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Choosing Selection: The Revival of Natural Selection in Anglo-American Evolutionary Biology, 1930-1970 Author(s): Stephen G.
Brush Source: Transactions of the American Philosophical Society, New Series, Vol. 99, No. 3 (2009), pp. 1-127, 129, 131-135, 137-183 Published by: American Philosophical Society Stable URL: http://www.jstor.org/stable/27757443 . Accessed: 29/03/2013 18:18
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Choosing
selection
The Revival ofNatural Selection in Biology, 1930-1970 Anglo-American Evolutionary
.-to'
by Stephen
Brush
AMERICANPHILOSOPHICALSOCIETY | PHILADELPHIA | 2009
CHOOSING
SELECTION
CHOOSING SELECTION
The <rR?vival Dâtural Selectionin of

zAnglo-American Evolutionary Biology,
1930-1970
STEPHEN G. BRUSH
AMERICAN
PHILOSOPHICAL "Philadelphia 2009
SOCIETY
TRANSACTIONS
of the AMERICAN
PHILOSOPHICAL
SOCIETY
Useful Knowledge Held at Philadelphia forPromoting Volume 99, Part 3
American Philosophical Society for its Copyright? 2009 by the

Transactions series, Volume 99. All rights reserved.
Text
Waters Titling display Set in Adobe Caslon with United States ofAmerica. Printed and bound in the
by Graphic Composition, Inc., Bogart, Georgia.
and cover design
by E. H. Graben.
isbn-13: us
978-1-60618-993-1 issn: 0065-9746
library
of
congress
cataloging-in-publication
data
Choosing cm.?
selection
Brush, Stephen G. : the revival of natural selection of the American
evolutionary biology,1930-1970 / StephenG. Brush,

Philosophical Society held
in Anglo-American at Philadelphia
forpromotingusefulknowledge, issn 0065-9746 ;v. 99, pt. 3)

Includes 1. Natural 2. Natural references bibliographical isbn 978-1-60618-993-1 and index.
(Transactions
selection?Research?Great selection?Research?United Britain.
Britain?History?20th States?History?20th States. 4. Biologists?United
century. century. I. Title.
3. Biologists?Great
qh375.b78 576.8'2?dc22
2009 2009020654
front
cover
illustration:
of Adelaide, ofR.A. Fisher, edited byj. H. Bennett (University 1971).Used by permissionof the UniversityofAdelaide.
R. A. Fisher,
from Collected Papers
CONTENTS
1. Introduction 1 2. Fisher: A New Language forEvolutionary Research 9 3. Wright: Random Genetic Drift, a Concept Out of Control 19 4. Haldane: A Mathematical-Philosophical BiologistWeighs in 5. Early Reception of the Fisher-Haldane-Wright Theory 30 6. Dobzhansky: The Faraday of Biology? 35 7. Evidence for Natural Selection before 1941 39 8. Huxley: A New Synthesis Is Proclaimed 46 9. Mayr: Systematics and the Founder Principle 51 10. Simpson: No Straight and Narrow Path forPaleontology 11. Stebbins: Plants Are Also Selected 60 12. Chromosome Inversions inDrosophila 68 13. Ford: Unlucky Blood Groups 14. Resistance toAntibiotics 74 15. Two Great Debates: 16. Dunkers and Drift 62 56
27
Snails and Tiger Moths 79
76
17. Gould: Why Did the Synthesis Harden? The Changing Views 81 ofDobzhansky and Wright 18. Why Did the Synthesis Harden? The Views ofOther 87 94 103 Founders and Leaders
19. The Peppered Moth 91 20. The Triumph ofNatural Selection? 21. Is Evolutionary Theory Scientific? 114 22. Context and Conclusions Acknowledgments Tables 131 Bibliography Index 171 137 129
NO There
SCIENTIFIC
THEORY
ISWORTH
ANYTHING
unless
it .. .
enables us to predict something which is actually going on. are still a number of not believe in the people who do not because it is an theory of evolution. Scientists believe in it, attractive theory, but because it enables them to make predic tions which come true. âldane 1940, 7-8
THE
FACT THAT NATURAE
planation of adaptation acceptance of Darwinian tion is very widespread, abundant
ralists have come to feel that adaptation ture of life requiring an explanation.
and some of it is very remarkable. So is it, and so marvelous are parts of it, thatmany natu is the outstanding fea
SELECTION offers a general ex is one of the chief reasons for the rapid theory among biologists. For adapta
?hulletal
1941,359
Introduction
it acts, selection, of the variations on which was those variations. Nevertheless, by 1970, Darwin a was correct when the that posing theory essentially were made. In came to believe particular, biologists natural
When did biologists accept Charles Darwins theory of evolution? One might argue that his theory, in its original form, was never was ac more than a handful of theory that biologists. The accepted by was based on a different of understanding cepted in the twentieth century and on the source of given credit for pro necessary corrections that natural selection
on small mutations, combined with Mendelian acting principles of hered cause of factors, such ity, is the major evolutionary adaptation. Additional as so and forth,may be in isolation, polyploidy, genetic drift, geographical volved in some cases (especially in speciation) and at some levels, but they
would
generallybelieved by historiansof biology to have been established in the

to the core of an "evolutionary synthesis," which brought as and such genetics, zoology, botany, gether previously separate disciplines a unified science of same time, it could into biology. At the paleontology 1940s. It was be considered many possible on the scholarly literature history of this synthesis.2 it excluded rather than a synthesis, because causes of evolution (Provine 1988). There is now a growing a constriction
in the twentieth cen little by themselves. As modified accomplish is called neo-Darwinism,1 Darwins tury, theory, sometimes (misleadingly)
Why did biologists accept themodern version ofDarwins theory?I

1. As
was the name used noted by Dobzhansky (1942) and Bowler (2003), "neo-Darwinism" to the to is the of which Weismann, only superficially similar designate theory of "Darwinism" natural selection and 1993). Fisher, Haldane, theory Wright (Gayon developed by itself is often misleadingly used in popular writings and in reception studies by historians to mean not Darwin's 1974). I will therefore theory of natural selection but simply "evolution" (e.g., Glick around 1900 avoid both of these terms except
2.Mayr and Provine (1980,1998); Delsol (1991); Smocovitis (1996);Gayon (1998);Gerson works citednext. The phrase (1998); Beurton (2002); Bowler (2003); Shanahan (2004); and other
synthesis" comes from the title of Julian Huxley's book (1942); see comments by
in quotations.
Junker(2002) on the terminology.
"evolutionary
CHOOSING
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discuss natural
in this book
what I will call the natural selection that (NSH): thehypothesis hypothesis
selection, with an ample supply of variation in heritable characters, is not only the major process involved in evolution (with the help ofgeographical in some cases), but also that Lamarckian isolation or random polyploidy effects, no genetic drift, and macromutations have essentially evolutionary significance. was a bare The NSH of majority accepted by evolutionary biologists for a short time (in the 1950s and 1960s). In were only particular I ask, what the reasons they gave for adopting that and for rejecting the hypothesis argument that drift plays a significant role in evolution? Although
the reception by evolutionists
and other biologists
of
selection should be considered a process rather than a cause. It does not "act"?rather, it is the result of heritable differences and occurs when variation among individuals arises due to mutation or other causes.3 natural This
I have generally used the concepts and terminology of evolu tionary biologists in the time period under discussion, these concepts and terms may be to a modern reader. Thus, John Endler argues that confusing
a to compare the reasons study is part of program why scientists new theories. in different fields accept (or in some cases reject) proposed was colored Is it true that "the reception of neo-Darwinism by prevail what specific empirical evidence scientists give to the confirmation was most persuasive? What
of thought"(Depew and Weber 1985b,240)? If so, ing empiricistcurrents

