120) (74)
60-80 35.8 38.0 46.0
(251) (328) (172)
Over 80 42.3 45.5 2.7
(131) cm) (1B4)

the figure is 32.8 yearn. I n o t h e r w o r d s , i t a d d e d al-

m o s t e x a c t l y 2 0 y e a r s t o t h e a v e r a g e l i f e of t h e first
g r o u p of p e o p l e t o h a v e e x t r e m e l y l o n g - l i v e d parents,
i n s t e a d of p a r e n t s d y i n g u n d e r a g e 6 0 . In each column
of t h e t a b l e t h e a v e r a g e d u r a t i o n of l i f e advance*! a s w e
proceed from top to bottomthat is, a s the father's age
at death increasesand in each r o w of the table the aver-
a g e expectation of life of the o f f s p r i n g i n c r e a s e s a s w e
p a s s f r o m left t o r i g h t t h a t i s , w i t h i n c r e a s i n g a g e of
the mother at death- H o w e v e r t h e m a t t e r fa t a k e n , a
c a r e f u l s e l e c t i o n of o n e ' s p a r e n t s i n r e s p e c t of l o n g e v i t y
i s t h e m o s t r e l i a b l e f o r m of p e r s o n a l l i f e i n s u r a n c e .
 THE INHERITANCE OF DURATION 161
H o w g r e a t a n d d e e p is t h e significance of t h e facts
shown in T a b l e 16 m a y b e s t be b r o u g h t h o m e t o t h e m i n d
b y m e a n s of a c o m p a r i s o n . S u p p o s e t h i s q u e s t i o n to
be a s k e d : b y h o w g r e a t a n a m o u n t w o u l d t h e a v e r a g e
expectation of life a t b i r t h (which i n a s t a b l e p o p u l a t i o n
is the s a m e t h i n g a s t h e m e a n d u r a t i o n of life) be i n c r e a s e d
if all t h e r e a s o n a b l y p r e v e n t a b l e d e a t h s w e r e p r e v e n t e d ?
If, s a y 75 p e r cent, of all t h e d e a t h s f r o m p u l m o n a r y t u b e r -
culosis did n o t o c c u r ; if 40 p e r cent, of t h e d e a t h s from
B r i g h t ' s d i s e a s e w e r e p r e v e n t e d ; a n d , i n g e n e r a l , if
all t h a t medicine a n d h y g i e n e k n o w s t o d a y w e r e p u t
i n t o r e a s o n a b l y effective o p e r a t i o n , a n d n o b o d y died
except w h e n a n d f r o m s u c h c a u s e s a s could i n no
w a y be influenced b y w h a t m e d i c a l science, g o o d envi-
r o n m e n t , etc., h a v e t o offer: b y h o w m u c h then would
t h e expectation of life be g r e a t e r t h a n i t n o w i s ? W e
h a v e seen t h a t t o h a v e o n e ' s p a r e n t s live t o 80 o r over
i n c r e a s e s t h e e x p e c t a t i o n of life 20 y e a r s , a s c o m p a r e d
w i t h t h a t of p e r s o n s w h o s e p a r e n t s d i e u n d e r 60 y e a r s of
age. B y h o w m u c h m o r e w o u l d t h e e x p e c t a t i o n of life
be extended if a l l r e a s o n a b l y p r e v e n t a b l e d e a t h s w e r e
prevented?
A t h o r o u g h a n d c r i t i c a l a n s w e r t o t h i s q u e s t i o n is
afforded by .an i n v e s t i g a t i o n of F o r s y t h ' s ) , c o n d u c t e d
along t h e m o s t e x a c t a n d a p p r o v e d a c t u a r i a l lines.
Some y e a r s a g o , P r o f e s s o r I r v i n g F i s h e r s e n t a l i s t of
some 90 d i s e a s e s t o a g r o u p of t h e m o s t p r o m i n e n t medi-
cal a u t h o r i t i e s i n t h i s c o u n t r y , a n d a s k e d t h e m t o desig-
n a t e w h a t p e r c e n t a g e of t h e d e a t h s d u e t o e a c h disease
t h e y considered p r e v e n t a b l e . T h e r e s u l t s of t h i s i n q u i r y
were tabulated in an extremely conservative manner,
w i t h t h e r e s u l t s e t f o r t h i n T a b l e 16a, w h i c h i s copied
f r o m F o r s y t h ' s p a p e r ( p p . 762-763).
li
162 BIOLOGY O F D E A T H
TABLE 16 a
Showing Fisher's ratios of preventability for the diseases enumerated in the
mortality statistics of the United States, together with the relative
importance of each disease as indicated by the percentage the
number of its deaths bears to the total number of deaths
Prominence of Ratio of
Causes of death disease. Percent, preventability.
of all deaths Per cent.
Premature birth 2.0 40
.55 0
3 Congenital malformation
Other congenital of the heart
malformations .3 0
4 Congenital debility 2.3 40
5 Hydrocephalus .1 0
6 Venereal diseases .3 70
7 Diarrhoea and enteritis 7.74 60
8 Measles .8 40
9 Acute bronchitis 1.1 30
10 Bronchopneumonia 2.4 50
11 Whooping cough 40
12 Croup 75
13 Meningitis 1.6 70
14 Diseases of larynxnot laryngitis .07 40
15 Laryngitis .06 40
16 Diphtheria 1.4 70
17 Scarlet fever .5 50
18 Diseases of lymphatics .01 20
19 Tonsillitis .05 45
20 Tetanus .19 80
21 Tuberculosisnot of lungs .17 75
22 Abscess .08 60
23 Aprendicitis .7 50
24 Typhoid fever 2.0 85
25 Puerperal convulsions .2 30
26 Puerperal septicaemia .4 85
27 Other diseases of childbirth .36 50
28 Diseases of tubes .1 65
29 Peritonitis .5 55
30 Smallpox .01 75
31 Tuberculosis of lungs 9.9 75
32 Violence 7.5 35
33 Malarial fever .2 80
34 Septicaemia .3 40
35 Epilepsy .29 0
36 General, ill-defined, and unknown causes (in-
cluding "heart failure," "dropsy," and "con-
vulsions") 9.2 30
37 Erysipelas .3 60
38 Pneumonia (lobar and unqualified) 7.0 45
39 Acute nephritis .6 30
40 Pleurisy .27 55
41 Acute yellow atrophy of liver .02 0
.6 25
42 Obstructions of intestines
 T H E I N H E R I T A N C E OF D U R A T I O N 163
TABLE 16 aContinued

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

Deaths Loss in Deaths Loss in

Age Not pre- Pre- Age Not pre- Pre-
vented vented Years Days vented vented Years Days
0., 49.44 62.11 12 245 25.. 39.31 46.18 318
1.. 56.03 66.26 10 84 26.. 38.56 45.31 274
2.. 56.84 66.28 9 161 27.. 37.82 44.45 230
3., 56.64 65.67 9 11 28.. 37.08 43.58 183
4.. 56.15 64.94 8 288 29.. 36.34 42.72 139
5.. 55.51 64.13 8 226 30.. 35.61 41.86 91
6.. 54.81 63.27 8 168 31.. 34.88 41.01 47
7.. 54.06 62.42 8 131 32.. 34.15 40.15 0
8.. 53.26 61.54 8 102 33.. 33.42 39.30 321
9.. 52.43 60.63 8 73 34.. 32.69 38.46 281
10.. 51.57 59.72 8 55 35.. 31.96 37.61 237
11.. 50.69 58.79 8 37 36.. 31.23 36.76 193
12.. 49.80 57.86 8 22 37.. 30.50 35.92 153
13.. 48.91 56.80 7 321 38.. 29.77 35.08 113
14.. 48.03 56.00 7 354 39.. 29.03 34.24 77
15.. 47.15 55.07 7 336 40.. 28.30 33.40 37
16.. 46.31 54.16 7 310 41.. 27.57 32.57 0
17.. 45.50 53.26 7 277 42.. 26.85 31.74 325
18.. 44.71 52.36 7 237 43.. 26.12 30.91 288
19.. 43.93 51.48 7 201 44.. 25.40 30.09 252
20.. 43.15 50.59 7 161 45.. 24.68 29.28 219
21.. 42.37 49.70 7 120 46.. 23.97 28.47 183
22.. 41.60 48.82 7 80 47.. 23.26 27.67 150
23.. 40.83 47.94 7 40 48.. 22.56 26.87 113
24.. 40.07 47.06 6 261 49.. 21.87 26.09 80

On. t h e b a s i s of the m o r t a l i t y e x p e r i e n c e of the Eegis-

tration A r e a for 11 y e a r s (1900-1910) Forsyth calculated
mortality tables on the assumption that the ratios of
preventability of Table 16a were actually in full opera-
tion. The results, so far as concerns expectation of life,
are set forth in Table 16b.
 THE INHERITANCE OF DURATION 165
F r o m t h e first line of t h i s t a b l e i t i s p e r c e i v e d t h a t
the. t o t a l i n c r e a s e i n e x p e c t a t i o n of life w h i c h w o u l d
r e s u l t if F i s h e r ' s r a t i o s of p r e v e n t a b i l i t y w e r e fully
realized is just under 13 years! How unfavorably this
c o n t r a s t s w i t h t h e 20 y e a r s i n c r e a s e s h o w n b y t h e t w o
TABLE 16bContinued
Deaths Loss in Deaths Loss in
Age Not pre- pre- Age Not pre- Pre-
vented vented Years Days vented vented Years Days
50. 21.17 25.30 47 71.. 8.82 11.15 120
51.. 20.47 24.52 18 72.. 8.36 10.59 84
52. 19.78 23.74 350 73.. 7.93 10.04 40
53. 19.09 22.97 321 74.. 7.50 9.51 4
54. 18.40 22.21 296 75.. 7.09 8.99 329
55. 17.74 21.46J 263 76.. 6.70 8.49 288
56. 17.08 20.72 234 77.. 6.31 8.00 252
57.. 16.45 20.00 201 78.. 5.98 7.53 201
58., 15.83 19.30 193 79.. 5.64 7.07 157
59. 15.23 18.61 139 80.. 5.32 6.63 113
60., 14.63 17.93 110 81.. 5.02 6.20 66
61., 14.05 17.27 80 82.. 4.74 5.78 15
62. 13.48 16.61 47 83.. 4.47 5.38 332
63., 12.92 15.96 15 84.. 4.23 4.99 277
64., 12.36 15.32 350 85.. 4.01 4.62 223
65., 11.82 14.69 348 86.. 3.79 4.25 168
66., 11.29 14.07 285 87.. 3.58 3.89 113
67., 10.77 13.47 256 88.. 3.39 3.56 62
68., 10.26 12.87 223 89.. 3.22 3.27 18
69.. 9.77 12.29 190 90.. 3.06 3.06 0
70., 9.29 11.71 153

