I

~
-
,
c
Figure 1. The componentsof the f"'-.. !'.:II
natural selectionfollowed a certai hI, ;11...
I
Darwin'sthinking that led him to ."'il 11
lation of his theory,
."!tIH' r I
0., II,', \1
,.1" 'II ;

Inference 1 --"I'I""!
Struggle for existence lo~. \',"'II'~
among individuals k""",
II '
(Author of interence: Malthus)
.'" " '1\
",), ,,111
1 ::::1 Inference 2 Inference 3 ..,..1'11'
~""I'Ir;1

pifferential surviv~I, Throu9h many gene j'" .I,lf.lIl

: observation, . I.e. naturalselection .. evolution
(Author at inference: Darwin) (Author of inference: D 1""1 II II'
iI"., 1 II 'r
, 1, rl;ll:
.1, "" I ;)1
;."t".M
'.. ;i , I II (~
'.. I II'IY
... t II;I!
--.,..lo'r;.I,
"'1, \Vl
.."I f 111'1
II,I II III'~

could Darwin have arrived at an idea ber 1838, when the theory of natural struct the trials and errors of Darwin". 1111'J
which not only was totally at variance selection was conceived. gradual approach, as reflected in .11'""t
with the thinking of his own time but successive entrjes in the Notebooka. ,.tll,lls f
which was so complex that even now, One fact, the importance of which has It also examines tentative ideas that j,I";lls (
139 years later, it is widely misun- not been reduced by the recent dis- he later rejected. Gruber, Kohn, and. .i,l.. fli).
derstood in spite of our vastly greater coveries, is the impact of Darwin's in part, Limoges have favored thit ,"II:: Ila
understanding of the processes of reading of Malthus. The interpreta- method. I".r\"(~c
variation and inheritance? This is the tion of the Malthus episode, however, '.II. .,1 s a
question which the historian of ideas has become the subject of consider- What were the components of Dar. I., "llli
wouldlike to seeanswered
(4). ablecontroversy
amongthe Darwin win's explanatorymodel?For my 11'111lie
scholars. According to some of analysis I found it most convenient to '"1,0/ in
Darwin's own version (in his auto- them-e.g, de Beer and S. Smith, recognize five facts and three infer, 1,'f..rl'll(
biography) was that contemplation ~f and, to a lesser extent, H. Gruber and ences, as diagrammed in Figure 1.1 11'.11 illr
the success of animal breeders in myself-it was merely the culmina- shall attempt to determine, first, at I 11;lrw
producing new breeds had provided tion in the gradual development of what time Darwin became aware ot I'" II'S i
himwiththecluefor themechanismDarwin'sthinking,a little nudgethat thesefivefacts,thenat whattimeh. '\ I" ,I.Il{i.
of evolution and was thus the basis for pushed Darwin across a threshold at made the three inferences, and !!'" Ilec;
his theoryof naturalselection.We whichhe wasalreadystanding,Ac- whetheror not theseinferences
had rIlrll.ill (
knowthat this is a vast oversimplifi- cording to others-e.g. C, Limoges already been made previously and "'Ir,' Ilf'
cation-a revision of our thinking and D. Kohn-it constituted a rather could be found in the literature,
which we owe to the rediscovery of drastic break, almost equivalent to a \ shi1
Darwin'sNotebooks.In July 1837he religiousconversion.Which of these When we consider the five facts in 1"'II;111CI
had started to write down (in a series two interpretations is nearer to the Figure 1 carefully, it becomes appa,. I ',rl,in S
of notebooks) all the facts as well as truth? ent that all of them were widely ! '" ,I al

his own brain waves and speculations known. Not only had they been in I"l is, h
"which borein any wayon the varia- Thereareessentiallytwo methodsby Darwin's own handswell beforethe '" I" of
tion of animalsand plants under do- which we can try to find an answer. Malthus episode,but they had been "I,ly~
mestication and [in] nature." Even Either we can attempt to analyze all available to Darwin's contemporaries, ,I,k" oy
thoughhe later cut out occasionalthe entriesin the Notebooks,in a only a singleone of whom,A. R. Illr,lt:!eJ
pages, to use them for his book man- chronological sequence, or we can try Wallace, used them in exactly the i,I",il,lOj
uscripts, Darwin never discarded to reconstruct Darwin's explanatory same way as Darwin. Merely havin,. , ,11!"('cti
these notebooks, and they were model of natural selectionanq then thesefactsobviouslywasnot enough. ," Ille gl
rediscovered in the 19508 among the study separately the history of each of They had to be related to each other 1"r"llt t

.
Darwin papers at the Cambridge its individual components. Myown in a meaningful manner; that is, they """rne
University Library (5). Darwin's choice is in favor of the second had to be placed in an appropriate" '1l,th.
day-by-day records throw an entirely method, placed in a chronological conceptual background. In other' I,'r(' is B
newlightonthedevelopment
andthe framework,althoughboth methods words,Darwinhadto beintellectually i ,III 'ren.
changesin his thought during the arenecessary
for a full understanding, prepared to see the connection. f,,;,1ive,
period fr~m ~~IY ~837 to 28.Septem- The first method attempts to recon- among thes~~a:ts, c\t; "f~,::;~O~