weight did the essential and accord science from
desideratum
of predictions, supposedly of the hypothetico-deductive scientific method, ing to Karl Popper, the basis of the criterion for demarcating
pseudoscience?4 That question
is especially appropriate in the case of evolutionary biol s own a since ogy, critique of Darwinism Popper provoked long-running debate among philosophers and biologists about whether biological theories
selection is afact, not a of anything (Wells, Hux set This be the view of 1931,1:604). may against philosopher Elliot Sober (1984a, Reisman and Forber that selection is a "force" (or "cause") natural 27,141), (2005), supported by that acts together with mutation, drift, and other forces. Dunn is one of the few biolo (1967,49) 3. Endler (1986,4,29; 1992). Natural ley, andWells
Matthen andAriew (2002);Nanay (2005); Skipperand Millstein (2005); (1989); Skipper (1999); Glymour (2006); Pigliucci andKaplan (2006); and Plutynski(2007a).
I am grateful to an anonymous Selection in theWild book, Natural time period covered by this book. referee who (1986), even excellent suggested that I consult John Endler's as a it is well outside the though primary source,
use that see also the 1944 comments in J. Cain (2004, 68). For gists who language; by Stebbins further discussion of selection-type theories in biology and other sciences, see Darden and Cain
I use theword "prediction" in its nontechnical towhat sense, corresponding 2002). Here physicists and some other scientists call "prediction in advance" and to what some call "novel philosophers next note for further discussion. prediction." See
4. Popper (1934,1959,1962) and Brush (1995 [and references cited therein], 1996, 1999a,
Introduction
"philosophy of science," allegedly biased toward in part from this debate. physics?resulted in biology is that A remarkable aspect of the debate about predictiveness its participants rarely mentioned the predictions, some of them confirmed, distinct from traditional actually made by evolutionary theorists. Perhaps they assumed that if bio to make testable it doesn't matter logical theories don't need predictions, if they do so. Or one
should be expected to make testable predictions, and whether more gen standards for biology should be different from erally, the methodological those of the physical sciences. The creation of "philosophy of biology"?as
might argue that the basic Popperian premise?that real scientific theories are always judged by the results of their predictions, rather than, for example, by their ability to provide plausible and consistent wrong. explanations?is Another problem with who proposes someone else would the emphasis on predictiveness is that the person a not agree that a theory may particular observation made by
Kuhn (1962) be a fair test of that theory.As Thomas and others have pointed out, it is rare to find genuine "crucial experiments" in science, the kind of testwhose outcome would force one side or the other to abandon its theory. In the controversy between Fisher andWright about often rejected the rel the role of random genetic drift in evolution, Wright even when evance of tests of his theory (which was itself alleged evolving), the results seemed will abandon
chromo Hunt Morgans that the most convincing evidence was Calvin Bridges' discovery of nondisjunction, yet they never mentioned that thiswas in fact a confirmed prediction (Brush 2002). Even Popper evo Darwinian eventually retracted his statement?that not scientific because it does not make testable predic is lutionary theory that such predictions had been made tions?but still did not acknowledge (1978) Popper's paradigmatic example of prediction-testing
her own experiment to confirm it. In the other biological case I studied, Thomas some theory of heredity, geneticists considered
a scientist to be favorable. It does happen, however, that or a favorite theory because of the unexpected failure of his
and tested. Conversely, Einstein's
contradicts his thesis, because general theory of relativity?also astronomers gave at least to the successful cal equal weight physicists and culation of another effect, the advance of the perihelion ofMercury, which had been observed novel prediction.5
inphysics?the discovery Albert ofgravitationallight bending, by predicted
for more than half a century and thus was clearly not a In fact, I have not yet found any examples in twentieth
use
5. Popper (1962, 117, 339-40; 1974, 36-37) and Brush (1989, 1999b). Physicistsgenerally
the term "prediction" to include the deduction of facts already known and use "prediction
CHOOSING
SELECTION
value as a secondary criterion in chemistry (Brush 1996,1999a). of science may wonder why I have focused on prediction Philosophers to the exclusion of other issues relevant to natural selection.6 philosophical I am not concerned with philosophical that scientists themselves worry about
the century physics in which novelty was generally thought to enhance evidential value of a confirmed prediction, although it seems to have some
in evaluating new ideas. This tiveness, which is a major part of the so-called scientific method as defined or not so by many scientists (whether they follow it themselves). It is widely an a believed that falsifiability is essential property of scientific theory that even the U. S. a landmark decision it in Court has cited Supreme justifying account studies of surprising thing about historical and philosophical on its reception in the Darwins theory is the huge quantity of literature itwas not nineteenth century, when itself (although evolution accepted was widely accepted), compared with the much smaller number of stud ies dealing with it in the twentieth century, when itwas accepted. Among is one of the "big reception studies by historians of science, Darwinism (Blackmun Another et al. 1993).
issues per se, but only to the extent them and perhaps take them into was case with certainly the predic
made;
were not known when want to the prediction was they specify that the facts the latter is what philosophers of science call "novel prediction." Elie Zahar redefined a fact that "did not to the "novel fact" to mean the which belong problem-situation governed construction of the hypothesis" that predicted it (Zahar 1973,103), but Iwill not need to use this in advance" when Some recent publications thesis and have the credibility of science have undermined by philosophers shown that explanation of a known fact (misleadingly called and as, or even superior to, novel prediction. See Hitchcock
definition.
of the "predictivism" "accommodation") Sober (2004)
that (2001). Peter Lipton, a leading predictivist, conceded not claim is normative rather than the superiority of novel predictions descriptive; that is, he did that scientists themselves give extra credit for novelty (Lipton 2005). See the book by Losee
may be as good and Scerri andWorrall
nor can we an mechanics, (Villee 1954, 4; 1957, 5). A physicist would predict point earthquake" out that quantum mechanics does make very accurate predictions of atomic properties that can was it predicted facts that were already actually be observed, but it originally accepted because known but not explained; its "predictions in advance" were confirmed was only after the theory accepted.
are tested in terms of their effectiveness in explain it A. and Claude Stone 2; added). 1952, emphasis happen it is not com Villee, noting that "there are some who claim that biology is not a science because even as the most 'scientific'of the sci pletely predictable," replies that physics, "generally regarded cannot make in the field of quantum We ences, is far from completely predictable.... predictions theory and the mechanisms or will predicting what has postulated (Patterson
of the relevance of "falsification" (2005) and its review by Allchin (2006) for further discussion and "predictiveness" to the behavior of scientists. as will be seen below, do not have a on Biologists, single clear-cut position prediction. John Patterson andW. S. Stone wrote: "As with all scientific theories, the validity of an evolutionary to
6.Hull (1974); Sober (1984a, 1984b,2006); and recentissuesofStudiesin HistoryandPhilos