corner diagonal cells of Table 16! No more striking

demonstration could be found of the overwhelming im-
p o r t a n c e of h e r e d i t y i n d e t e r m i n i n g d u r a t i o n o f l i f e . For
if all the d e a t h s w h i c h r e a s o n will j u s t i f y o n e i n suppos-
i n g p r e v e n t a b l e o n t h e b a s i s of w h a t is n o w k n o w n , w e r e
prevented in fact the resulting increase in expectation
of life falls seven years short of what might reasonably
be expected to follow the selection of only one generation
of ancestry {the parental) for longevity.
S o m u c h f o r B e l l ' s a n a l y s i s of l o n g e v i t y i n t h e Hyde
166 BIOLOGY OF D E A T H
family. W e h a v e seen t h a t i t d e m o n s t r a t e s w i t h t h e ut-
m o s t clearness a n d c e r t a i n t y t h a t t h e r e i s a n h e r e d i t a r y
influence between p a r e n t a n d offspring affecting t h e ex-
pectation of l o n g e v i t y of t h e l a t t e r . B e l l ' s m e t h o d of
handling t h e m a t e r i a l does n o t p r o y i d e a n y p r e c i s e meas-
u r e of t h e i n t e n s i t y of t h i s h e r e d i t a r y influence, n o r does
it f u r n i s h a n y i n d i c a t i o n of its s t r e n g t h i n a n y b u t the
direct line of descent. Of c o u r s e , if h e r e d i t y is a factor
in the d e t e r m i n a t i o n of l o n g e v i t y w e should expect its
effects t o be m a n i f e s t e d a s b e t w e e n b r o t h e r s a n d sisters,
or in t h e a v u n c u l a r r e l a t i o n s h i p s , a n d i n g r e a t e r o r less
degree i n all t h e o t h e r c o l l a t e r a l a n d m o r e r e m o t e direct
degrees of k i n s h i p . H a p p i l y , w e h a v e a p a i n s t a k i n g
analysis, w i t h a q u a n t i t a t i v e m e a s u r e of t h e r e l a t i v e in-
fluence of h e r e d i t y i n t h e d e t e r m i n a t i o n of longevity,
which w a s c a r r i e d o u t m a n y y e a r s before B e l l ' s w o r k on
the H y d e family, b y t h e p i o n e e r i n t h i s field, P r o f . K a r l
P e a r s o n . H i s d e m o n s t r a t i o n of t h e i n h e r i t a n c e of longev-
ity a p p e a r e d m o r e than, t w e n t y y e a r s before t h a t of
Bell. I h a v e called a t t e n t i o n t o the l a t t e r ' s w o r k first
merely because of t h e g r e a t e r simplicity a n d directness of
his d e m o n s t r a t i o n . W e m a y n o w t u r n t o a consideration
of P e a r s o n ' s m o r e d e t a i l e d r e s u l t s .

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)

Relatives Correlation Ratio of

coefficient coefficient to its
X probable error
y
Father Adult son 0.135 db .021 6.4
Father Minor son .087 .022 4.0
Father Adult daughter .130 .020 6.5
Father Minor daughter .052 =b .023 2.3
Mother Adult son .131:4= .019 6.9
Mother Minor son .076 =fc .024 3.2
Mother Adult daughter .149 =b .020 7.5
Mother Minor daughter .138=b .024 5.7
Elder adult brother Younger adult brother .229 .019 12.1
Adult brother Adult brother .285 db .020 14.3
Minor brother Minor brother .103=b .029 3.6
Adult brother Minor brother -.026 .025 1.0
Elder adult sister Younger adult sister .346 d= .018 19.2
Adult sister Adult sister .332 db .019 17.5
Minor sister Minor sister .175 .031 5.6
Adult sister Minor sister -.026 .029 .9
Adult brother Adult sister .232 .015 15.5
Minor brother Minor sister .144=fc .025 5.8
Adult brother Minor sister -.006 .035 .2
Adult sister Minor brother -.027 db .024 1.1
The cases above the horizontal line are all direct lineal inheritance;
those below the line collateral inheritance.

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 casesnamely, 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 charactersstature,
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:

Substituting the ascertained values w e have

1. F r o m parental correlations.
0.1365 = .4675 (1-p)2
(1-p) = .292
(1-p) = .54
2. F r o m fraternal correlations
0.2831 = .5156 (1-p) *
(1-p) = .74
F r o m these figures i t may be concluded, and P e a r s o n
does so conclude, that from 50 to 75 p e r cent, of the
general death rate within the group of the population on
which the calculations are based, is determined funda-
mentally by factors of heredity and is not capable of
essential modification or amelioration by any sort of
environmental action, however well intentioned, however
costly, or however well advertised. Mutatis mutandis
the same conclusion applies to the duration of life. I have
 T H E I N H E R I T A N C E OF D U R A T I O N 177
preferred t o s t a t e the conclusion i n terms of death rates,
as i t w a s originally stated by Pearson, because of the
bearing i t has upon a g r e a t deal of the public health
propaganda so loosely flung about. I t need only be re-
membered that there i s a perfectly definite functional
relation between death rate and average duration of life
in a n approximately stable population group, expres-
sible by an equation, in order to sec that any conclusion
as to the relative influence of heredity and environment
upon the general death rate must apply with equal force
to the duration of life.

THE SELECTIVE DEATH BATE IN* MAN

If the duration of life w e r e inherited i t would logical-
ly be expected that some portion of the death rate m u s t
be selective in character. F o r inheritance of duration
of life can only m e a n that when a person dies i s in p a r t
determined by that individual's biological constitution or
makeup. A n d equally it i s obvious that individuals of
weak and unsound constitution must, on the average,
die earlier than t h o s e of strong, sound, and vigorous con-
stitution. W h e n c e i t follows that the chances of l e a v i n g
offspring will be g r e a t e r f o r those of sound constitution
than for the weaklings. T h e mathematical discussion
which has j u s t been given indicates t h a t from one-half
to three-fourths of t h e death rate i s selective in char-
acter, because t h a t proportion is determined by hereditary
factors. J u s t i n proportion a s heredity determines
the death rate, s o i s the mortality selective. The reality of
the f a c t of a selective death rate i n m a n can be easily
shown graphically.
I n F i g u r e 4 4 are seen, the g r a p h s of some d a t a f r o m
E u r o p e a n royal families, w h e r e no neglect of children,
12
178 BIOLOGY OF D E A T H
degrading environmental conditions, or economic want
can have influenced the results. These data were COin-