:122 AmericanScIentist,Volume65 2~.- :~~

 "
:;~;

eory of ~fThis leadsus to the most interesting 3. A changing attitude toward the many of Darwin's favori~eand most
rder in but also the most difficult question: balanceof nature. This affectedtwo frequentlyr-eadauthors,lIke Erasmus
formu- What had beengoingon in Darwin's aspects;whetherthe balanceis purely Darwin,CharlesLyell, Alexandervon
'mind in the one and a half years prior static or, rather, dynamic, and Humboldt, and William Paley. Fur-
to the Malthus episode? All the indi- whether the balance is maintained by thermore, Malthus's principle was
cations are that it was a period of benign adjustments or by constant widely discussed in the essay litera-
unprecedentedintellectual activity in war. ture {)f the period. Why then did this
Darwin's life. Precisely what the suddenly impress Darwin so pro-
changes in Darwin's thinking were 4. A gradual loss of his Christian foundly on 28 September 1838?
and how they were connected with faith. It is apparent that Darwin lost
each other has not yet been investi- his faith in the years 1836-39, much Three reasons might be suggested-
gated nearly as fully as it deserves. of it clearly prior to the reading of the first one being, as pointed out by
Gruberand Kohn haveexaminedthis Malthus (7). In order not to hurt the Gruber(10),that Darwinhadlearned
'ations: problem more fully than anyone else, feelings of his friends and of his wife, on the three preceding days (between
'arwin) but the Darwin correspondence of Darwin often used deistic languagein ~eptember 2?.and 27) of the unbe-
that period and other manuscript his publications, but much in his llevable fertilIty of protozoans by
materials that have not yet been an- Notebooks indicates that by this time reading Ehrenberg's work on the
alyzed are bound to provide new in- he had become a "materialist" (more subject. This quite likely primed
sights. My own tentative conclusions or less = atheist). Darwin's receptivity for Malthus's
may, therefore, turn out to be incor- thesis. The second reason is that
rectoHowever, my reading suggeststo These four changes in Darwin's when Malthus applied the principle
me that Darwin's beliefs changed thinking are to some extent inter- to man, a specieswith relatively few
moderately or drastically in four connected. Since they were largely offspring, Darwin suddenly realized
areas,which I shall simply list here unconscious, they are usually re- that a potentially exponential in-
and then discuss in the context of flected in Darwin's Notebooks only in crease of a population was entirely
Darwin's model. subtle changesof wording, and there independent of the actual number of
is considerable leeway in possible in- offspring of a given pair. The third
arwin's 1. The gradual substitution of the terpretation. However, keeping these reason is that the Malthus episode
ted in concept of the uniqueness of indi- four points in mind will sharpen our came at a time when population
:books. viduals for one that assumesall indi- awareness of possible changes in thinking had begun to mature in
as that viduals of a speciesto be essentially Darwin's thinking in the years prior Darwin's mind. .
n, and, alike (6). This is the belief, ultimately to reading Malthus, while we make a
~d thi~ going back to Plato (eidos), that the point-by-point analysis of Darwin's The second fact in Figure 1-popu-
observed variability of phenomena explanatory model (Figure 1). lation stability-was not in the
reflectsa limited number of constant, slightest controversial. No one ques-
~f Dar esse~ces-:th~ant- The strug gle for existence
disconti!1uous tioned~hatthe numberof species
. or m! onym beIng "populatIon thInkIng," a and, asIde from temporary fluctua-
nient ill beliefin the importance of individual When recording his reaction to read- tions, the number of individuals in
e infer differences and the reality of the ing Malthus on 28 September 1838, every species maintained a steady-
ure I, I variation within a population. Most Darwin makes it quite clear that it state stability. This is implicit in the
first, 111 of Darwin's earlier statements on was not Malthus's general attitude conceptof plenitude of the Leibniz-
ware III" speciesand varieties were strictly that had acted as a catalyst on his ians and in the harmony-of-nature
time hi' typological. It is my impression that thoughts, but one particular sentence, concept of the natural theologians. If
!s, ami they becamemore "populational" as for he says that "yet until the one there is any extinction, it is balanced
ces hall Darwin delved deeper into the liter- sentence of Malthus no-one clearly by speciation, and if there is high
sly ami ature of the animal breeders. perceived the great check a"mongst fertility it must be counterbalanced
Jre. men" (8). De Beer succeeded in de- by mortality. In the end everything
2. A shift from soft toward hard in- termining what Malthus's crucial adds up u) a steady-state stability.
facts ill heritance. In his earlier statements sentence was: "It may safely be pro-
; app,lr Darwin seemedto assumethat most, nounced, therefore, that the popula- The third fact-limitation of re-
wid!:'I, if not all, inheritance was "soft"- tion, when unchecked, goes on dou- sources-again, was not at all con-
been ill that is, he assumedthat the material bling itself every twenty-five years, or troversial, being very much part of
fore tllc' basisof inheritance is not unchan- increases in a geometrical ratio" (9). the balance-of-nature concept of
ad bt:'('IJ geablyconstant but can be modified From then on, Darwin stressedthat it natural theology, so dominant in
;>orari('", alike by the inheritance of acquired was Malthus's demonstration of the England in the first half of the nine-
1, A. Ii characters("use and disuse," etc.), by exponential increase of populations teenth century.
ctly 1.11" physiologicalactivities of the body, by that was decisive in his discovery of
I' havil4: . direct influence of the environment the importance of natural selection Darwin's first great inference,derived
enou~11 on the genetic material, or by an in- ("Fact 1"). from these three facts, was that ex-
ch oth('r herent tendency to progress and to ponential population growth com-
tis, thl'\ become more perfect. With the Yet there is a puzzling difficulty. Why bined with a fixed amount of re-
ropri:ll, growth of his population thinking, did it take Darwin so long before sources would result in a fierce
n oth"1 thereis an increasingstresson genetic recognizing the evolutionary signifi- struggle for existence (Figure 1). We
lectuall\ differences among individuals, in- cance of the Malthusian principle? must ask, Was this inference original
necti<III" dicativeof a growing awarenessof the The prodigious fertility of animals with Darwin, and if so, what part of it
needtopostulate"hard"inheritance. and plants had been pointed out by did he owe to Malthus? This is per-