and Biomedical Sciences and of Biology and Philosophy.
ophy ofBiological
Introduction
selection in particular. By contrast, I am not the acceptance of the of evolution by biologists, fact which was almost universal in the twentieth century (see Numbers 2004), but with their debates about how evolution works. with little attention to natural concerned here with
others being Einstein's relativity and Freud s psycho at or before of the Darwinian Almost all analysis.7 reception studies stop 1900 (unless they cover the Scopes Trial and other aspects of the creation versus evolution debate), and in general, they usually discuss evolution three" theories?the
Natural
three canonical works
Selection in the 1930s
I startthe storyat thebeginning of the 1930s, with thepublication of the

J. B. S. Haldane, of modern evolutionary theory by Ronald A. Fisher, and Sewall Wright.8 At that time, natural selection was as a factor in evolution but was not considered suf recognized
generally ficient by itself to account for the observed facts. It was sometimes argued that natural selection is only a negative factor: it can eliminate bad genes but good genes.9 atHarvard Uni Director of the Zoological Parker, George Laboratory ... have one one in asserted that "most modern evolutionists 1925 versity, by come to the conclusion in time that natural selection, which Weismanns was in evolution, may after all be of little real on which significance." Parker favored de Vries's theory of large mutations, natural selection could act, possibly with the help of a little Lamarckism.10 declared to be all-sufficient does nothing to create
was
But Lamarckism, though supportedinEngland byE.W. MacBride

losing credibility. A weak version was periodically
(1929),
revived, then and
Libraries
(through the Research Bibliography using the words "reception," "response," and "reaction" suggests that about 20 percent of reception studies are devoted to Darwinism, 18 per a more cent to to of and 15 Freudian search percent relativity, psychoanalysis; comprehensive the humanities and social science literature would increase the proportion of Freud probably Information Network electronic database) studies studies. In any case, these three have now become in the history of science (Glick and Henderson data see table 1 (p. 131). For established 2001). history emphasizing the earlier as canonical
7. A
search of the online
version
of the Isis Cumulative
and Darwin reception 8. For
cases for
works ofFisher (1918) andHaldane (1924), see Sarkar (2004). 9. work dis Morgan (1932,130-1); G. E. Allen (1968,1980); Punnett (1938,221 [andother cussedbyKimler 1983b,301-2]); Dalbiez (1927,189); Lindsey (1931,70,113); Newman (1932, McKerrow (1937). 395-6); Osborn (1932, 60); and
"An analysis of textbooks and of the evolutionary literature shows that as late as 1930 se in both Britain and the United lectionism was still a minority viewpoint States" (Mayr 1998, cites a number of (1957) gave no details of this analysis. Lack preface). Unfortunately, Mayr to natural selection. objections 10. Parker (1925,113-44; see especially 119).
personal
a revisionist
CHOOSING
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on Lecture, delivered to the Linnaean Society of London March Bio 14,1929, by E. J.Allen, F.R.S., F.L.S., Director of theMarine was of Association the United Allen there stated that logical Kingdom. influence on the heredity of organ plenty of evidence for environmental in the Hooker
later,under names like the "Baldwin effect" or "canalization."11 Saltationism and several versions of orthogenesis were still (de Vries s large mutations) advocated by some biologists.12 A good authoritative summary of the uncertain situation can be found
isms, including Heslop Harrisons work on the production ofmelanic forms inmoths and H. J. Mullers Lamarck x-ray experiments on Drosophi/a.Yet ian theories of this influence had been discredited, and orthogenesis failed to to to a provide the flexibility needed adapt changing environment. This seem to have an excellent an ex situation would provided opportunity for so far failed to a planation by natural selection, but that theory had provide account.13 plausible to natural So in the 1930s, there was no widely accepted alternative selection but instead a need for that process to be clarified and elaborated
radiation-induced
swork on in order to serve as the major process in evolution. H. J. Muller s research on natural pop mutations and Dobzhansky
11. On 12. On
hausen (1949); see discussionby Simpson (1953b),Richards (1987), andLustig (2004). for evolution,seeOlby (1981); Gould (2002, 425-50); Bowler (1978; 2003, 268-72, 327-30);
and Provine (1971,13,56-57, 64-69,91-92). According to Ernst Mendelians had (Bateson, DeVries, Johannsen, Goldschmidt, little use for natural selection" (Mayr 1992b, 179). Mayr, and Morgan) in the 1920s were the leading "saltationists who the pre-1930 arguments about whether large or small mutations are more important
canalization,
see Waddington
(1957,
1960);
on
"stabilizing
selection,"
see Schmal
and Kane (1988). the germ-plasm itself be directly or indirectly acted upon by physical or chemical are in the environment in such a way that variations, or, to be more precise, mutations, changes in the which are capable of being transmitted to future genera produced offspring?variations can be tions [?] is now, I think, Well-substantiated evidence that the germ-plasm forthcoming 13. "Can
and Richardson
seeBowler (1979; 1983,144,162,178-81); Holmes (1948a); Jepsen(1949); On orthogenesis,
so acted upon ..." (Allen 1930,133). Allen asserted that Lamarckian theories, despite the sup now seemed in of the Graham Kerr's argument: port ofMacBride, implausible, especially light "Is there any possible that, were it actually the case that impressed escape from the conclusion as to characters are transmitted, we should have before us a mass of evidence so overwhelming leave not the slightest doubt in any one's mind as to the occurrence of such transmission" (Kerr 62 [as "Natural Selection, as conceived by Darwin andWal 1930,133]). quoted by Allen a as lace, still seems to most competent potent factor, but the number who biologists regard it seem to be must even face all-sufficient would that diminishing. We fearlessly the possibility as a main cause of the whole (Allen 1930, conception must be abandoned organic evolution" 1926: 137). Yet Graham Kerr, in the book of it. just cited, made a rather strong case for natural selection and refuted several criticisms
Introduction
ulations
variation
on similar used different mathematical dane, andWright approaches based to arrive at results that were, with some minor assumptions exceptions, in on the rel with each other. Fisher and good agreement Wright disagreed as it genetic drift to the process of evolution actually in nature, and this disagreement was perceived as a controversy had retreated from his by Fisher and others. However, by 1960 Wright earlier pro-drift position and asserted that he had not earlier and did not now to believe in his deny the primacy of natural selection. (He continued of random occurred "shifting balance" theory, involving random drift in subdivided large popu lations, up to his death.) In this book, I focus on the competition between natural selection and in detail evance
to broaden the definition of natural selection to include helped as a basic a component rather separate factor.14 In their development of the natural selection hypothesis, Fisher, Hal
not discuss period; I do of drift after 1965. This
driftas perceivednot only byFisher and Wright but also by others in that
the shifting balance theory or the revival focus may leave the reader with an unfairly nega ofWrights contributions, an impression that should be
in arguments made about natural selection by biologists and philosophers recent decades, because an not seem to important role in the they did play of natural selection itself before 1970. acceptance the other major alternatives to selectionism had been By the mid-1960s so one discredited, may say that the strong selectionist view had become established knowledge among the leaders of evolutionary theory. (This is So we edge can then ask the question: when, and towhat extent, did this knowl of natural (the primacy selection) infiltrate the writing and teaching some of the other theories for at least a couple as my I have studied, which remained or of decades, selectionism was chal
tive impression remedied by looking at the publications of Provine and Skipper. Moreover, the NSH considered here was rather simplistic compared with later dis cussions of "levels of selection"; I avoid many of the more sophisticated
what Stephen Jay Gould
of the synthesis.) [1983] called the "hardening"
of otherbiologists?
Unlike thodox doctrine I have taken 1970
directionsbeginning in the late 1960s (Dietrich 1994). lengedfrom several