so
A5

*
35
b
30

25
: 20
15 MOTHER
FATHER AND
AND CHILDREN
CHILDREN

10

16 26 36 56 66 76 66 and over

AGE AT DEATH OF PARENTS

FIG. 44.Diagram showing the influence of age at death of parents upon the percentage of
offspring dying under 5 years. (After Ploetz).
piled by the well-known German eugenist, Professor
Ploetz of Munich. The lines show the falling per-
 THE INHERITANCE OF DURATION 179
centage of t h e i n f a n t i l e d e a t h r a t e a s t h e d u r a t i o n of
life of t h e f a t h e r a n d m o t h e r i n c r e a s e s . A m o n g t h e chil-
d r e n of short-lived f a t h e r s a n d m o t h e r s , a t t h e left end
of each line, is f o u n d t h e h i g h e s t i n f a n t m o r t a l i t y , while
a m o n g the offspring of long-lived p a r e n t s t h e lowest
infant m o r t a l i t y occurs, as shown a t t h e r i g h t - h a n d end of
the d i a g r a m .
T h e r e s u l t s so f a r p r e s e n t e d r e g a r d i n g a selective
d e a t h r a t e a n d i n h e r i t a n c e of d u r a t i o n of life, h a v e c o m e
from selected c l a s s e s : t h e a r i s t o c r a c y , r o y a l t y o r Q u a k e r s .
None of t h e s e classes c a n be f a i r l y said t o r e p r e s e n t t h e
general p o p u l a t i o n . C a n t h e conclusion be t r a n s f e r r e d
safely f r o m t h e classes t o t h e m a s s e s ? T o t h e d e t e r m i n a -
tion of t h i s p o i n t one of P e a r s o n ' s s t u d e n t s , D r . E . C. Snow,
a d d r e s s e d himself. T h e m e t h o d which h e u s e d w a s , f r o m
the necessities of t h e case, a m u c h m o r e c o m p l i c a t e d a n d
i n d i r e c t one t h a n t h a t of P e a r s o n a n d P l o e t z . I t s essen-
tial i d e a w a s t o see w h e t h e r i n f a n t d e a t h s w e e d e d o u t t h e
unfit a n d left a s s u r v i v o r s t h e s t r o n g e r a n d m o r e r e s i s -
t a n t . All t h e i n f a n t s b o r n i n a single y e a r w e r e t a k e n
a s a cohort a n d t h e d e a t h s o c c u r r i n g i n t h i s c o h o r t i n suc-
cessive y e a r s w e r e followed t h r o u g h . R e s o r t w a s h a d
t o t h e m e t h o d of p a r t i a l o r n e t c o r r e l a t i o n . T h e v a r i a b l e s
c o r r e l a t e d i n t h e c a s e of t h e P r u s s i a n d a t a w e r e t h e s e :
1. <p0 = Births in year a given cohort started.
2. a?i = Deaths in the first two years of life.
3. co2 = Deaths in the next eight years of life.
4. xz = Deaths in the ten years of all individuals not included in
the particular cohort whose deaths are being followed.
I n t h e case of t h e E n g l i s h d a t a t h e v a r i a b l e s w e r e :
a0 = Births in specified year.
<vx = Deaths in the first three years of life of those born in
specified year.
x2 = Deaths in fourth and fifth years of life of those born in
specified year.
at = The "remaining" deaths under 5.
 180 BIOLOGY OF D E A T H
The u n d e r l y i n g i d e a w a s t o g e t t h e p a r t i a l or n e t
correlation between xx a n d x2, while x0 a n d a?8 a r e h e l d
constant. If the m o r t a l i t y of i n f a n c y is selective, i t s
amount should be n e g a t i v e l y c o r r e l a t e d t o a s i g n i f i c a n t
degree with the m o r t a l i t y of t h e n e x t e i g h t y e a r s w h e n
the b i r t h s i n each d i s t r i c t c o n s i d e r e d a r e m a d e c o n -
stant a n d when the g e n e r a l h e a l t h e n v i r o n m e n t is m a d e
constant. U n d e r t h e c o n s t a n t c o n d i t i o n s specified a
negative correlation d e n o t e s t h a t t h e h e a v i e r the i n f a n -
TABLE 22
Snow'8 results on selective death rate in man. Engluh and Prussian
rural districts
Actual correlation Expected correlation
Data r if no election
I2.03
Males:
English Rural (1870) -0.4483 -0.0828
Districts (1871) - .3574 - .1014
(1872) - .2271 - .0807
Prussian Rural (1881) - .9278 - .0958
Districts (1882) - .6050 - .0765
Females:
English Rural (1870) - .4666 - .0708
Districts (1871) - .2857 - .0505
(1872) - .5080 - .0496
Prussian Rural (1881) - .8483 - .0933
Districts (1882) - .6078 - .0705
tile d e a t h r a t e i n a c o h o r t of b i r t h s t h e l i g h t e r will b<
t h e . d e a t h r a t e of l a t e r y e a r s , a n d vice versa. The l a s t
variable, # 8 , is t h e one chosen, a f t e r c a r e f u l c o n s i d e r a t i o n
and m a n y trials, to m e a s u r e v a r i a t i o n i n t h e h e a l t h e n v i -
r o n m e n t If a n y y e a r is a p a r t i c u l a r l y u n h e a l t h y o n e a n
epidemic y e a r f o r e x a m p l e t h e n t h i s t m h e a i t h m e s s
should be accurately reflected in t h e d e a t h s of those m e m -
bers of t h e p o p u l a t i o n n o t i n c l u d e d in t h e c o h o r t
u n d e r review.
 T H E I N H E R I T A N C E OF D U R A T I O N 181
S n o w ' s r e s u l t s for E n g l i s h a n d P r u s s i a n r u r a l d i s -
t r i c t s a r e s e t f o r t h i n T a b l e 22. F r o m t h i s t a b l e i t i s
seen t h a t in e v e r y case t h e c o r r e l a t i o n s a r e negative, and
t h e r e f o r e i n d i c a t e t h a t t h e m o r t a l i t y of e a r l y life is selec-
tive. F u r t h e r m o r e , t h e d e m o n s t r a t i o n of t h i s f a c t is
c o m p l e t e d b y s h o w i n g t h a t t h e o b s e r v e d coefficients a r e
f r o m 3 t o 10 t i m e s as g r e a t a s t h e y w o u l d be if t h e r e w e r e
n o selective c h a r a c t e r t o t h e d e a t h r a t e . T h e coefficients
f o r t h e P r u s s i a n p o p u l a t i o n , i t will be n o t e d , a r e of a
d i s t i n c t l y h i g h e r o r d e r of m a g n i t u d e t h a n t h o s e f o r t h e
E n g l i s h p o p u l a t i o n . T h i s d i v e r g e n c e is p r o b a b l y d u e
chiefly t o differences i n t h e q u a l i t y of t h e f u n d a m e n t a l
s t a t i s t i c a l m a t e r i a l i n t h e t w o cases. T h e P r u s s i a n m a -
t e r i a l is f r e e f r o m c e r t a i n defects i n h e r e n t i n t h e E n g l i s h
d a t a , which c a n n o t b e e n t i r e l y g o t r i d of. T h e difference
i n t h e coefficients f o r t h e two successive P r u s s i a n c o h o r t s
r e p r e s e n t s , i n S n o w ' s opinion, p r o b a b l y a r e a l fluctua-
t i o n i n t h e i n t e n s i t y of n a t u r a l selection i n t h e one g r o u p
a s c o m p a r e d w i t h t h e o t h e r . H o w significant S n o w ' s
r e s u l t s a r e i s s h o w n g r a p h i c a l l y i n F i g u r e 45.
S n o w ' s o w n c o m m e n t s o n his r e s u l t s a r e significant.
He says:
The investigations of this memoir have been long and laborious, and
the difficulties presented by the data have been great. Still, the general
result cannot be questioned. Natural selection, in the form of a selective
death-rate, is strongly operative in man in the early years of life.. Those
data which we believe to be the best among those we have usedthe Prus-
sian figuresshow very high negative correlation between the deaths in
the first two years of life and those in the next eight, when allowance is
made for difference in environment. We assert with great confidence that
a high mortality in infancy (the first two years of life) is followed by a
corresponding low mortality in childhood, and conversely. The English
figures do not allow such a comprehensive survey to be undertaken, but,
so far as they go, they point in the same direction as the Prussian ones.
The migratory tendencies in urban districts militate against the detection
of selective influences there, but we express the belief that those influences
182 BIOLOGY OF D E A T H
are just as prevalent in industrial as in rural communities, and could be
measured by other means if the data were forthcoming.

MALES
.7

.3

"EXPECTED CORRELATION - NO SELECTION

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
peoplethe 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-
tiveeliminating 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

INHEBITADSTCE OF DUBATION OF LIFE IN DROSOPHILA

I N t h e l a s t c h a p t e r t h e r e w a s p r e s e n t e d indubitable
proof t h a t i n h e r i t a n c e is a m a j o r f a c t o r i n d e t e r m i n i n g
the d u r a t i o n of life i n m a n . T h e evidence, while entirely
convincing a n d i n d e e d in t h e w r i t e r ' s opinion critically
conclusive, m u s t be, i n t h e n a t u r e of t h e case, statistical
in its n a t u r e . E x p e r i m e n t a l i n q u i r i e s i n t o t h e d u r a t i o n
of h u m a n life a r e obviously i m p o s s i b l e . I t is always
important, however, as a general principle, and particu-
larly so in t h e p r e s e n t i n s t a n c e , t o check o n e ' s statistical
conclusions b y i n d e p e n d e n t e x p e r i m e n t a l evidence. This
can be successfully done, w h e n o n e ' s p r o b l e m is longevity,
only b y c h o o s i n g a n a n i m a l w h o s e life-span r e l a t i v e t o
t h a t of m a n is a s h o r t one, a n d i n g e n e r a l t h e briefer it
is, the b e t t e r s u i t e d will t h e a n i m a l be f o r t h e p u r p o s e .
A n o r g a n i s m which r a t h e r completely fulfils the r e -
q u i r e m e n t s of t h e case, n o t only i n r e s p e c t of the short-
ness of t h e life s p a n , b u t also i n o t h e r w a y s , such as
ease of h a n d l i n g , feeding, h o u s i n g , etc., is t h e common
" f r u i t " or " v i n e g a r " fly, DrosopMla melanogaster.
This insect, which e v e r y one h a s s e e n h o v e r i n g a b o u t
bananas and other fruit in fruit shops, has lately attained
g r e a t f a m e a n d r e s p e c t a b i l i t y a s a l a b o r a t o r y animal,
as a r e s u l t of t h e b r i l l i a n t a n d e x t e n d e d i n v e s t i g a t i o n s
of M o r g a n a n d h i s s t u d e n t s u p o n it, i n a n a n a l y s i s of the
m e c h a n i s m of h e r e d i t y . Drosophila is a small fly, p e r -
 STUDIES ON T H E DURATION OF LIFE 187
h a p s one f o u r t h a s l a r g e a s t h e c o m m o n h o u s e fly. I t
h a s s t r i M n g r e d eyes, a b r o w n i s h body, a n d w i n g s of
length a n d f o r m v a r y i n g i n different s t r a i n s . I t lives
n o r m a l l y on t h e s u r f a c e of d e c a y i n g f r u i t of all s o r t s ,
but because of a m o r e o r less well-marked p r e f e r e n c e f o r

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

the t w o cases. M a n ' s y a r d s t i c k is o n e y e a r long, w h i l e

DrosophUa's is o n e day long. A fly 9 0 d a y s old is j u s t

as decrepit a n d senile, f o r a fly, a s a m a n 9 0 y e a r s o l d

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

life table f o r a d u l t flies o f b o t h s e x e s . T h e s u r v i v o r -

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.

Experimental group Mean duration Number of

of life in days flies
Sterile (Loeb and Northrop) 28.5 70
Non-sterile, males, all genetic lines (Pearl) 31.3 2620
Non-sterile, females, all genetic lines (Pearl) 33.0 3216
Non-sterile, both sexes, all genetic lines (Pearl) 32.2 5836
Difference in favor of non-sterile 3.7
Probable error of difference about db 1.0
"We reach, t h e conclusion t h a t b a c t e r i a - f r e e Drosophila
live n o l o n g e r on t h e a v e r a g e , a n d i n d e e d p e r h a p s e v e n
a little less l o n g , t i n d e r o t h e r w i s e t h e s a m e c o n s t a n t
.202 BIOLOGY OF DEATH
environmental conditions, than do normal non-sterile
indeed germ-ladenflies. This result is of great inter-
est and significance. I t emphasizes in a direct experi-
mental manner that in a broad biological sense bacteria
play but an essentially accidental role in determining
length of the span of life in comparison with the influence
of heredity.
POVEBTY AND DTJKATION OF LIFE
But we must take care lest we seem to convey the
impression that no sort of environmental influence can
affect the average duration of life. Such a conclusion
would be manifestly absurd. Common sense tells us
PERSONAL PROPERTY TAX IN PARIS 1911 -1313
I 1 PAYING i EXENPT

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 -

ness, b y t h e e m i n e n t Swiss, statistician, H e r s c h , m a y

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).

right in broader bars for the four groups of arrondisse-

ments separated by wider spaces in the detailed dia-
gram, and finally for Paris as a whole. It will be seen
that the poverty of the population, m e a s u r e d b y the per-
sonal property yardstick, is least at the left-hand end of
the diagram, where the smallest percentages of fam-
ilies are exempted from the tax, and greatest at the
right-hand end, w h e r e scarcely any of the population is
well enough to do to p a y this tax.