1977 May-June 323

~_L ;'c;i?

haps the most controversial question researches" (12). Lamarck expressed a species.This is the interpretation of, had the
raised by the analysis of the selection similar sentiments. the struggle for existence in the lit. (Ioes no
theory.The maindiffic~ty is that the ". . erature of natural theol°r;Y' up to .de evolutioJ
term "struggle for exIstence" and Was thIS benIgn Interpretation of the Candolle and Lyell, and ISthe major clevelop
similar synonymous terms were used struggle for existence unanimous? emphasis of Darwin's Notebooks up , mo 1
in different sensesby different au- Unfortunately,eventodaywehaveno to the Malthus reading. The main ;:oP~~ti
thors. reliableanalysisthat wouldgiveusan function of this struggleis to correct c'achin
answerto this question.My impres- disturbancesin the balanceof nature, ',bility tc
Beforewecananalyzethem,wemust sion is, however,that as the interac- but it can never lead to changes;on ;,l1da pI
dealwith oneotherconcept,the idea tion of predators and prey, of para- the contrary,it is a deviceto preserve Inatear
of a perfec~balanc~of nature,an idea sitesa?d t~eir victims,the frequency a ste.ady-statecondition. As such it 'I'he;eci
prevalentm the eIghteenthcentury: of extInction, and the struggles of continued even after 1838to be an <Iuality
nothingin natureis toomuch,noth- competingspeciesbecamebetter important componentof Darwin's II >rtanc
ing too little, everythingis designed known,the strugglefor existencewas thinking, particularly in his biogeo. !l;story:
to fit with everything else.Rabbits moreand morerecognizedasa "war" graphic discussions(e.g.determina. mostdr
and hareshavelots of youngbecause or fight, a strugglefor survival, "red tion of speciesborders). ill West
food must be availablefor foxesand in tooth and claw" asTennysonlater '.Illthe
other carnivores.The wholeeconomy expressedit. Bonnet (1781)and de It is only whenoneappliespopulation ::ni ue'
of nature forms a harmoniouswhole Candolle(1820)(13)emphasized that thinking to the strugglefor existence Ilill ~o~
that canin no waybedisturbed.This this war amongspeciesconsistednot that one can make the crucial con.
is why Lamarck,whowasvery much merely of a predator-prey relation- ceptualshift to recognizinga struggle 'I'herei
an adherentof this concept,couldnot ship but of competition for any and for existenceamongindividuals of a II >pula1
conceiveof extinction.Cuvierlikewise all resources(14).However,it wasn<?t single population. This, as Sandra ('I~ormo
had adopted.it, a,ss~own in corre- at all a~preciated h?w fier~e thIS Herbert (17)wasth.efirst to.r~cognize Malthu
spondencewith hISfriend Pfaff. He struggle IS,and Darwm admIts that clearly, was Darwm's decIsIvenew 'llrious'
transferredthe sameconceptto the "even the energeticlanguageof .de insight resulting from his readingof ; hus's~
structure of an organism, which he Candolle does not convey the warrIng Malthus. If most individuals of every I)Opulat
visualized as a "harmonious type" in of the speciesas [convincingly as does species are unsuccessful in every whats04
which ,?ot~ing could be changed. the). inference from Malthus" (15). generation, th.e? there must be a ~o. t!le sub
Everythmg m such a complex system lossal competitIve struggle for eXlS. Malthu
is so perfect that any change would Nevertheless, it is highly probable tence among them. It was this con. ;'xpone
lead to deterioration, that Darwin had been gradually con- clusion whi~h made Darwin think at .'ver a
ditioned by his reading to a far less once of varIOUSother facts that had i,l Cbal
This type of thinking was still domi- benign interpretation of the struggle been slumbering in his subconscious i Iltrodu
nant in Darwin's day.,not only among for existence th.an that held by the but for which, up to that moment, he 1)(lsitei
the natural theologIans of England natural theologIans. The mere fact had had no use. ,'Iaims'
but also on the Continent. Indeed, that Darwin had adopted evolution t !lus st
one can find a number of entries in must have made him aware of the Darwin's reading of Malthus was Illaxim
Darwin's Notebooks which seem to frequency of extinction and of the dramatic and climactic, and it does Illilt yo
reflectthiskindof thinking.But was unbalancesand adaptationallags not matterwhetheroneinterpretsit II il:ety ~
Darwin still a wholeheartedsupporter caused by evolutionary changes. as. a ~omplete reversal of .Darwin's Illuxim
of the concept of a harmonious bal- From Aristotle to the natural theo- thmkmg,or whether one believesthat 'lIme 0'
ance of a benign nature? This is very logians it was considered axiomatic "the e~Idenc~ sugges.ts that the (Iesirab
much a question, and it is a very im- that a belief in a harmonious universe change m choIce of UnIt was a pro. I'reater
portant question because it affects and "perfect adaptation" in nature, tracted proces~,stretched over a year ~(Irts 0
the interpretation of what Darwin or in a creator continuously active in o~re, and linked to other aspects 1';lllnot
understood under the term struggle correcting imperfections and imba- o~ hiSthought" .(18). I myself hold II) the
for existence. lances,was incompatible with a belief wIt~ the latter VIew, be.causethe ca. I rIle t)
in evolution. By necessity, accepting pacIty to be able to Interpret t~e 'I.II1~0
For us moderns the term means a evolutionary thinking undermined a Malthus stateme!1t on exponential tl,'sirab
rather fierce fight with no holds continued adherence to a belief in a gro~th.o~ populatlon~ and to appl~ it ~reater
barred. But for the natural theolo- harmonious universe (16). to. m~Ividuals r~qulres. populatIon 111(lefin
gians the struggle for existence was a thinking, and this Darwm had been
beneficial feedback device, the func- Struggle among species gra~ually acquiring during the pre. \\'here
tion of which was ~o maintain the or individuals? ce?mg year and a hal~. T?at every.' IJllllula:
balanceof nature. It IS,as Herder (11) thing came to a dramatIc climax on 28 ,1"lltlY4
calledit, "the balanceof forceswhich Of far greaterimportanceis a second Septe.mber1838,however,is beyondr IllS au'
brings peaceto the creation." Lin- question: Betw~en whom does the questIon. ~ !,'Ilers,
,nae.usdevoted an entire ess~yto the str':lggle fo.r existence take place} . f ,11:;lin
Policeof Natureand emphasIzed that ThIs q~estlonallows for two dras~I- Umqueness of the 1 l'I"'I>arf