impossibly to from endpoint in order keep this monograph not discuss later research on evolution, ex I do long.
becoming
man ous
14. Muller (1927); Pattersonand Muller (1930); andDobzhansky (1981).According toRo

(1988), discovery. one should also give some credit to Lewis J. Stadler (1928) for his almost simultane
CHOOSING
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cept for historical 1970. This American
and philosophical
writings
relevant to the period before
as Theodosius and Ernst Mayr, prominent evolutionists such Dobzhansky who emigrated from other countries in the 1930s. But it does not include of Russia, Germany, and France; one may obtain a general understanding what was going on there from some excellent recent historical studies.15 limitation, not so easily remedied: I have mostly used published rather than attempting to examine all the correspondence of the works, hundreds of biologists whose views on natural selection might be relevant. Another
as well study is limited in space community. Of biological
as in time: it covers only theAnglo course this includes community
Also, I have omitted some biologists who made important contributions to the were not on evolutionary synthesis but directly involved in research is an example. I have not examined the natural selection; Edgar Anderson of biologists who, as an anonymous referee of this laboratory notebooks book suggests, may have been testing predictions without saying so in their or published reports, investiga conversely, may have been "reconstruct[ing] so as to make them tions in an overly rational way" in their publications look like tests of predictions, "when in fact they had other designs." I can more only hope that other scholars, acquainted and their archives, will fill in this gap. with individual scientists
Br?mer et al. (2000);Harwood (1985);Hossfeld (2000); Junker(2000, 2002,2004); Junker and et and Hossfeld Laubichler and Reif al. (1999); (2006); (2000,2002); Junker Engels Mayr (1999); (2000). For Russia, seeAdams (1970) and Kolchinsky (1999, 2000a, 2000b). Krimbas (1995)
provides on influences of Dobzhansky survey of the direct and indirect personal comprehensive see same in in several also the other articles the book countries; (Levine 1995). Jona biologists was writes that all of these works refute "claims that the than Harwood evolutionary synthesis an achievement" (Harwood 2002, 369-70). largely Anglo-American a
15. For France,
see Buican
(1983,1984,1989)
and Grimoult
(2000,2001).
For Germany,
see
Fisher: A New Language forEvolutionary Research
the modern theory early reception?or perhaps nonreception?of was the conditioned of natural selection presentation of by strongly The to most Anglo-American bi the theory in a language incomprehensible am not to the fact that some features of the I important referring ologists. evolutionary Chetverikov were in Russian by Sergei S. originally published synthesis mean I his that Fisher es and (1926) colleagues. Rather, andWright, reasoned and wrote and to a lesser degree Haldane
pecially, in the language cannot
biology, look up individual words
of nineteenth-century theoretical physics. That language into of be translated the language easily early twentieth-century as in because any language, one cannot simply understanding in a dictionary; one must learn something about
the background knowledge and styleof thought of thosewho use the