Figure 51 shows the death rates from all c a u s e s for

the same arrondissements and the same groups. It is
at once apparent that the black bars in this g r o u p r u n in
a general manner parallel to the preceding one. The
204 BIOLOGY OF D E A T H
p o o r e s t d i s t r i c t s h a v e t h e h i g h e s t d e a t h r a t e s , t h e richest
d i s t r i c t s t h e lowest d e a t h r a t e s , a n d d i s t r i c t s i n t e r m e -
d i a t e in r e s p e c t of p o v e r t y a r e also i n t e r m e d i a t e in r e s -
p e c t of m o r t a l i t y . O n t h e face of t h e evidence t h e r e
w o u l d seem t o h e h e r e complete p r o o f of t h e overwhelm-
i n g l y i m p o r t a n t influence u p o n d u r a t i o n of life of degree
of p o v e r t y , w h i c h is p e r h a p s the m o s t p o t e n t single envi-
r o n m e n t a l f a c t o r affecting civilized m a n to-day. But,
a l a s , pitfalls p r o v e r b i a l l y l u r k i n s t a t i s t i c s . B e f o r e we
c a n accept t h i s so a l l u r i n g r e s u l t a n d go a l o n g w i t h our
a u t h o r t o h i s final s o m e w h a t s t u p e n d o u s conclusion t h a t
if t h e r e w e r e no p o v e r t y t h e d e a t h r a t e f r o m c e r t a i n im-
p o r t a n t causes, a s f o r e x a m p l e t u b e r c u l o s i s , would forth-
w i t h become zero, w e m u s t exercise a l i t t l e inquisitive
c a u t i o n . W h a t evidence is t h e r e t h a t t h e i n h a b i t a n t s of
t h e d i s t r i c t s s h o w i n g a h i g h p o v e r t y r a t e a r e n o t biologi-
cally a s well a s economically differentiated f r o m the in-
h a b i t a n t s of d i s t r i c t s w i t h a low p o v e r t y r a t e ? A n d
a g a i n w h a t is t h e evidence t h a t i t is n o t such biological
differentiation r a t h e r t h a n t h e economic which d e t e r m i n e s
t h e d e a t h r a t e differences i n the t w o cases ? U n f o r t u n a t e l y ,
o u r a u t h o r gives u s n o w h i t of evidence on t h e s e obviously
s o i m p o r t a n t p o i n t s . H e m e r e l y a s s u m e s , because of t h e
f a c t s shown, t h a t if s o m e o m n i p o t e n t spook w e r e t o t r a n s -
p o s e all t h e i n h a b i t a n t s of t h e M e n i l m o n t a n t a r r o n d i s s e -
m e n t t o t h e E l y s e e a r r o n d i s s e m e n t , a n d vice versa for
example, and w e r e to p e r m i t each group to annex the
w o r l d l y goods of t h e d i s p o s s e s s e d g r o u p , t h e n t h e d e a t h
r a t e s w o u l d be f o r t h w i t h i n t e r c h a n g e d . T h e r e is no r e a l
evidence t h a t a n y s u c h r e s u l t w o u l d follow a t all. One
c a n n o t s h a k e i n t h e s l i g h t e s t d e g r e e f r o m i t s solidly
g r o u n d e d f o u n d a t i o n t h e critically d e t e r m i n e d fact of
t h e p a r a m o u n t i m p o r t a n c e of t h e h e r e d i t a r y f a c t o r i n
d e t e r m i n i n g r a t e s of m o r t a l i t y , w h i c h h a v e b e e n s u m m a -
 ON T H E DURATION OF LIFE 205
1:ri
- 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 b y a n y s u c h evi-
t h a t of H e r s c h .
TABLE 24
*^_^^^As in Paris (1911-15 0 by classes ofarrondissements (Hersch)
a s Absolute figures Stillbirths
* s<etS o f Arrondissement8 per 100 living
Stillbirths Living births births
I 1,004 12,313 8.2
II 1,390 19,998 7.0
III 7,279 82,821 8.8
IV 3,024 30,853 9.8
Paris 12,679 145,985 8.7
, indeed, lie himself finds to be t h e f a c t w h e n h e
t h e e x t r e m e l y s e n s i t i v e i n d e x of h e r e d i t a r y
b i o l g " i o L i c o n s t i t u t i o n f u r n i s h e d "by t h e s t i l l b i r t h r a t e .
l ^ 2 4 gives t h e d a t a . W e see a t once t h a t t h e r e is n o
s t r i k i n g i n c r e a s e i n t h e foetal m o r t a l i t y a s w e p a s s
f r o X t x " t t t e r i c h e s t class of d i s t r i c t s , as w a s s h o w n i n t h e
d e a - t t i r a t e f r o m all c a u s e s . I n s t e a d t h e r e is p r a c t i c a l l y
no < 3 l x a . : n _ g e , c e r t a i n l y n o n e of significance, a s w e p a s s
f r o x x x o n e class of d i s t r i c t s to a n o t h e r . T h e r a t e is 8.2
per- 1 O O l i v i n g births* i n t h e r i c h e s t c l a s s a n d 9.8 i n
the p o o r e s t
O t 3 b . o i r definite evidence t h a t such conclusion as t h o s e
of H e a r s o i l c a n n o t b e accepted a t a n y t h i n g like t h e i r f a c e
v a l u e i s . a f f o r d e d b y t h e w o r k of G r e e n w o o d a n d B r o w n
o n "fclxe r e l a t i o n of p o v e r t y a n d t h e i n f a n t d e a t h r a t e .
T h e y dBLxid, g i v i n g s u b s c r i p t s t h e following m e a n i n g s :
STiTbscript 1 = Birth rate
Six*fc>script 2 = Artificial feeding rate
S"ixt>script 3 = Poverty rate
SxxTbscript 4 = Infant death rate
th.a.t

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

This coefficient m e a n s t h a t t h e differences in i n f a i

mortality r a t e in the different d i s t r i c t s of L o n d o n , w h c
the b i r t h r a t e is m a d e constant, a r e n o t a s s o c i a t e d w H
differences in p o v e r t y between t h e s a m e d i s t r i c t s t o a
extent sufficient to lead to a c o r r e l a t i o n coefficient s e n * !
bly different f r o m zero.
Finally, Stevenson h a s , since t h e a p p e a r a n c e <
H e r s c h ' s p a p e r , studied t h e s a m e p r o b l e m s on t h e b a n
of the London data, for t h e s a k e of c o m p a r i s o n w i
the results from P a r i s . H e t a k e s a s t h e index o f e c
nomic s t a t u s t h e n u m b e r of d o m e s t i c s e r v a n t s ( o f h o
sexes) p e r 100 of population, a n d h a s e x a m i n e d t h e den
r a t e s from all causes, i n f a n t m o r t a l i t y , a n d t u b e r c u l a r
for the identical y e a r s t h a t H e r s c h u s e d . T h e r e s u l
a r e set f o r t h in T a b l e 24a.
Commenting on the facts r e g a r d i n g g e n e r a l m o r t a l !
from all causes i n London, S t e v e n s o n s a y s :
"These bear an altogether different aspect from the Parisian
Whereas the latter increase so regularly with poverty that the
rate for any district in one group never exceed** the lowest for any dint!
in the next poorer group, in London the gradation, even for the %rt*
themselves, is irregular, the lowest death-rate not being returned for
 STUDIES ON T H E DURATION OF LIFE 207
TABLE 24 a
Mortality of London boroughs grouped by wealth. (From Stevenson)

(both sexes) per cent,

Infant mortality Deatll-rate

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 .

EXPEKIMENTS ON TEMPEEATUKB AND DURATION OF MFB

A l t o g e t h e r i t i s p l a i n t h a t w e n e e d a n o t h e r k i n d of
evidence t h a n t h e s i m p l e u n a n a l y z e d p a r a l l e l i s m which
Hersch demonstrates between poverty and the general
d e a t h r a t e if w e a r e t o g e t a n y d e e p u n d e r s t a n d i n g of t h e
influence of e n v i r o n m e n t a l c i r c u m s t a n c e s u p o n t h e d u r a -
tion of life o r t h e g e n e r a l d e a t h r a t e . W e shall d o well
to t u r n again to the experimental method. About a
dozen y e a r s a g o L o e b ,
starting from the idea that chemical conditions In the organ bin art* tnw
of the main variables in this case, raided the question whether ther# wa a
definite coefficient for the duration of life and whether thin temperature
coefficient was of the order of magnitude of that of a chemical reaction.
The first experiments were made on the unfertilized and fertilized
 STUDIES ON T H E DURATION OF LIFE 209
of the sea urchin and could only be carried out at the upper temperature
limits of the organism, since at ordinary temperatures this organism lives
for years. In the upper temperature region the temperature coefficient
for the duration of life was very high, probably on account of the fact
that, at this upper zone of temperature, death is determined by a change
of the nature of a coagulation or some other destructive process. Moore,
at the suggestion of Loeb, investigated the temperature coefficient for the
duration of life for the hydranth of a tubularian at the upper temperature
limit and found that it was of the same order of magnitude as that
previously found for the sea urchin egg. In order to prove that there
is a temperature coefficient for the duration of life throughout the whole
scale of temperatures at which an organism can live, experiments were
required on a form whose duration of life was short enough to measure
the duration of life even at the lowest temperature.
A suitable o r g a n i s m w a s f o u n d i n Drosophila. This
w a s g r o w n u n d e r a s e p t i c conditions, a s a l r e a d y described.
The general r e s u l t s a r e s h o w n i n T a b l e 25.
TABLE 25
Effect of temperature on duration of life of Drosophila.
(After Loeb and Northrop)
Duration (in days) of
Temperature Total duration
Larval stage Pupal stage Life of of life from egg
imago to death
C
10 57 PupaB die 120.5 177.5 + x
15 17.8 13.7 92.4 123.9
20 7.77 6.33 40.2 54.3
25 5.82 4.23 28.5 38.5
27.5 (4.15) 3.20 .... ....
30 4.12 3.43 13.6 21.15
F r o m t h i s t a b l e i t is seen t h a t a t t h e l o w e s t t e m p e r a -
t u r e t h e d u r a t i o n of life i s longest, a n d a t t h e h i g h e s t t e m -
p e r a t u r e s h o r t e s t . Cold slows u p t h e r a t e of l i v i n g f o r
the fly. H e a t h a s t e n s it. O n e g a t h e r s , f r o m t h e a c c o u n t
which L o e b a n d N o r t h r o p give of t h e w o r k , t h a t a t low
t e m p e r a t u r e t h e flies a r e s l u g g i s h a n d i n a c t i v e i n all
14
210 BIOLOGY OF D E A T H
t h r e e d e v e l o p m e n t a l s t a g e s a n d p e r h a p s live a l o n g time
because t h e y live slowly. A t h i g h t e m p e r a t u r e s , o n the
o t h e r h a n d , t h e fly is v e r y a c t i v e a n d l i v e s its life t h r o u g h
quickly a t the p a c e t h a t k i l l s . " T h e s e r e s u l t s a r e exactly
c o m p a r a b l e to t h e effect of a r e g u l a r i n c r e a s e of t e m p e r a -
t u r e upon a chemical r e a c t i o n . I n d e e d , L o e b a n d N o r t h -
r o p consider t h a t t h e i r r e s u l t s p r o v e t h a t
With a supply of proper and adequate food the duration of the larval
stage is an unequivocal function of the temperature at which the larvae are
raised, and the temperature coefficient i of the order of magnitude of that
of a chemical reaction, i. e.> about 2 or more for a difference of 10 C. It
increases at the lower and is less at the higher temperature**. The duration
of the pupal stage of the fly in also an unequivocal function of the tempera-
ture and the temperature coefficient is for each temperature practically
identical with that for the larval stage. The duration of life of the imago
is, with proper food, also an unequivocal function of the temperature and
the temperature coefficient for the duration of life is, within the normal
temperature limits, approximately identical with that for the duration of
life of the larva and pupa.