.
"those laws of n~tur~ by which the cally ~If~er~n~
answers.In the entI~e indi vid ual } '111<lyil
number of speCIes m the natural essentlallstlc literature the struggle IS ~ l,rf'pdiJ
kingdoms is preserved undestroyed, cons!dered to take place am°n.g The w~ole. c.oncept of comp~titio " Illsistei
and their relative proportions kept in spe.cles: The ba~ance of nature. ~s amon~ mdlvlduals. wo~l.d be Irrele.!ii 11:lrwiI
proper bounds are objects extremely mamt.-amedby thIS struggle: eve.nIf It vant If. all t~ese l,ndlvI~uals wer t I"l hin,
worthy of our attentive Pursuit and occasIonall
y causesthe extInctIon of typologIcally Identical-if they a 'I I II III t
I
c', 324 American
Scientist,
Volume
65,- _cJ .
 -

tion.of had the same essence. Variability Notebooks until about three months lected-only if at least part of it is
he lIt. does not become meaningful in an after the Malthus reading. For myself, heritable ("Fact 5"). Like the animal
) to .de evolutionary senseuntil a concept has I am rather convinced that Darwin's breeders from whom he got so much
maJor developed that allows for differences own presentation is nevertheless es- of his information, Darwin took this
)ks ~p among the individuals of the same sentially correct. heritability completely for granted.
mal11 population. According to this concept
:orrecl each individual may differ in the If we ask ourselves what Darwin And this assumption can be held
iature. ability to tolerate climate, to find food would be likely to enter in his Note- quite independently of the assump-
~es;011 and a place in which to live, to find a books, we would certainly say new tions concerning the nature of the
'ese~~e mate, and to raise young successfully. fact£ or new ideas. Hence, since it was genetic material and of the origin of
;uc II The recognitionof the role of indivi- not a new subject, animal breeding newgeneticfactors.Darwin'sideason
be.a,11 duality is not only of the utmost im- surely would not qualify. Darwin's these subjects were quite confused,
I~wm S portancefor an understandingof the bestfriendsat CambridgeUniversity but fortunately Darwin tr-eatedge-
)Io~eo. history of biology but it is one of the were the sons of country squires and netic variation as a "black box." As a
rmma- most drastic conceptual revolutions of owners of estates. As Himmelfarb naturalist and reader of the animal
in Western thought ("Fact 4"). We called them, they were the "horsy set" breeding literature, he knew that
I . call the concept which emphasizesthe of Cambridge University, riding or variation was always present, and this
~ atlol1 unique distinctness of every individ- hunting with dogs on every occasion. is all he had to know. He was also
Isten('(' ual population thinking. One can hardly doubt that all of them convinced that the supply of variation
al COI1 to a greater or lesser degree were in- was renewed in every generation and
t~ug1}e There is little doubt that Darwin's tensely interested in animal breeding. thus always abundantly available as
~s ~ a population thinking received an They must have argued a great deal material for natural selection. In
::)an.ra enormous boost through reading among themselves about Bakewell other words, a correct theory of ge-
COgnIZ(' Malthus at that right moment. Yet, and Sebright and the best methods of netics was not a prerequisite for the
d~ new curiously, when we go ~hrough Mal- breeding and improving dogs, horses, establishment of the theory of natural
mg ()I thus's writings we find no trace of and livestock. selection.
)f ever:. population thinking. There is nothing
l ever.v whatso~ver
.even faintly relating to How ~lse-other th.anthat it had a Natural selection
Je a c!) the subJectm those early chapters of great mterest for him-can one ex-
J! exis Malthus that gaveDarwin the idea of plain that Darwin, in the excessively
The next question we have to answer