language.16
was far 16. As noted by J. F. Crow, among the founders of the evolutionary synthesis, "Fisher He solved very difficult problems, and and away the most skilled and inventive mathematically. "is the most important book on evo almost always with elegance and grace." His GeneticalTheory in Bennett But "his papers could Kimura, May 3,1956, (1983,229-30). most mathematicians, with the result that they were often not trusted by it clear what his assumptions were, when and how he was approximating, Fisher hardly ever made A more danger and how to get from one equation to the next" (Crow and Dove 2000,142,144). letter from Fisher not be understood toM.
lutionsince Darwin" (Crow 1992c,99); see alsoCrow (1990b) andEdwards (2000). See also the
a desirable result: "Discussions ous aspect of Fisher's work iswhat Haldane of Dar considered winism will come to be classed with circle-squaring ignorant of mathematics by persons wholly and earth-flattening" See also Ford (2005). (Haldane 1931,475). of Darwin's As hypothetical history, one might ask how the development theory and of post have if he had studied the works of Darwinian would been different Quetelet, Adolphe biology a statistical s law of s Malthus critique of population growth, and had adopted including Quetelet to natural selection (Ariew 2007). Instead, most meth biologists resisted mathematical approach "Hardy-Weinberg to be 'discovered'" law," which, (1988,473). as James Crow points out, "is so self-evident that it hardly needed
ods until after World War II.This is (to me) illustrated gave to the by thegreat significance they
CHOOSING
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be reliably computed. It may be helpful to note that a theoretical model does a scientistmay make the not make empirical claim that empirical claims, but a ismore or lesswell described by themodel. Mod particular phenomenon on the models ern based developed by Fisher, Haldane, evolutionary theory as an can be view of theo andWright example of the "semantic interpreted now than the in rather the of science, ries," (no-longer) philosophy popular
successful theoretical physicist often chooses a simple model, deliber world so dear to the natu ately ignoring the rich complexity of the natural can a set in whose mathematical of favor of consequences ralist, postulates A
"received" view of theories, according to John Beatty (1981). fact: he stud In Fisher s case it is essential to note one biographical ied theoretical physics, including the kinetic theory of gases, with James the following sentence, which appears halfway through the Jeans.17 Hence Natural Selection: first chapter of The Genetical Theory of The particulate theory of inheritance resembles the kinetic theory of gases with its perfectly elastic collisions, whereas the blending theory resembles a in which some outside theory of gases with inelastic collisions, and agency is required to be continually at work
to keep the particles astir.
(Fisher1958,11)
In the particulate theory, "elastic" interactions tend to maintain a statistical distribution (of speeds, or genes). The combination ofmany random causes may produce a single clear-cut effect.18Fisher's statistical approach implied that fluctuations from an average value become relatively insignificant for a
as the pressure of a gas at a fixed temperature in a large population, just fixed volume remains constant unless the density is extremely low. So when that random mating of and A. C. Hagedoorn A. L. Hagedoorn proposed organisms objected has a greater effect than natural selection, Fisher immediately s random effect" (a version of Sewall Wright that this "Hagedoorn
in June examination the Cambridge (1978, 33) writes that Fisher completed Tripos in Schedule B_Because he had been as "a in the with distinction paper optical Wrangler in October for a year of in he was able to return to Cambridge awarded a studentship physics, the theory and under statistical mechanics and of James Jeans quantum theory graduate study 17. Box 1912
is found in the Fisher in a 1927 lecture at Leeds, a summary ofwhich theory of gases of the influ further For discussion this Hall for Leeds thank file information). (I papers, Nancy and Mor ence of statistical see and Weber 10); (1995, (1992); chapter Hodge Depew physics, His published papers are reprinted in Fisher (1971-1974). rison (1997,2000,2004). the kinetic
M. Stratton." Jeans authored a standard text {The Dynamical of errors under F. J. Theory ofGases) a and popular books on science, contributed to the theory of black-body radiation, and proposed with selection natural Darwinian also compared theory of the origin of the solar system. Fisher
18.Fisher and Stock (1915, 60) andFisher (1922,321).
10
Fisher: A New Language for Evolutionary Research
genetic drift, discussed
tions (Fisher 1921,1922). Fisher (1930a) had already Darwins theory based on pointed out that in
inheritance,
next) must be insignificant
except in small popula
blending
bisexual reproduction will tend rapidly to produce uniformity [and there fore] if variability persists, causes of new variation must be continually at work; [and hence] the causes of the great variability of domesticated ... species, of all kinds and in all countries, must be sought for in changed conditions and increase of food.19
and/or envi So, the variability must be maintained by frequent mutations ronmental changes in a theory based on blending inheritance.20 or was essential to evolution be increasing the variability Preserving cause ofwhat Fisher called the "fundamental theorem of natural selection." Fisher was muddled, and the theorem itself original proof of the theorem as is it as literally incorrect printed; evolutionary biologists simply accepted no serious with technical discussion until many years qualitatively correct, later. I will therefore simply state the original without commenting on its derivation: "The rate of increase in fitness of any organism at any time is equal s
to its at that time."21 genetic variance in fitness to R. Price (1972), the theorem is correct if one de George According due to natural fines "change in fitness" as change in the fraction oftotal change one a not corrects and if the effects of environment, selection, large including
number of errors and ambiguities. There have been many other attempts to means and how valid it is,22but for my purposes, explain what the theorem the qualitative statement that the speed of evolution is an increasing func
in brackets were inserted in the 1958 edition; see Fisher (1999). Fisher evi words to felt the need that these statements, originally presented as a list of separate dently emphasize were connected. postulates, logically 20. He did not mention Maxwell's velocity distribution, much less discuss itsvariance-preserving property, but probably took it for granted that anyone familiar with the kinetic theory of gases 19. The
(e.g., anyone who studied for the Cambridge University Mathematical see the of the between gene frequencies point immediately analogy et al. (1986,9-11,281-2,288-95,320-1). Garber 21. Fisher variable
Tripos examination) would and molecular velocities. See
theorem as stated is dimensionally incorrect, since it has a time (1999, 35). The on the left but not on the one is to assume that time ismea right; presumably, supposed sured in number of generations. For further discussion of the theorem, see Price (1972); Olby a (2006). Ayala (1965,1968a) provided qualitative experimental verification. On the ambiguity of the concept of fitness, see Depew andWeber (1995); Ariew
(1981); Endler (1986, 28-29); Leigh (1986); Ruse (1996, 316); Crow (2002); and Plutynski
and Lewontin
articles H. Beatty and by Sober,3-39]). (2004); Sober (2006 [reprinting by S.K.Mills and J. 22. Lessard (1997); Plutynski(2006); andworks citedby theseauthors.
11
CHOOSING
SELECTION
undertake
use of the he called attention to the misleading in the evolutionary literature. This for existence" struggle phrase "struggle on the "excessive was of blamed offspring, supposedly production usually This man to be observed throughout organic nature" (Fisher, 1930a, 46). as a cause of on ner of overpopulation speaking, which places the emphasis attention the impor from diverts between individuals, away competition tance of the quantitative aspects of reproduction: understanding natural selection when There is something like a relic of creationist philosophy in arguing from the observation, let us say, that a cod spawns a million eggs, that therefore its offspring are subject to [njatural [s]election; and ithas the disadvantage of excluding fecundity from the class of characteristics ofwhich we may attempt to appreciate the aptitude. It would be instructive to know not a just apportionment ismade be only by what physiological mechanism tween the nutriment devoted to the gonads and that devoted to the rest of the parental organism, but also what circumstances in the life-history and environment would render profitable the diversion of a greater or lesser share of the available resources toward reproduction.23 s view
of genetic variance is sufficient; it is not necessary to a technical discussion of the theorem here, since the reception of natural selection before 1970 did not much depend on the theorem. Fisher introduced another important feature of the modern theory of tion of the amount
Fisher (1942,
tionary battle with other organisms to a definition of fitness in terms of differential success (similar remarks inThomson 1920). Ac maybe found reproductive toDarwinism to Peter J.Bowler (2004), T. H. Morgans opposition cording "was dictated by his moral The objections to the picture of a world dominated
of the operation of selection, endorsed by Dobzhansky the general shift in the conception of "evolu 397), foreshadowed fitness" away from survival of the individual organism in a brutal
shift away from a "struggle for survival" to "struggle to reproduce" was to prove useful in the harsh image of natural selection, combating as well as for "social blamed by some critics for the excesses of the Nazis famous line "Nature, red in tooth and claw" (1850, Darwinism."Tennyson's
(Bowler 1978,55). by struggle"
but section LVI, fourth quatrain) was frequently quoted. Less memorable s statement in his 1893 Romanes was T. H. more Lec Huxley appropriate ture: Survival of the "fittest" does not mean survival of the "best" in the ethical sense. On the contrary, "The existence struggle for tends to elimi
Fisher (1930b). 23. Fisher (1930a,47); cf.
12
nate those less fitted to to the circumstances of their exis adapt themselves tence. The strongest, the most self-assertive, tend to tread down theweaker" writers invoked the counterculture
im sexual rather than asexual reproduction is quantitatively Moreover, to to out while Fishers that He evolution, according portant theory. pointed natural selection would allow an asexual organism to evolve, an otherwise similar sexual organism would evolve much faster, depending on "the num are ber of different loci in the sexual species, the genes in which freely a sexual two Even with genes would organism interchangeable.... only 24.Moody (1962, 355); Rhodes (1962, 277); Savage (1963, 50); C. Wallace (1969, 125); Weisz (1965, 430; 1966, 352-4; 1968, 196).According toGerson (1998, 356), this shift began
after World War a clerical now to Ernst I.According "Travel Notes, 1954": "In Germany? Mayr's unpublished movement is particularly likeMc state?the anti-evol[utionary] strong_Just was war thus evolution liberalism and after the communism, synonymizes synonymized typological selectionism, (1963,
not war" slogan "Make love, by calling natural selection a "peaceful force" that promoted sex rather than a destruc tive one that promoted death.24
(T.H. Huxley 1947, 81-82). But in the 1960s, someAmerican textbook
Carthy with the most
225). For a contrary view, see Huxley in (1993,240-3). Dronamraju C. H. Waddington, commenting
and biology with Nazi racism" (quoted by Junker 2002, and his letters to Simpson introduction) published
on the statement of T. H. in the text,wrote Huxley quoted so to us that "he was writing under the spell of that extraordinary incomprehensible impulsion, to transmute the of the major contribu today, which forced the Victorians simple mathematics a biology into battle-ground the conclusion that evolution
for their sadism" (Waddington 1942,17). Wad to "an increase in bloodiness, fierceness and leads dington rejected self-assertion." On the contrary, he asserted, "We must accept the direction of evolution as good it is good according to any realist definition of that 1942, simply because concept" (Waddington to intimidated by the tendency of evolutionists 18). Karl Popper, "feeling somewhat suspect any one of obscurantism who does not share their emotional attitude toward evolution"?exemplified assertion?conceded, by Waddingtons tion of the relevant facts" (Popper "I see in modern 106-7). This Darwinism concession 1960, the most successful explana from did not prevent Popper
tion of theoretical
status of Darwinism (see chapter 21). denying the scientific to Dar Fisher (1947) ascribed the late nineteenthand early twentieth-century opposition an was felt to be "a harsh and ruthless sort of to winism it sources": theory" and "psychological determinism. But now, he wrote, the physicists have liberated us from example of materialistic determinism
causes "lie in and we can recognize that natural selection is a creative process whose the day to day incidents in the innumerable lives of innumerable plants, animals and men"?not "set inmotion on 'the firstmorning of creation or determined by heredity or environment" (Fisher
1947,619,620); see alsoFisher (1950).