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

A.ge in months Rat No. 1 No. 4 No. 2 No. 3

at death Miles Miles
Miles Miles
25 1265
26 1391
32... 2098
34 5447
a g e . " F r o m t h i s e x p e r i m e n t it clearly a p p e a r s t h a t t h e
g r e a t e r t h e t q t a l w o r k d o n e , o r total e n e r g y o u t p u t , t h e
s h o r t e r t h e d u r a t i o n of life, a n d vice versa. Or, p u t
a n o t h e r w a y , if t h e t o t a l activity p e r u n i t of t i m e i s in-
c r e a s e d b y some m e a n s o t h e r t h a n i n c r e a s i n g t e m p e r a -
t u r e , the same r e s u l t s a p p e a r a s if the i n c r e a s e d a c t i v i t y
i s caused b y i n c r e a s e d t e m p e r a t u r e . I t a p p e a r s , i n
s h o r t , to be a c t i v i t y per $e, and not the t e m p e r a t u r e per
se t h a t is of r e a l significance. There i s o t h e r evidence,
f o r which s p a c e lacks h e r e , p o i n t i n g i n t h e s a m e direction,
A n entirely different, a n d extremely s u g g e s t i v e l i n e
of evidence i n f a v o r of t h e v i e w h e r e set f o r t h , lias "been
g i v e n b y P r o f e s s o r M a x B u h n e r , t h e d i s t i n g u i s h e d G-er-
m a n s t u d e n t of t h e e n e r g y relations of the l i v i n g o r g a n -
i s m . S t u d y i n g a c o n s i d e r a b l e r a n g e of a n i m a l s , h e lias
f o u n d t h a t all t r a n s f o r m n e a r l y t h e s a m e t o t a l a m o u n t
of energy, per kilo of tody weight, in t h e w h o l e p e r i o d
f r o m t h e i r b i r t h to t h e i r n a t u r a l death. T h e m e a n v a l u e
of t h e c o n s t a n t R u b n e r finds t o be 191,600 calories, t h e
v a l u e s for different species r a n g i n g between 141,090 a n d
214 BIOLOGY OF D E A T H
265,500 calories. S m a l l animals, w i t h a n i n t e n s i v e m e t a -
bolism live a r e l a t i v e l y s h o r t t i m e ; l a r g e animals with
m o r e sluggish m e t a b o l i s m live a l o n g e r t i m e . B u b n e r ' s
view is t h a t a definite s u m of living action ( e n e r g y t r a n s -
f o r m a t i o n ) d e t e r m i n e s t h e physiological end of life.
T h i s is p r e c i s e l y t h e view suggested h e r e except t h a t it
is h e r e p o s t u l a t e d t h a t t h e definite sum, f o r individual
or species, is f u n d a m e n t a l l y d e t e r m i n e d b y heredity,
working through the structural make-up.
I f we m a y be p e r m i t t e d to m a k e a s u g g e s t i o n r e g a r d -
i n g the i n t e r p r e t a t i o n of L o e b a n d N o r t h r o p ' s r e s u l t s in
conjunction w i t h o u r own on Drosophila, i t would be to
this effect. A n y g i v e n genetically p u r e s t r a i n of Droso-
phila is m a d e u p of individual machines, constructed to
t u r n out, b e f o r e b r e a k i n g down, a definite limited a m o u n t
of energy i n t h e f o r m of work, mechanical, chemical and
other. T h i s definitely limited t o t a l e n e r g y o u t p u t is
p r e 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 constitution of t h e indi-
vidual which fixes t h e kind of physico-chemical m a c h i n e
t h a t t h a t i n d i v i d u a l is. B u t the rate p e r u n i t of time of
the energy o u t p u t m a y be influenced between wide limits
b y e n v i r o n m e n t a l circumstances i n g e n e r a l a n d t e m p e r a -
t u r e in p a r t i c u l a r , since increased t e m p e r a t u r e i n c r e a s e s
r a t e of metabolic chemical changes i n a b o u t t h e s a m e
r a t i o , as d e m o n s t r a t e d by a w e a l t h of w o r k on t e m p e r a -
t u r e coefficients, a s i t i n c r e a s e s o t h e r chemical changes.
B u t if t h e r a t e of e n e r g y o u t p u t p e r u n i t of time is changed,
the total t i m e t a k e n f o r t h e t o t a l o u t p u t of a p r e d e t e r -
mined a m o u n t of e n e r g y , a s work, m u s t c h a n g e i n i n v e r s e
p r o p o r t i o n t o t h e c h a n g e of r a t e . S o w e should expect
j u s t precisely t h e r e s u l t s on d u r a t i o n of life t h a t Loeb
a n d N o r t h r o p got, a n d so f a r from t h e s e r e s u l t s b e i n g in
contradiction to o u r s u p o n heredity, t h e y m a y be looked
 STUDIES ON T H E DURATION OF LIFE 215
u p o n a s a n e c e s s a r y consequence of t h e m . L o e b a n d
N o r t h r o p ' s final conclusion i s : " T h e o b s e r v a t i o n s on t h e
t e m p e r a t u r e coefficient f o r t h e d u r a t i o n of life s u g g e s t
t h a t t h i s d u r a t i o n i s d e t e r m i n e d b y t h e p r o d u c t i o n of a
s u b s t a n c e l e a d i n g to old a g e a n d n a t u r a l d e a t h , o r b y t h e
d e s t r u c t i o n of a s u b s t a n c e o r substances, w h i c h n o r m a l l y
p r e v e n t old a g e a n d n a t u r a l d e a t h . " T h e view w h i c h I
h a v e h e r e suggested, completely i n c o r p o r a t e s t h i s view
within itself, if w e s u p p o s e t h a t t h e t o t a l a m o u n t of h y p o -
thetical " s u b s t a n c e o r s u b s t a n c e s which n o r m a l l y p r e v e n t
old a g e a n d n a t u r a l d e a t h " w a s essentially d e t e r m i n e d
by heredity.
T h i s view I t a k e t o be i n n o w a y n e c e s s a r i l y o r f u n d a -
m e n t a l l y c o n t r a d i c t o r y t o t h a t s e t f o r t h i n this w o r k .
"Whatever t h e f a c t o r w h i c h d e t e r m i n e s specific longev-
i t y m a y b e ; w h e t h e r a specific chemical s u b s t a n c e , as
L o e b a n d N o r t h r o p suggest, o r m o r e generally, a s I h a v e
suggested, the kind of m a t e r i a l , i n t h e sense of i t s biologi-
cal fitness, c o m p o s i n g t h e m u l t i c e l l u l a r body, a n d t h e
n a t u r e of t h e o r g a n i z a t i o n ( i n detail) of t h a t m a t e r i a l
to f o r m t h e m u l t i c e l l u l a r b o d y ; i t seems to m e t h a t w e
h a v e n o w a sufficient m a s s of critical evidence t o s a y
t h a t i t is p r o v e d t h a t q u a n t i t a t i v e l y t h e effective m a g n i -
t u d e of this specific l o n g e v i t y f a c t o r i n e a c h p a r t i c u l a r
case is determined hy heredity. T h i s I t a k e t o be of
g r e a t e r i m p o r t a n c e t h a n t h e p r e c i s e n a t u r e of t h e specific
longevity f a c t o r itself, a b o u t which we a r e , a d m i t t e d l y ,
entirely i g n o r a n t . I c a n see n o t h i n g i n t h e a v a i l a b l e evi-
dence which definitely m a k e s L o e b ' s s u g g e s t i o n i n h e r e n t l y
m o r e p r o b a b l e t h a n m i n e . I t does, h o w e v e r , s e e m clear
t h a t , b y definitely s h o w i n g t h e significance of t h e h e r e d i t y
element i n t h e p r o b l e m , h e l p h a s been r e n d e r e d t h e p r o g -
r e s s of f u t u r e r e s e a r c h i n t h e field.
216 BIOLOGY OF DEATH
I t would seem, a t first t h o u g h t , t h a t one should be
able t o t e s t t h e t h e o r y h e r e s u g g e s t e d , t h a t r a t e of e n e r g y
e x p e n d i t u r e i n t h e b u s i n e s s of l i v i n g i s n e g a t i v e l y c o r r e -
l a t e d w i t h the t o t a l d u r a t i o n of life, b y a n e x a m i n a t i o n of
t h e m o r t a l i t y r a t e s f o r p e r s o n s i n different occupations
a s set forth, f o r e x a m p l e , i n t h e well k n o w n p a p e r of
Bertillon. W h e n one e n d e a v o r s t o m a k e such a test,
however, h e i s a t once c o n f r o n t e d w i t h a s e r i e s of diffi-
culties which p r e s e n t l y convince h i m t h a t t h e p r o j e c t
is v i r t u a l l y a n i m p o s s i b l e one, if h e w i s h e s critical r e s u l t s .
I n t h e first place, m e a n a g e a t d e a t h will n o t do as a
criterion, b e c a u s e of t h e g r e a t differences i n t h e a g e dis-
t r i b u t i o n s of t h o s e e n g a g e d i n different occupations.
This p o i n t h a s l a t e l y b e e n t h o r o u g h l y discussed by Collis
and Greenwood, i n t h e i r book " T h e H e a l t h of t h e I n d u s -
t r i a l W o r k e r . ' 9 I n d e e d , t h e i r w h o l e t r e a t m e n t of t h e p r o b -
lem of occupational m o r t a l i t y is b y f a r t h e m o s t sound
a n d critical w h i c h t h e p r e s e n t w r i t e r h a s y e t seen. One
m u s t deal w i t h a g e a n d s e x specific d e a t h r a t e s , o r mor-
t a l i t y indices b a s e d u p o n them-
I n t h e second place, t h e r e a r e specific h a z a r d s , direct
or indirect, i n v a r i o u s occupations, q u i t e a p a r t f r o m a n y
question of e n e r g y e x p e n d i t u r e involved i n t h e case.
T h e s e h a z a r d s will, obviously, t e n d t o obscure a n y direct
effects of t h e e n e r g y r e l a t i o n s involved.
I n t h e t h i r d place, w e haVe only t h e m e r e s t suggestion
of q u a n t i t a t i v e l y a c c u r a t e k n o w l e d g e as t o t h e a v e r a g e
e n e r g y o u t p u t involved i n d i f f e r e n t t r a d e s a n d occupations.
On the l a s t p o i n t , a b e g i n n i n g t o collect i n f o r m a t i o n
h a s been m a d e b y W a l l e r a n d Ms co-workers. I n a r e -
cent p a p e r W a l l e r a n d D e D e c k e r h a v e g i v e n t h e m e a n
calory output, p e r h o u r , p e r s q u a r e m e t e r of b o d y surface
for a small s a m p l e of w o r k e r s i n a f e w t r a d e s . B u t t h e r e -
 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 217
suits a r e f a r too m e a g e r , and, statistically, too u n r e p r e -
s e n t a t i v e t o w a r r a n t any a t t e m p t a t g e n e r a l i z a t i o n f r o m
the p r e s e n t p o i n t of -view.
A s in so m a n y o t h e r cases t h e e x p e r i m e n t a l m e t h o d i s
likely to s h e d f a r m o r e critical light on this p r o b l e m t h a n
is the p u r e l y s t a t i s t i c a l m e t h o d d e a l i n g w i t h h u m a n d a t a .
T h e r e a r e too m a n y factors in the latter material t h a t
c a n n o t be controlled.