ll~ COI1- exponential growth. There is, how- busy period after the return of theis how Darwin arrived at the actual
,hmk ,II ever, a reference to animal breeding Beagle, devoted so much of his timeconcept of natural selection on the
1at ?a<l in Chapter 9, but here the subject is to studying the literature of the ani-
basis of the stated five facts and his
nscio~~ intr.oduce~ to prove exac~ly the op- m~l bree~ers? To be .sure, Da!v:in's
firs~ inference (Figure 1). In his a~-
lent, ( poslte pomt. After referrIng to the prImary mterest wasm the ongm of . toblography (pp. 118-20) Darwm
claims of the animal breeders, Mal- variation, but in the course of his stressesthat he "collected facts on a
thus states, "I am told that it is a reading Darwin could not help ab- wholesale scale,more especially with
l~S;as maxim among the improvers of cattle sorbing the important lessonfrom the respect to domesticated productions,
It 0(:, that you may breed to any degree of breeders-that every individual in by printed enquiries, by conversation
:pre~s ~1 nicetyyou please,andthey found'.~is the herd was different from every with skillful breedersandgardeners,
)arwm , maxim upon another, which is, tl t other one, and what extreme care had and by reading. . . . I soo~ perceived
ves t~al someof the offspring will possessth~ to be used in selecting the sires and that selection was the key-stone of
iat t It' desirablequalitiesof the parent£in a dams from which to breed the next man'ssuccessin makingusefulraces
s a prt) greaterdegree." He then produces all generation. I am quite convinced that of animals and plants. But how se-
~ra year sort£ of facts and reasons why this it is no coincidence that Darwin lection could be applied to organisms
aspe('I, cannot possibly be true, leading him studied the literature of the animal living in a state of nature remained
el~ holtl to the conclusion that "it cannot be breeders so assiduously exactly dur- for some time a mystery to me" (21).
~t e (',I true, therefore, that among animals, ing the six months before reading In 1859 he wrote to Wallace (22), "I
)ret t~~t; so~e of the o~f~pringwill possess.the M.althus. In this I agreeentirely with came to th~ c?nclusion that selection
onenltl'l desirablequalItIes of the parent£ m a Michael Ruse (20). was the prInciple of change from the
app y 11 " greaterdegree;or that animals are study of domesticproductions;and
.o~~lt)11 indefinitely affectable" (19). It was not the processof selection but then, reading Malthus, I saw at once
a eell the fact of the unique differences how to apply this principle." To Lyell
the prl' Where, then, did Darwin get his among individuals which Darwin re- he wrote, with reference to Wallace's
It ever:, population thinking, since he evi- membered when suddenly becoming theory, "We differ only {in] that I was
lb on !.,~ dently did not get it from Malthus? In aware of the competition among in- led to my views from what artificial
; eYOl1t his autobiography and in various dividuals, of the struggle for existence selection had done for domestic ani-
letters,Darwin emphasizedagain and among individuals. Here we have the mals." Traditionally, these state-
IIgain that he had been mentally fortuitous coming together of two ments were accepted by the Darwin
preparedfor the Malthus principle by important concepts-excessive fer- students as a correct representation
.tudying the literature of animal tility and individuality-which of the facts.
.. breeding.Recent commentators have jointly provide the basis for an en-
lP~tltl;)~) insis~d that this must be a lapse of tirely new conceptualization. This interpreta.tion, however: has
Ie Irre t l)arwm's memory becausethere is beenchallengedm recentyearsm the
al~ w~r;; f)()thingabout selection and very little Variation can b~ of evolutionary sig- wake of the discovery of Darwin's
t ey J "oout animal breeding in Darwin's nificance-that is, it can be se- Notebooks. Limoges and Herbert