Fisher's concern with which
is of course directly related to his interest in eugenics, reproduction was a motivation to B. Norton for his research on natural selection; (1978,1983) according it takes up about a third of the text of his Genetical remarks: "Curiously, factors that Theory. Mayr success were contribute to reproductive largely neglected by evolutionists until around 1970. At that time naturalists rediscovered Darwin's that females may play a important finding (1871) role in the choice
of their mates" He attributed the earlier neglect to (Mayr 2004,139). as the than the "eliminated the rather individual the of selection," which gene target "adopting difference between natural and sexual selection" (Mayr 1992a, 17). decisive
13
CHOOSING
SELECTION
a manifest apparently possess from an approximate doubling [njatural [s]election."25 A result that illustrates more tion is that large mutations can contribute mutations Let point A space, where are
over its asexual advantage competitor of the rate with which it could respond sharply Fisher smathematical
... to
represents the value of some attribute of the to students of the kinetic is like the organism. (This "phase space" familiar at distance d from Ay represent the set of val theory of gases.) Let point O, ues of those attributes to the best to the corresponding possible adaptation environment. A mutation closer to O. Suppose will be favorable mutations are random
represent each dimension
sophistica or lethal, so only small generally nonadaptive was to evolution. Fisher s reasoning geometric: the position of the organism in an ^-dimensional
moves the organism only if it correlated with (not adaptive a mutation to is is that favorable ness).The probability simply proportional the amount of space that is closer to O than A is. For those who think in three-dimensional space, it seems reasonable that for a very small mutation?moving the organism a distance r away from A?the chance of ending up closer to O is nearly 50 percent, since the surface of the sphere centered at O is nearly a plane. The organism is more to move farther away from the slightly likely sphere. On the other it?an unfavorable mutation. If r isgreater than 2d (the diameter of land inside the sphere; the probability of a favorable thinks in ^-dimensional
hand, ifr is large (compared to d), it may go throughthe sphereand land

outside the sphere), it cannot mutation is zero. For someone
space, it is obvi ous that the same argument even greater force: the greater the applies with number of dimensions of the (attributes organism), the smaller the prob like Fisher who a course itwas ability that large mutation will be favorable.26 Of precisely the effort to think outside the three-dimensional box of everyday experi ence that was one of the driving forces of early twentieth-century out, the idea that most mutations
and art (Miller2001).

As Fisher pointed
physics is not
are harmful
to J. F. Crow, "Muller shares with R. A. Fisher 25. Fisher (1930a, 121-3). According (1930) statement of the value ofMendelian credit for the first convincing segregation and recombina even of sexual reproduction, tion," as illustrated by this explanation though "the Muller-Fisher idea is no production,
of life"(1992c, 85, 88 [citing Muller 1932]). history
as the reason for the evolution and maintenance of sexual re major longer regarded at least in multicellular at an eukaryotes, although itmay have been early time in the
26. Fisher (1999, 38-39). It has been suggested that Fisher developed his facility for visual Crow (1990c) writes that space because of his poor eyesight (Box 1978,15). izing ^-dimensional "Fisher had an insight into multidimensional geometry that was little short of occult" (Crow and 2000,142).
Dove
14
Fisher: A New Language for Evolutionary Research was as obvious original but "regarded by the naturalists" who believed that are to their environments. It was the well organisms generally very adapted to theory that made it necessary popularity of the de Vries large-mutation reinforce that view.27 seem If natural selection acts primarily on very small mutations, it might that itmust be a very slow process. But Fisher showed that a rare gene with
in every generation. For example, if the selective advantage is 1 percent (and one rate in one thousand million is then "in the mutation [per generation]), a a come to in in which each 1,000,000,000 species generation maturity, a one rate in mutant mutation in every of 1 thousand million will produce
a small selective small mutation that produces no ob only advantage?a a servable effect in the first generation?can spread rather quickly through same a at mutation is repeated if the finite rate) population (randomly but
[on average], and thus establish the superiority of the new type generation in less than 250 generations, and quite probably in less than 10, from the first occurrence of the mutation, whereas, if the new mutation started with rate of 1 in 1,000,000 thewhole business would be settled, with a considerable margin to spare, in the first generation." Nat is very small.28 ural selection will be much less effective if the population familiar mutation Fisher's theoretical conclusion that small mutations?those whose ef the more
more in one generation?are important in confirmed by later experiments. An example was the (Fisher s first genetical assistant when he study by Kenneth Mather became Galton Professor at University College London) and L. G. Wigan. They found that the several small mutations that combine to determine a polygenic character, such as the number of chaetae (hair or bristles) on the
fects are not easily observable evolution than large ones was
fluctuations. Under ous
abdomen of aDrosophila fly, masked by produce an effectthat is initially