GO1TADS AND DTJBAriON OF XIFB

T h e r e is a n o t h e r a n d q u i t e different line of e x p e r i -
m e n t a l work on t h e d u r a t i o n of life "which m a y b e t o u c h e d
u p o n "briefly. T h e daily p r e s s h a s lately h a d a g r e a t d e a l
t o s a y a b o u t r e j u v e n a t i o n , accomplished "by m e a n s of
v a r i o u s s u r g i c a l procedures, u n d e r t a k e n u p o n t h e p r i m a r y
sex o r g a n s , p a r t i c u l a r l y in t h e m a l e . T h i s n e w s p a p e r
n o t o r i e t y h a s especially c e n t e r e d a b o u t t h e w o r t of
Yoronoflf and Steiixa^h. T h e only e x p e r i m e n t s which, a t
t h e p r e s e n t t i m e , p r o b a b l y d e s e r v e serious c o n s i d e r a t i o n
a r e t h o s e of Steinach. H e h a s w o r k e d chiefly w i t h -white
r a t s . H i s t h e o r y is t h a t , "by c a u s i n g t h r o u g h a p p r o p r i a t e
o p e r a t i v e p r o c e d u r e , a n extensive r e g e n e r a t i o n , i n a sen-
ile a n i t a a ! a b o u t t o d i e , of c e r t a i n g l a n d u l a r elements of
t h e testis, senility and n a t u r a l d e a t h will f a r a time be
p o s t p o n e d because of the- i n t e r n a l secretion p o u r e d i n t o
the blood by the r e g e n e r a t e d "puberty glands " a s h e calls
t h e m . T h e o p e r a t i o n w h i c h h e finds to- "be m o s t effective
is to l i g a t e firmly t h e efferent d u e t of the testis, t h r o u g h
which the s p e r m n o r m a l l y p a s s , close Tip t o t h e testis
itself, a n d before t h e coiled p o r t i o n of t h e d u c t is r e a c h e d .
T h e r e s u l t of t h i s , a c c o r d i n g to S t e i n a c h ' s account, is to
b r i n g a b o u t in h i g h l y senile animals a g r e a t e n l a r g e m e n t
of all the, sex o r g a n s , a r e t u r n of s e x u a l a c t i v i t y , p r e v i o u s l y
218 BIOLOGY OF D E A T H
lost t h r o u g h old age, and a g e n e r a l loss of senile bodily
characteristics a n d a r e s u m p t i o n of t h e conditions of
full a d u l t vigor in those r e s p e c t s , t o g e t h e r w i t h a consid-
erable i n c r e a s e in the t o t a l d u r a t i o n of life.
S p a c e is lacking to go into t h e m a n y details of
S t e i n a c h ' s work, m u c h of which is indeed chiefly of inter-
est only to the technical biologist, a n d f r o m a wholly
different s t a n d p o i n t t h a n the p r e s e n t one. I should,
however, like to p r e s e n t one e x a m p l e f r o m his experi-
ments. A s control, a r a t was taken, i n t h e l a s t d e g r e e
senile. H e w a s 26 m o n t h s old when the e x p e r i m e n t be-
gan. H e w a s obviously emaciated, h a d lost m u c h of his
hair, p a r t i c u l a r l y on t h e back a n d h i n d q u a r t e r s . H e
w a s weak, inactive and drowsy, a s indicated by t h e fact
t h a t his eyes w e r e closed, a n d were, one infers from
Steinach, k e p t so m u c h of the t i m e .
A l i t t e r b r o t h e r of t h i s a n i m a l h a d t h e efferent ducts
of t h e t e s t e s ligated. T h i s animal, we a r e told, w a s , a t
the t i m e of t h e operation, in so m u c h w o r s e condition of
senility t h a n his b r o t h e r , above described, t h a t i t w a s n o t
t h o u g h t w o r t h while even to p h o t o g r a p h him. H i s con-
dition w a s considered hopeless. T o the s u r p r i s e of t h e
operator, however, he came back, slowly b u t s u r e l y a f t e r
the operation, a n d a f t e r t h r e e a n d a half m o n t h s p r e -
sented a perfect p i c t u r e of l u s t y y o u n g r a t h o o d . H e
was i n full vigor of e v e r y sort, including sexual. H e
outlived his b r o t h e r by 8 m o n t h s , a n d himself lived 10
months after t h e operation, a t which t i m e h e w a s , accord-
ing to Steinach, practically m o r i b u n d . T h i s r e p r e s e n t s a
p r e s u m p t i v e lengthening of h i s expected s p a n of life b y
roughly a q u a r t e r t o a t h i r d . It is to he remembered,
however, that Slonaker's rats to which nothing was done
lived to an average age of 40 months.
 S T U D I E S ON T H E D U R A T I O N OF L I F E 219
* presumption that Steinach's experiments have
b r o u g h t a b o u t a s t a t i s t i c a l l y significant l e n g t h e n -
l i f e is l a r g e , a n d t h e b a s i s of a s c e r t a i n e d fact
A f t e r a careful e x a m i n a t i o n of S t e i n a c h ' s b r i l -
oxLtribution, one is compelled t o t a k e t h e v i e w t h a t ,
s i r i n t e r e s t i n g t h e r e s u l t s m a y be f r o m t h e s t a n d -
o f functional rejuvenation in the sexual sphere,
3 e i s n o t p r o v e n t h a t a n y r e a l l y significant l e n g t h -
o f t h e life s p a n h a s occurred. I n o r d e r t o p r o v e
l e n g t h e n i n g we m u s t , first of all, h a v e a b u n d a n t a n d
<fce q u a n t i t a t i v e d a t a a s t o t h e n o r m a l v a r i a t i o n of
1 r a t s in r e s p e c t of d u r a t i o n of life, a n d t h e n show,
; r e g a r d t o t h e p r o b a b l e e r r o r s involved, t h a t t h e
i x i r a t i o n of life a f t e r t h e o p e r a t i o n h a s b e e n signi-
y l e n g t h e n e d . T h i s S t e i n a c h does n o t do. H i s
I s s i n g u l a r l y b a r e of s t a t i s t i c a l d a t a . W e m a y well
a d e q u a t e q u a n t i t a t i v e evidence before a t t e m p t i n g
^xxeral i n t e r p r e t a t i o n of h i s r e s u l t s ,
l e e d , one m a y n o t e i n p a s s i n g t h a t t h e case does
e m entirely c l e a r i n r e s p e c t of S t e i n a c h ' s r e s u l t s
; p n r e l y sexual sphere, Thus Romeis has repeated
: p e r i m e n t s , a n d finds, f r o m c o m p a r a t i v e histologi-
i d i e s on t h e g e n i t a l o r g a n s of r a t s , b e f o r e a n d after
's o p e r a t i o n , t h a t t h e r e i s no evidence of a n y
in L e y d i g ' s i n t e r s t i t i a l cells, a n d hence n o n e
s o - c a l l e d i'interstitial or puberty gland.'' Romeis
XLO increase i n s e x u a l d e s i r e a m o n g h i s r a t s a f t e r
> e r a t i o n . T h e h y p e r t r o p h y of t h e s e m i n a l vesicles
x r o s t a t e , d e s c r i b e d b y S t e i n a c h following t h e o p e r a -
v a s also s e e n b y R o m e i s , b u t found, b y t h e l a t t e r , t o
s r e l y the r e s u l t of t h e s t a s i s of t h e secretions nec-
L l y consequent upon the operation, and not a t r u e
L o n a l h y p e r t r o p h y a t all.
220 BIOLOGY OF D E A T H
THE PITUITABY GLAND AND DURATION OF LIFE
R o b e r t s o n h a s b e e n e n g a g e d f o r a n u m b e r of y e a r s
p a s t on a n extensive s e r i e s of e x p e r i m e n t s r e g a r d i n g t h e
effect of v a r i o u s a g e n t s u p o n t h e g r o w t h of w h i t e m i c e .
T h e e x p e r i m e n t s h a v e been c o n d u c t e d w i t h g r e a t c a r e
a n d a t t e n t i o n t o t h e p r o p e r h u s b a n d r y of t h e a n i m a l s .
I n consequence, t h e r e s u l t s h a v e a h i g h d e g r e e of t r u s t -
w o r t h i n e s s . I n t h e c o u r s e of t h e s e s t u d i e s h e f o u n d t h a t
t h e a n t e r i o r lobe of t h e p i t u i t a r y body, a small g l a n d a t
t h e b a s e of t h e b r a i n , n o r m a l l y s e c r e t e s i n t o t h e blood-
s t r e a m m i n u t e a m o u n t s of a n active s u b s t a n c e w h i c h h a s
a m a r k e d effect u p o n t h e n o r m a l r a t e of g r o w t h . B y c h e m i -
cal m e a n s , R o b e r t s o n w a s able t o e x t r a c t t h i s a c t i v e s u b -
stance f r o m t h e g l a n d i n a f a i r l y p u r e s t a t e , a n d g a v e to it
the n a m e tethelin. I n l a t e r e x p e r i m e n t s , t h e effect of
tethelin, g i v e n by t h e m o u t h w i t h t h e food, w a s t r i e d in
a v a r i e t y of w a y s .
I n a recent paper, Robertson and R a y have studied
t h e effect of t h i s m a t e r i a l u p o n t h e d u r a t i o n of l i f e of t h e
w h i t e m o u s e w i t h t h e r e s u l t s shown in. T a b l e 27,
TABLE 27
Effect of tethelin on duration of life in days of white mice,
(Robertson and Ray)
Both
MALES FEMALES mxm
toother
Chance Chance Chance
Cl&8B0iAverage Dv. Dov. 6&Y. WA Avoragft
H Dv. dav. WAS dav. was
duration from duration from acciden- acci-
animals of life normal P. B. Acciden- of life normal P.M. tal dental
tal
Normal 767 719
Tethelin 866 +99 3.00 1:22.25 800 f81 2.25 1:6.75 1:150.2
P r o m t h i s table, it i s a p p a r e n t t h a t t h e a d m i n i s t r a t i o n
of t e t h e l i n w i t h t h e food f r o m b i r t h t o d e a t h p r o l o n g e d
 STUDIES ON T H E DURATION OF LIFE 221
life to a d e g r e e which, in t h e case of t h e m a l e s , m a y be
r e g a r d e d a s p r o b a b l y significant statistically. I n t h e
case of t h e females, w h e r e t h e r a t i o of t h e d e v i a t i o n to
its p r o b a b l e e r r o r ( D e v . / P . E . ) falls t o 2.25 t h e case
is v e r y doubtful. T h e p r o c e d u r e by which t h e chance of
1:150.2 t h a t r e s u l t s i n b o t h sexes t o g e t h e r w e r e acciden-
tal, w a s obtained is of doubtful validity. P u t t i n g m a l e s
a n d females t o g e t h e r f r o m t h e original table, I find t h e
following r e s u l t s .
TABLE 28
Duration of life of white mice, both sexes taken together
(From data of Robertson and Ray)
No.of No. of deaths
Age deaths of tethelin
Group of normals fed
(Both sexes) (Both sexes)
200-299 3 Tethelin fed: Mean age at death =839=b20
300-399 2 Normal fed: Mean age at death =74317
400-499 2 1 Difference == 96db26
500-599 9 3
600-699 7 9 Difference = 3.7
PE
700-799 15 Diff.
800-899 10 10
900-999 10 6
1000-1099 6 9
1100-1199 1
64 39
One concludes f r o m t h e s e figures t h a t t e t h e l i n c a n be
r e g a r d e d as h a v i n g l e n g t h e n e d t h e s p a n of life t o a de-
g r e e which is j u s t significant statistically. One would
expect, f r o m t h e v a r i a t i o n of r a n d o m s a m p l i n g alone, t o
get a s d i v e r g e n t r e s u l t s a s t h e s e a b o u t 1*4 t i m e s i n e v e r y
100 t r i a l s w i t h s a m p l e s of 64 a n d 39, r e s p e c t i v e l y .
I n a n y event i t is a p p a r e n t t h a t , m a k i n g o u t t h e b e s t
case possible, t h e differences i n a v e r a g e d u r a t i o n of life
222 BIOLOGY OF DEATH
p r o d u c e d by a d m i n i s t r a t i o i i of t c t h c l i n a r e of a wholly
different a n d s m a l l e r o r d e r t h a n t h o s e which h a v e been
shown, in t h e e a r l i e r p o r t i o n of t h i s c h a p t e r , t o exist be-
t w e e n p u r e s t r a i n s of DrosophUa which a r e baned u p o n
h e r e d i t a r y differences.
P u t t i n g t o g e t h e r all the r e s u l t s which h a v e been r e -
viewed 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 , it a p p e a r s t o
be clearly a n d firmly established t h a t i n h e r i t a n c e in t h e
f a c t o r of p r i m e i m p o r t a n c e in d e t e r m i n i n g t h e n o r m a l ,
n a t u r a l d u r a t i o n of life. I n c o m p a r i s o n with thin factor,
t h e influence of e n v i r o n m e n t a l forces (of fmb-lftthal im-
m e d i a t e i n t e n s i t y of c o u r s e ) a p p e a r s in g e n e r a l to be
less m a r k e d .
 CHAPTER VIII