1977 May-June 325

- - - -

point out that in the first three animal breeding had prepared his is happening is for the common good slill to
Notebooks Darwin nowhere refers to mind to appreciate the role of the in- and helps to maintain the perfect (lark as
selection or to the selective activities dividual and its heritable qualities. harmony of the world. The world of mind ir
of animal breeders,particularly in the Indeed I am convinced, and in this I Malthus was a pessimistic world: ois bec(
production of new domestic races. agree with Ruse (20), that the many there are ever-repeated catastrophes, 1837) ~
They claim that Darwin was inter- years during which Darwin had been an unending, fierce struggle for exis- reading
ested in domestic animals only be- exposed to the ideas of the animal tence, yet the world essentially re- lIasbec
causehe hoped to find evidence con- breedershad preconditioned his mind mains the same. However much shift in
cerning the occurrence of variations to appreciate the importance of the Darwin might have begun to question Ilermit1
and the mechanisms of their pro- Malthus principle. This dormant the benign nature of the struggle for kll0wn
duction, matters that are difficult to knowledge was actualized under the existence, he clearly did not appreci-
study in wild populations. impact of reading Malthus (2.5). ate the fiercenessof this struggle until I )ne of
reading Malthus. And it permitted ('(illsistl
It is true that the term selection does The natural selection of individuals him to combine the best elements of who ha
not occur in Darwin's Notebooks; it is with particular heritable qualities, Malthus a~d of natural.theology: it loe his1
first found in his 1842 sketch in the continued over many generations, brought him t? the b~lIef that the selectio
words "natural means of selection" automatically leads to evolution, as in struggle for ex~stence~snot a hope- tent tho
(23). Darwin here refers to artificial, Inference 3 (Figure 1). In fact this less steady-state condition a la I;lith af
selection by-the term "human selec- process is sometimes used as the Ma!thus but the very means b'y work 01
tion." Actually, in the Notebooks definition of evolution. In this con- wh~ch the harmony of the world ~s selectio
Darwin not infrequently refers to the nection it must be emphasized once ac~iev~d and maintained. Adap- t oat th
process of selecting, but he uses a more that Darwin's inference is ex- tat~on ~sthe result of the struggle for I)t~liefs
different term-picking. actly the opposite of that of Malthus, existence. Ii; I )arwin
who had denied that "some of the ~::{ tois acc
I am willing to grant to the recent offspring will possess the desirable The eve~ts on 28 September 1838ar .Aft toe yea
critics that thereis no evidencein the qualities of th~ parents in a greater of grea~mteres~to studentsof theory - ,I;muar
Notebooks of a simple application to degree." Indeed, Malthus used his form~tlon. S~emg to what ~n extent I )arwin
the evolutionary processof the anal- entire argument as a refutation of the Darwm wa~ m the ~ossesslon ?f al! i\doptiJ
ogy between selection by man and thesis of Condorcet and Godwin of th~ other ~lecesof his theory p~lor to loan tb
selection by nature. This is quite ev- human perfectability. The Malthus- this date, It becomesclear that m the 1;lctor I
ident when one readsthe crucial entry ian principle, dealing with popula- ~a~ of a coIl?p'lexlystrucwred theory, I()rmerl
in the Notebooks on 28 September tions of essentialistically identical It.ISnot suffIcIent to have most of the ,ign W
1838(here reproduced in the original individuals, causesonly quantitative, pieces; no, one ~~st ha,,:ethe~ ~ll, Ilowevl
telegraph style): not qualitative, changes in popula- Even a sm,~" d~fIc~~ncy,lIke.defm~ng "Iution
tions (26), the word varIety typologically m- 111inin
,
Take Europe on an averageevery speCIes stead . . of populationally ' might be .
I ) arwm
must havesamenumberkilled yearwith How great was Darwin's sl!ffI,cIent to prevent the correct (-rucial
year by hawks, by cold etc.-even one d b M 1h ? piecIng t~gether of t~e components, ;1(lopt
speciesof hawk decreasingin number e t to a t us. Equally Important IS the general "oard"
must~ffectinstantaneo~slyallt?erest,- That it was the Malthus reading ide,ological attitude. of the the?ry- (I)nstel
The fInal causeof all this wedgIng, must which acted as a catalyst in Darwin's builder. A, pe~son like ~lyth might which!
be
. to sort out proper structure, and adapt.
. mmd m . prod ucmg
. th e th eory0f nat - have had m hIS Possessionthe very {'( I ,
It to changes-to do that for form, which . b d. d d samecomponentsof the theory as
Malthusshowsis the final effect (by ural selection
h . cannot
d b De Ispute
. h. anIf Darwm
. but would havebeenunable
meanshoweverof volition) of this popu- was emp aSlze y arwm Imse . '1' 1le t ra
. d ' H h to pIece t h em toget h er correct I y
lousnesson the energyof man.Onemay agaIn an agaIn. owever, w en we. , ... 1111I Igat E
saythere is a forcelike a hundredthou- analyze the componentsof the theory, owIng ~oan mcompa~Ible ~deoiogical toeory'
sandwedgestrying [to} forceeverykind as we havejust done, we find that it is commItme~t. Nothmg Illustrates (('ractio
of adaptedstructureinto the gapsin the primarily the insight that competition b,etter how Important the general at- ilnporu
economy?fnature,or ratherforminggaps is among individuals rather than trtude and conceptual !ramework ~f toe rea
by thrustIngout the weakerones(24), among species that is clearly a Mal- the maker o~ a theory IS than the SI- ~rowth
thusian contribution. To be sure this multaneous, Independent pr.oposalof (Illite m
The simile here is "wedging" not in turn led Darwin to a reevalu~tion the theory of natural selection by A, (I.mple
"selecting." Thus it.a.ppearsthat ~he of other phenomena, such as the na- R.. Wallace. He was one of the few
argumen~ ,?fthe CritICShave consld - ture of the struggle for existence, but people,.pe!hapsthe only one,w~o had ;\ 11dit i
erable valldl~~. !l°wever~ the analogy only as second-order consequences.I ha~ a slml~ar set of past experl.ences: 'I)Onsib
between ar~lflc~al selection and nat- agree with those who think that the a life dedlcat~d to natu~al ~Istory, toe the<
ural selectIon IS not necessary for Malthusian thesis of exponential years of c?"ectmg on tropical Islands, ~enerat
Darwin's conclusions.Inference 1 and growth was the capstone of Darwin's and readIng Malthus. lifetimE
Fact 4 automatically ~esult i~ Inf~r- theory. "The one sentence of Mal- . . , (I.mpe
ence2 (natural selection).It ISquite thus" acted like a crystal dropped Preparation of Darwm s ;Imong
likely that Darwin did not see the into a supercooledfluid. mind ::enesi
obviousanalogybetweenartificial , . IJl~stkn,
andnaturalselectionuntil sometime There is, however,alsoa secondand It ISobvIousthat we havelearneda IJl'twee
after the Malthusreading.Yet, I have moresubtle MalthusianimP.act.The ~reat ~eal in recent ye.arsabout ~h,e ('xplan
little doubt that the copiousreading world of the natural theologianswas Ingredientsthat went Into Darwms ;1rate S
which Darwin had done in the field of an optimistic world: everything that theory of natural selection,but we are Ii nally