or a sudden jump change
Wigan
Mather 1943). 1942; see also
the influence of selection, theymay produce a continu that mimics a macromutation (Mather and effectiveness of natural selection was first revealed in 1915 used a (Nor
quantitative calculations by H.T. ton and Punnett citing is a mechanism without a
The
in
J.Norton, published by R. C. Punnett 1915). In later years, many evolutionists
source but attributed to Fisher: "Natural specific for generating a high degree of improbability."}. S. Huxley Fisher where he had published
phrase selection this, to

of mim 109 in the to adopt
reported
that he "asked Professor
the phenomenon (1999,41). Turner (1985) discusses how Fisher handled see considered evidence for also (1987). strong icry, formerly large mutations; Leigh itmeant 28. Fisher (1999, 78). At that time, the word "billion" was ambiguous; United States, but itmeant 1012 in the United Kingdom, which subsequently decided the American definition.
27. Fisher
15
CHOOSING
SELECTION
which
he replied that he had merely stated it verbally at a meeting. How ever, it sums up the situation so pithily that I feel it should be recorded in Later, Fisher printed his own version of the 1951,595). print" (J. S. Huxley statement: natural selection is the "process by which contingencies a priori are an increasing until given, in the process of time, probability, improbable it is their nonoccurrence rather than their occurrence which becomes highly improbable."29 The above results show that natural
selection may be an effective cause of evolution but does not appear to lend itself to empirical tests in a short or the mutation rate is large. period of time unless the selective advantage situations inwhich stable gene ratios are maintained Fisher also discussed a balance of selective forces: "One gene has a selective advantage only by is established, while for higher ratios it is at a until a certain gene-ratio In such cases the gene ratio will be stable at the selective disadvantage.
in action will tend to restore it to this value limiting value, for the selection it happens to be disturbed from it in either direction." Of course whenever the conditions of stability must themselves be transient during the course of evolutionary change, but the temporary stability provides a good oppor
over both a selective situation, gote has homozygotes."30 This advantage later known as a "balanced polymorphism," was investigated extensively by Fisher sOxford colleague E. B. Ford and his students and provided impor tant evidence While Fishers for the validity of the theory. theory is sometimes described to Darwin's Men application of selection, he also proposed as an
indicates that tunity to study in detail how natural selection works. Theory "a single factor may be in stable equilibrium under selection if the heterozy
a theory of natural in Mendel's theory. The attribute of genes called domi significant change not fixed, but was itself a result of evolu nant or recessive was, he argued, delian genetics
to be of great importance, it seems to have had no influence on hypothesis the reception of his general theory,with a few exceptions (e. g., J.Huxley 1947,167). An excellent account of the hypothesis and its empirical test is
tion (Fisher 1930a, chapter 3). Although Fisher himself considered this
given in chapter8 of thebiographyby his daughterJoanFisher Box,who

concludes: nance
of domi "Among geneticists today the concept of the evolution it has played a significant part in is almost unknown. Nevertheless, in respect of evolution by natural selection. transforming scientific opinion
29. Fisher enormous Muller
(1954,
91). I thankWill
Provine
degree of that improbability was, in 1929," but he did not citeMuller's (1930a, 99-100); see Fisher
himself stated that "the for this reference. Huxley in a I think, first clearly demonstrated general way by publication. for an earlier statement of this principle.
30. Fisher
(1922)
16
of natural research
It has opened thewhole field of ecological genetics bywhich the efficacy

selection indicated (Box 1978, 233). Later fully demonstrated" is not supported by that Fisher s theory of dominance has been
empiricaldata (Skipper2000,105-15).
Fisher cofounded Heredity: An International Journal of Genet Darlington, ics. did most of the editorial work, Fisher "was con Although Darlington a was tent that Darlington providing inHeredity ready platform for the selectionist crusade of his prot?g? E. B. Ford and Fords students Bernard
are indifferent or hostile. In 1947, together with theBritish cytologist Cyril
a new field, it is to be able to In developing important publish one's results quickly, even when the editors and referees of established journals
Arthur Cain, and Philip Sheppard." (Harman 2004,209-10). Kettlewell,

Readers seeking a "social construction" interpretation of Fisher s
related to his eugenic and religious views.They should also inspect Fishers own text to check the accuracy of Norton's assertion (1983,27) that Fisher was not interested
should consult the papers by BernardNorton (1978, 1983), Robert Olby Moore (2007) for discussion of how Fishers theory is (1981), and James
theory
he came first alphabetically or chronologically (his firstmajor con in 1918), but because he established a tribution was published simple prop natural selection acting on small osition, subject to certain assumptions: mutations can produce a significant change in the genetic composition of a because was a population much faster than previously believed. Thus, theory based on natural selection to is be able therefore worth evolution and may explain criticized Fisher s model (as defined by his assump pursuing. Biologists
species (our own) moons cuss an speciation, suggesting analogy with the fission theory of the son origin proposed by Charles Darwin's George Howard Darwin.31 In summary, Fisher was the most important of the three founders, not
in speciation because his motive was to improve one rather than change it to another. In fact Fisher did dis
tions) for being unrealistic?"bean bag genetics."32 But from the viewpoint of theoretical physics, that is not a criticism at all. It is often by starting with an idealized model whose we can be properties accurately calculated that can start to understand how nature works.33 Of course it takes a while for
to and that, as used by Fisher Wright, of our thinking ..." (Mayr 1992b, 21). former career as a theoretical ized models, whereas Fisher
31. Fisher (1930a, chapter6); Olby (1981,256-7); and Brush (1996a, chapter 4.2). withMayr (1959),who admittedthat itdid not apply 32.The phrase apparently originated
and Haldane, itwas "a necessary step in the development
33. Lewontin (2000) and Plutynski(2001,2004, 2006). Full disclosureon thispoint: inmy
physicist, I followed just this approach, working some to test his theories experiments actually did only with ideal et al. (e.g., Brush
1966).
17
CHOOSING
SELECTION
other scientists to be persuaded that this approach can work in biology, and so it was very fortunate that Fisher attracted followers like E. B. Ford who could apply and propagate his ideas. The previous paragraphs may give the impression that Fisher was an armchair theorist who left it to other scientists to do the empirical work needed to test his theories. But he did take an active role in empirical re
search, by himself and in collaboration with E. B. Ford and others, as may be seen from the list of his papers on genetics, evolution, and eugenics at the beginning of his Collected An early example is the confirmation Papers ?A of a prediction, and quantitatively derived suggested by Charles Darwin from Fisher s
it "pushed evolutionary synthesis," because to the to into it attention the field" natural "drew study people populations; selective adjustment of the effects of genes." in Fisher 's fame as a leader in the development of statistical methods
shouldbe proportionalto the size of itspopulation (Fisherand Ford 1926, was 1928).According toFord (1980,338), Fishers 1927 paper onmimicry
"the true start of the modern
theory of natural
selection,
that the variability
of a species
NSH; his method of double-blind random trials is often called the gold
in medicine standard for testing new substances and procedures and ag of science, his critical riculture.35 For some historians and philosophers data showed that one can never assume that empiri scrutiny ofMendel's cal results are independent of the theoretical bias of the observer; Fisher's statistical analysis to be true.36 suggested thatMendel's published results are too good
research undoubtedly
helped
to call attention
to his development
of the
34. Fisher
therein.
35.N. L. S.Hall (2007) andBox (1978). 36. Fisher (1936a); fordiscussion,seePilpel (2007), Franklin et al. (2008), andworks cited
(1971-1974,1:14-17).
18
Wright: Random Genetic Drift, a Concept Out of Control
Sewall
tance of size in evolution.37 His earlier work on inbreeding, as population of the U.S. of Agriculture, indicated that ran employee Department dom fluctuations are important in small populations. They will generally an tend to decrease the
in his comprehensive 1931 article, showed that many of Wright, his results were consistent with those of Fisher but stressed the impor
fixed (Wright 1931,158). In such a smallpopulation there is
heterozygosity (presence of different genes; e.g.,^f and a at the same locus on chromosomes of an organism) at a corresponding rate means that traits to the size N. This inversely proportional population determined by the homozygous genes (e.g., A andî) will tend to become
little variation, little effect of selection and thus a static condition modi fied occasionally by chance fixation of new mutations degeneration and extinction. On the other hand, leading inevitably to
in a very large interbreeding population,
there is
great variability but such a close approach to complete equilibrium of all gene frequencies that there is no evolution under static conditions. The most a favorable situation for evolution divided
Complete
is into partially isolated local