NATURAL DEATH, PUBLIC HEALTH, AND THE

POPULATION PROBLEM.

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 togetherthe 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-plasmthe innate
inherited constitutional make-upof 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 superiorby our present standardsto 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

CONTROLLABLE CAUSES OF DEATH

IflOO r==

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 deathnamely:
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

NON - CONTROLLED CAUSES OF DEATH

1,000

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 FVR

^ *
\
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.

the role of i m m e d i a t e a n d l a r g e i m p o r t a n c e w h i c h the

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 -

standing, let m e s a y q u i t e s p e c i f i c a l l y a n d categorically,

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

i d e a that public h e a l t h w o r k i s n o t desirable, or t h a t a

 NATURAL DEATH, PUBLIC HEALTH 237
laissez-faire policy would be better, or that public health
efforts have not been enormously valuable in connection
with typhoid fever and diphtheria. My purpose is quite
other, being solely a desire to emphasize two things, viz:
1. That the trend of human mortality in time is an

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 generalto 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* mosquitoesA. quadrimaculatus Say,
A. punctipenms Say, and A. oruzians Wiedermannwhich 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, 1910The 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.

THE POPULATION* PBOBLEM

T u r n i n g to a n o t h e r p h a s e o f t h e p r o b l e m , i t is a p p a r -
ent t h a t if, a s a r e s u l t o f s a n i t a r y a n d h y g i e n i c activi-
ties a n d n a t u r a l e v o l u t i o n , t h e a v e r a g e d u r a t i o n of
h u m a n life is g r e a t e r n o w t h a n i t "used t o be a n d is g e t t i n g
g r e a t e r all t h e time, t h e n c l e a r l y t h e r e m u s t be m o r e
people on t h e e a r t h a t a n y t i m e , o u t of a g i v e n n u m b e r
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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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173 A \

450
/
f
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125
' A
100
/ Af if*

75

50.

1
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1
^n! change in percentage which deaths were of births in each of the
>;<*