326 American Scientist, Volume 65

 ---

;ood till to a considerableextent in the in the 19308and 1940s,during the vol. 2, nos.2--6(abbrev.NBT); G. de Beer,
fect.. 'd ar k as t 0 what went on m
. Darwin's so-called "evolutionary synthesis."
...
M. J. Rowl~nds,and B. M. Skramovsky.
1967.Darwm'sNotebooks ontransm\lta-
d of .~mind in the year and a half between Yet, the res~stan.ceto selectiomsm tionof species,
pt. 6.Op.cit.,vol.3,no5.
)rld: ';fhis becoming an evolutionist (March amon~ non biOlogIStsand lay pe<}ple 6. Forindications
ofashiftin Darwint.<-ward
.hes, J:1837)and the day of the Malthus has still not been overcome entirely populationthinking,seealso Ghiselin
~xis- 'reading on 28 September 1838.What even in our day. (note4),pp.57-59.
, re- has becomeclear is that it required a 7. SeealsoGruber(note1),pp.208-14.
I~ch shift in at least four major c.oncepts, References 8. NBT,5 (Note~k~, p. 135). ..
Ibon permitting Darwin to see five well- 9. Found~y Darwmm the .6t~editionof
e for known facts in a new light. 1. Darwin'searliertheories
ofevolution,
prior Mal~hus s EssayonthePrIncIpleof Pop-
reci- to the Malthusepisode,havebeenper- ulallon(1836),
vol.1,p.6.
~ntil . ceptivelyanalyzed byH. E.Gruber.1974. 10.Gruber(note1),pp.161--63.
One .of these .shlfts has been rather DarwinonMan.Dutton;C.Limoges. 1970. 11.J. G.Herder.1784.ldeenzurPhilosophie
tted, consistently sidestepped by all those La selectionnaturelle. Paris: Presses der Geschichte
derMen.~chheit, II, 3, p.
ts of who have occupied themselves with Universitaires
deFrance(Englishedition' 89.
y: it. the history of the theory of natural to appearin 1~7?);andE. D. Kohn.1975. 12.C. Linnaeus.[1781).Policeof Nature
the selection.It is the questionof the ex- C:harles.DarwIns Path to Nat~ral Selec- [Amoen.Academicae],trans.F. J. Brand,
ope- tent t hat Darwm . t Ian
., s 1ass 0f ChrlS . lion. Urnv.of Massachusetts
dlss.
.' pp 131-32. London .
j La e ' th f ~ ted
th t 1f 2. Nora Barlow, ed. 1958.The AutobIography 13, A. S. de Candolle. 1820.Essai Elementaire
.
'.
by lal
work a lec
h. h the concep ua rame-
th f t ie of Charles Darulin, 1809-1882. London: d GeographIe . B0t amque.
.
Ld is
. 1 on. w IC e eory 0 na ura
seectlonr~ts. Th. ey have.over1~ ked
Collins.
3. As for Instance
. J. Monod(1971)m
.' the l~st
14 E
lengthy
L II
ven ye list ofemp
h . d th O
the causes
d h.
aslze of extinction
IS, an is
IS
iap- that the discussion of his religious two chapters of Chanceand NecessIty strikingly in conflict with the conceptof a
e for beliefs is an important chapter in (Knop.f): . perfectlyadapted,well-balancedworld.
Darwin's autobiography (27). From 4. In addition to Kohn,Gruber,.andLlmog~s 15. NBT, 5 (NotebookD, pp. 134-35).
, . . (note 1), those who have discussedthis .
this account It becomes evident that bl ' t S S .th 1960 16. See also Limoges (note 1), pp. 40, 44-45, 70,
3 are pro em In recen years are , mi.. d 79 80
the years between October 1836 and Theoriginof "TheOrigin."Advancement an -.
eory January 1839were the years in which of Science64:391-401;
G. deBeer.1965. 17.S'l:Ier.bert(note4), pp.209-17.Seealso
'tent. . fal..th (28)
Darwm. 1ost muc h 0f hIS CharlesDarwin'. A Scienti'ic