B u t in 1919, it p r o m p t l y d r o p p e d back to the n o r m a l value

of 57.98, a l m o s t identical w i t h the 1917 figure of 57.34.
In England and Wales, the provisional figure indi-
cates that 1920 will show a lower value for the vital
index than that country has had for m a n y years.
So we see that neither a highly destructive war, nor
the most destructive epidemic since the Middle Ages,
serves more than to cause a momentary hesitation in the
steady onward march of population growth.
 NATURAL DEATH, PUBLIC HEALTH 247
T h e first t h i n g obviously n e e d e d i n a n y scientific
a p p r o a c h to t h e p r o b l e m of p o p u l a t i o n is a p r o p e r m a t h e -
m a t i c a l d e t e r m i n a t i o n a n d e x p r e s s i o n of t h e l a w of p o p u -
l a t i o n g r o w t h . I t h a s b e e n seen) t h a t t h e m o s t d e v a s t a t i n g
c a l a m i t i e s m a k e b u t a m o m e n t a r y flicker in the s t e a d y
p r o g r e s s of t h e c u r v e . F u r t h e r m o r e , p o p u l a t i o n g r o w t h
is p l a i n l y a biological m a t t e r . I t d e p e n d s upon, in l a s t
a n a l y s i s , only the b a s i c biological p h e n o m e n a of f e r t i l i t y
a n d m o r t a l i t y . T o t h e p r o b l e m of a n a d e q u a t e 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 n o r m a l g r o w t h of p o p u l a t i o n s ,
m y colleague, D r . L o w e l l J . R e e d , a n d I h a v e a d d r e s s e d
o u r s e l v e s f o r some t i m e p a s t . T h e k n o w n d a t a u p o n which
w e h a v e to o p e r a t e a r e t h e p o p u l a t i o n c o u n t s given b y
successive c e n s u s e s . V a r i o u s a t t e m p t s h a v e been m a d e
i n t h e p a s t t o g e t a m a t h e m a t i c a l r e p r e s e n t a t i o n of t h e s e
i n o r d e r to p r e d i c t successfully f u t u r e p o p u l a t i o n s , a n d
t o g e t e s t i m a t e s of t h e p o p u l a t i o n i n inter-censal y e a r s .
A n o t e w o r t h y a t t e m p t of t h i s s o r t is P r i t c h e t t ' s fitting
of a p a r a b o l a of the t h i r d o r d e r to t h e U n i t e d S t a t e s p o p u -
l a t i o n f r o m 1790 t o 1880 inclusive. T h i s g a v e a f a i r l y
g o o d r e s u l t o v e r t h e p e r i o d , b u t w a s obviously p u r e l y
e m p i r i c a l , e x p r e s s e d n o r e a l biological l a w of change,
a n d i n f a c t failed b a d l y i n p r e d i c t i o n a f t e r 1890.
W e h a v e a p p r o a c h e d t h e p r o b l e m f r o m a n a priori
basis, set u p a hypothesis as to the more important
biological f a c t o r s involved, a n d t e s t e d t h e r e s u l t i n g
e q u a t i o n a g a i n s t t h e facts f o r a v a r i e t y of c o u n t r i e s .
T h e h y p o t h e s i s w a s b u i l t u p a r o u n d t h e following
considerations:
1. I n a n y g i v e n l a n d a r e a of fixed limits, as b y
political o r n a t u r a l b o u n d a r i e s , t h e r e m u s t n e c e s s a r i l y be
a n u p p e r l i m i t t o t h e n u m b e r of p e r s o n s t h a t can be s u p -
p o r t e d on t h e a r e a . T o t a k e a n e x t r e m e c a s e , it is obvious
248 BIOLOGY OF D E A T H
that not so many as 25,000 persons could possibly stand
upon an acre of ground, let alone live on it. So, similarly,
there must be for any area an upper liiniting number of
persons who can possibly live upon it. I n mathematical
terms this means that the population curve munt have
an upper limiting asymptote.
2. A t some time i n the more or less remote past the
population of human beings upon any given land area
must have been nearly or quite zero. S o the curve m u s t
have somewhere a lower limiting asymptote.
3. Between these two levels we assume that the rate
of growth of the population, that is, the increase in
numbers in any given time unit, is proportional to two
things, namely:
a. The absolute amount of growth (or size of population) already
attained;
b. The amount of as yet unutilized, or reserve, means or aourcos of
subsistence still available in the area to support further
population.
These hypotheses lead directly to a curve of the form
shown i n F i g u r e 60, i n which the position of the asymp-
totes and of the point of inflection, when the population
is growing at the most rapid rate, are shown in terms
of the constants. I t i s seen that the whole history of a
population, a s pictured by this curve, is something like
t h i s : I n the early y e a r s following the settlement of a
country the population g r o w t h i s slow. P r e s e n t l y it
begins to grow faster. After i t p a s s e s the point where
half the available resources of subsistence have been
drawn upon and utilized, the rate of growth becomes
slower, until finally the maximum population wMch the
area will support i s reached.
 NATURAL DEATH, PUBLIC HEALTH 249
This theory* of population growth makes it possible
to predict what the maximum population in a given area
will be, and when it will be attained. Furthermore, one
can tell exactly when the population is growing at the
maximum rate. To test the theory, we have only to fit

Fio. 60.Showing a theoretical curve of population growth.

this theoretical curve to the known facts of population
for any country by appropriate mathematical methods.
If the hypothesis fits well all the known facts for a variety
of countries in different stages of population growth, it
may well be regarded as a first approximation to a sub-
stantially correct hypothesis and expressive of the bio-
logical law according to which population grows. In
making this test the statistician has somewhat the same
* The mathematical hypothesis here dealt with is essentially the same
as that of Verhulst, put forth in 1844. As Pearl and Reed pointed out
in theirfirstpaper on the subject it is a special case of a much more
general law. A comprehensive general treatment of the problem we are
publishing shortly in another place. The generalization in no way alters
the conclusions drawn here from a few illustrative examples.
250 BIOLOGY OF D E A T H
kind of p r o b l e m t h a t c o n f r o n t s t h e a s t r o n o m e r calculat-
i n g t h e complete o r b i t of a comet. T h e a s t r o n o m e r n e v e r
h a s m o r e t h a n a r e l a t i v e l y few o b s e r v a t i o n s of t h e posi-

175
UNITED STATE'S
150
115
100
76
60
Z3
1 I _.LJ_
/7OO 20 4O 60 80 J6OO JX>4OO 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

MOO O 4-0 60 6O J7O0 O <*O 6O dO IQOO 2D AO 6O 60 J900 20 40 6O 6O 0OO

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

/
SERW\
/
A
/
e

1
A
\ 1 J...J-4--<*\ I
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.
JZ5
/
era /
m

TH or OH 7SOPH0.A
ORCMT
/WUATION J
tzs 7
5 too /
j
/
/
f
73
SO
*
0 6 / M &I JO 4 t n
OCT. NO*
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 -

lation p r o b l e m of the U n i t e d S t a t e s i n the light of

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

d i d o r will the p o p u l a t i o n c u r v e of this c o u n t r y p a s s t h e

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 -

m e r i c a l values, the U n i t e d S t a t e s h a s a l r e a d y p a s s e d its

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.
 BIBLIOGRAPHY
The following list of literature in no sense aims at completeness
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250
 260 BIBLIOGRAPHY
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 BIBLIOGRAPHY 261
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 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, <$3
Conklin, E. G., 29, 44, 260 Donaldson, H. II, 20, tm
Controllable causes of death, 230,233 Drosophila mtUinopaMter, 180*202,
Correlation coefficient, 168 208-211, 214, 222, 225, 22S, Z&%,
Correlations in duration of life, 168- 2f>4
177 Dublin, U I., 82, 113, 260, 261
Crossett, 239, 241 du Noy, l\ L., 77
Croup, 230, 231 Duration of life, corn*!ntion in, 16B-
Crura, F. 8., 184, 260 177
Culture of tissues in vitro, 58-78
Curve, mortality, graduation of, 94- of, im^tt
101 \nfimtm* of activi-
specific death rate, 114, lltt ty mi, 2U-217
Curves, logarithmic plotting of, 114 of t-ni
Cyclops, 39 t on, 20B-
Cytomorphosis, 28 217
Cytoplasm, 29, 44 inh<ritanci of, M,
Darwin, L., 257 imim
Davis, W. H., 113 In man, 7D-04, 150.
Dawson, J. A., 73, 260 185
 INDEX 271
Duration of life of domestic fowl, 63 Expectation of life, effect of selection
of parents and off- on, 94
spring, 155-157 hypothetical, 164
rOle of bacteria in, in ancient Egypt,
43, 199-202 87-89
variation in, 21, 22, in ancient Home,
68, 80-82 90-92
Dysentery, 230, 231 in Hispania and
LuHitania, 91-
East, E. M., 195, 257, 261 92
Ebeling, A. 1L, 60, 61, 74, 76, 77, 78, in Roman
260, 261 Africa, 92-93
Ectoderm, 138-149 Experimental atudy of duration of
Effects of public health work, 112, life, 186-222
227-242 Eye color, 174, 175
Egypt, expectation of life in, 87-89
Elephant, longevity of, 22 Ferxnat, 82
Embryology and mortality, 138-149 Fertiltain, 67
Embryonic juice, 74 Finn, longevity of, 22
Endocrinal system, 107, 108, 112, FiHher, A., 101, 149, 184, 261
133, 134 PiHher, I., 161, 162, 165
Endoderm, 138-149 Kiftiiion, 32, 33, 35, 40, 41
Endomixis, 30, 33, 71-73 Fitting the mortality curve, 94-101
Energy, 213-217 Food requirements, 256
England, 106, 108-111, 139, 140, 235, Fowyth, C. H., 161, 164, 261
244-246 Fowl, duration of life of, 63
Enriquea, P., 73, 261 France, 244, 245, 251, 252
Environment, 2Z$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 Ini*ctH, longvity 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 cdlf 210
Hersch, L., 202, 203, 205, 206, 208, In vertebra ten, longevity of, 22
262 In vitro culture of ti*ti<**, fH 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 Rigitration 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 ytem, 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, 20
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
HeneKcenee, 27-30, 4o\ 70-78
theorie* of, 43-50
Senile changes in nerve cells, 27-2D
Senility as eaune of death, 100
in plants, 44, 74, 75
Septicaemia, 231, 232
Serbia, 252, 253
Serum, influence on tiH*m* culture,
76, 77
Sex organs, 107, 108, 111, 121-125,
217-219
Sexual reproduction, 37-41
Shell, J., 26, 27
Skeletal nyHtem, 107, 108, 112, 127,
128
Skin, 107, 108, 110, 112, 131, 132
Klonaker, J. H., 212, 213, 218, 228,
267
Klotnpohtki, BM 33, 267
Snowf E. C, 179-183, 225, 267
Softening of th hrain, 231 , 232
Soma, 40
Somatic cells, immortality of, 58-78
Span, 174, 175
HpiegelWg, W.t 87
Spiritualmm, 18-20
Kple<?n, 61
Sponge*, 02
Stature, 174, 175
Htelnach, E., 217-210, 207
Btenoatomum, 35, 36
Btnvunnon, T. H. C., 206-208, 267
Still hirthn, 205
8trongylocentrotu$ purpuratus, 38,
56
Sttmmary of renultw, 223-227
gurvivornhip Hum of DromphHa,
188, 102, 195
Syphilis, 123
275
Table, life, 79-82
Temperature, 208-217
Tethelin, 70, 220-222
Theories of death, 43-50
Theory of population growth, 249
Thyroid gland, 61
TiHsue culture in vitro, 5H-78
Transplantation of tumors, 64, 05
TutmrcukmiB, 161, 204, 208, 230, 2:U,
238
Tumor tranHplaniation, 64, 65
Typhoid fever, 230, 231, 235, 2.'W
United States, growth of, 250-252,
254-257
Urostyla grantlis, 72
Van Buren, G. H., 113, 260
Variation, genetic, 190
Venereal diabases, 123, 124
VerhulHt, I\ F., 249, 267
Verworn, M., 44, 267
Vienna, 245, 246
Voronoff, 217
Waller, A. I)., 216, 267
Waiworth, H. H., 152, 267
War, 243
Wedekind, 33, 267
Weiamann, A., 26, 43, 65, 267
Whale, longevity of, 22
Wilton, H. V., 62, 267
Wittatein, 09
Woodruff, U L.t 30, 33, 72, 73, 267,
268
Woods, F. A., 38, 39, 268
Ydlow fever, 240, 242
Voting, T. E., 23-25, 268