".. Biography . Ghlselln(note4), p. 59.
If all. Ad ' 1 1 . h DoubledayAnchor;M. T. Ghlselm.1969. 18. Gruber(note 1),p. 165.
optIng natura se ectlon rat er TheTrium
or to th h h d f G d h t . p h Of the Darwinian Method. 19. Malth us. 1798[seenote 9]. Chap.,9 p.
h an t e an 0 0 as t e ac Ive Univ.of CaliforniaPress;R. M. Young. 163.
1t e factor respon~ible for. all that was 1969.Malthusandtheevolutionists.
~a~t 20. MichaelRuse.1975.CharlesDarwinand
eory, formerly considered evidence for de- and Present43:109-41; 1971.?Darwl~s artificial selection.J. Hist. Ideas 36:
fthe sign was of course the last step. metaphor:Doesn~tureselect. Mon~st 339-50.
1 all. " 55:443-503;P. Vorzlmmer.1969.Darwm, ,
, . ,However, even the acceptance of ev- Malthus and the theory of natural selec- 21. AutobIography(seenote2), pp. 118-20,
m~ng olution was already a fatal under- tion. J. Hist.ldeas 30:527-42;S. Herbert. 22. Darwin to Wallace,6 April 1859.A. R.
Y m- mining of natural theology. That 1971.Darwin, Malthus, and selection.J. Wallace.Letters and Reminiscences, vol.
t be Darwin lost his faith in the same Hist. Bioi. 4:209-17; P. Bowler. 1974. 1,p. 136.
rrect
ents. CU'
r cia 1 y ears in .w hI.ch . be . began to Darwin's
Med. andconcepts of variation.
Allied Sci. J. 1976.
29:196-212; Hlst. 23. Darwin. 1842.Sketch.
TheFoundations InOrigin
of the F. Darwin.1909.
of Species,
adopt population thInkIng (and Malthus, Darwin, and the concept of p. 46.CambridgeUniv. Press.
leral "hard" inheritance) is a synergistic struggle.J. Hist.ldeas 37:631-50. Seealso 24. NBT,5 (NotebookD, pp. 134-35).
. ht
!ory- constellation ' the im p ortance of J. C. Greene.
fD 1975.
. Reflections
d . J H . on
t B'the
I 25. Gruber (noe
t 1),po118.
11g which should not be underestimat- progress0 arwmstu les. . IS. 10. .
8:243-73. - 26. Limoges (note1),p. 80;alsodeBeerand
very
'y as ed. 5. G. de Beer, ed. 1960--61.Darwin's Note- :
others
' - bookson transmutation of species,pts. 1-5. 27. AutobIography (see note 2), pp. 85-96.
,ab:e The traditional Interpretation, pro- Bull.Brit. Mus.(Nat.Hist.),Hist.Series, 28. Gruber(note1),pp.208-14.
e~t y mulgated by Darwin himself, that his
gical theory was the result of a simple in-
~ates leraction between a knowledge of the
11at-, importance of artificial selection and
rk 01 the realization of the exponential
Ie si
oal01
-, growth of populations is, however,
quite misleading. The theory is far too
-
>y A. complex for such a simple origin.

~
!few
>had And it is this complexitywhich is re-
nces: IIponsiblefor the enormousresistance
tory, thetheoryencounteredin subsequent '\ .~t
f~ ~~
mds, ~enerations.It led, still in Darwin's ~ ~~
lifetime, to the proposal of many
, competing explanations of evolution,
S Amongwhich saltationism, ortho-
Kenesis,and Neo-Lamarckism are
ledh
a I>eHt
known. To report on the struggle
between
Darwinismandtheseother ~ / !
. r-.zt
t ~ ,e t';xplanatorytheories would be a sep- 'i
wm H urate story. Evolutionary biologists "One reason this condition is so hard,to cure is that it's
'e are , 'ffinally cameto completeagreement 111111111. caused
byoneof the teeniestviruseS"'.n , ..n..