case ... originates new
large population,
of small size....
and subdivided
isolation
races
in this
species differing for the most part in nonadaptive
respects, but is capable
37. with otherpapers in Wright (1931), reprinted Wright (1986). See the comprehensive biography byProvine (1986) and articles byCrow (1994),Hill (1990), and Park (1991).
19
CHOOSING
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of initiating an adaptive radiation as well as parallel orthogenetic lines, in accordance with the conditions. (Wright 1931,158) of Genetics (in Ithaca, New York, Congress a more version of his theory. shorter, 1932), Wright presented qualitative This, according to his biographer William Provine, "was probably the most to show introduced diagrams influential paper he ever published."38 He At the Sixth International various hypothetical "fields of gene combinations." The species will gener a a region around ally occupy "peak" representing the optimum combination of gene frequencies in a particular environment. Isolated small local popu lations will evolve by random genetic drift: The chances are good that one at least will come under the influence of another peak. If a higher peak, this race will expand in numbers and by
new crossbreeding with the others will pull thewhole species toward the position. The average adaptiveness of the species thus advances under in tergroup selection, an enormously more effective process than intragroup
selection.39
to describe electrical and Faraday qualitatively by Michael magnetic phenomena but without the mathematical sophistication of James or R. A. Fisher. Clerk Maxwell "fields" used Robert A. Skipper Jr.has given a comprehensive philosophical-historical as follows:40 analysis of Wright's theory and summarizes it Phase causes genetic drift subpopulations within the global population to lose fitness. I. Random semi-isolated
evo s was called the "shifting balance" process of Wright general theory lution. It is another example of introducing the language of nineteenth century theoretical physics into twentieth-century biology: it invokes the
Phase
II. Selection on complex genetic interaction systems raises the fit ness of those subpopulations. III. Interdemic selection then raises the fitness of the large or
Phase
global population. 38. Provine in Wright (1986,97). Ruse (1996b) arguesthat Wright's use ofdiagrams,though
logically necessary, helped to persuade other scientists to adopt his theory. See also Skipper
not
(2004). inBrousseau (1967, 68-78 [quotationon 75]). 39. Wright (1932); reprinted 40. Skipper (2000; 2004,1177).
20
a Wright:Random GeneticDrift, Concept Out ofControl comments:
James Crow
to drift across a valley to a point where it came under the domain of at traction of a higher peak. The contrast between Fisher and Wright is that mass selection to carry a the of inability Wright regarded population from one stable state to another whereas Fisher believed that everwould hardly a a no in in that al?ele be such population position change frequencies s Wright says intergroup selection he really had inmind directional migration be
subgroups.41
as a way of It is important to realize that Wright thought of his process one are unfit. state to intermediates from another when going adapted In his metaphor, random drift could occasionally cause the population
could cause an increase in fitness. Fisher thought that Wright were more likewaves and an ocean_When valleys troughs in
tween
peaks and
This mechanism "an essentially
for evolution
following
was used Provine (1983), favoredgenetic drift because it Wright originally

by several evolutionists
species.
one" although nonadaptive as the "Sewall became known genetic drift, Wright to explain nonadaptive
called isolation, which Wright it involved selection as well as effect."42According to
differences between
similar
statements that he did not think genetic drift Despite Wright's frequent by itselfwould produce evolution in the absence of selective forces, his ef
on an earlier version of this book, reviewed for the American 41. Crow, commenting Philo see also Crow et al. (1990). sophical Society, June 2005; 42.Wright and Provine Provine credits the Hagedoorns (1983,1985,1986). (1986,168-9) in 1921 (A. L. Hagedoorn with pointing out the importance of random fixation in evolution and A. C. Hagedoorn (1931) recognized unknown 1921; Wright 1986,108). Apparently a similar effect (see also Dubinin and Romashov (1970,231). Even earlier, H. Muller J. (1918) to Wright, N. P. Dubinin subse 1932), and itwas "drift" under
See also quentlyoccasionally called the Wright-Dubinin effect. Wright (1931);Mayr (1963,
204);
in Muller (1962); and longername. See his paper in Huxley (1940,185-268), partlyreprinted
see Lewontin Here's (1974, an example 87-90). of an
and Dobzhansky
introduced
and pairing of "The accidental meeting early textbook explanation: individuals will usually result in some deviation from the expected result. A gene that ought theo to occur in 25 per cent of the individuals may to be in 28 per cent or retically only easily happen in the next generation hap 22 per cent solely by chance. Should the deviation from expectation pen to be in the same direction, the difference is accentuated. Different parts of a range may thus to the same still belonging by groups of individuals which, while species, even a have their genes in different proportions_Possibly divergence great enough to mark two separate manner. species may take place in this purely random Beyond this degree of to be differentiation probably other factors enter. The most important of such factors is believed nevertheless natural selection." (Shull et al. 1941, 355). come to be inhabited
21
CHOOSING
SELECTION
explain (supposedly) himself 1940, 324). may have en nonadaptive Wright as that the distribution of couraged this incorrect usage by statements such the three allelomorphs for human blood groups is "apparently nonadaptive" characters (Lack
feet and his term "drift"were most
often invoked
to
(Wright1931,127) and that
in the human species, the blood group al?eles are neutral as far as is known. The frequencies vary widely from region to region and in such a way as to indicate that the historical factor (i.e., partial isolation) is the deter The frequency distribution indicates a considerable amount mining factor. of random differentiation even among the largest populations.
(Wright
1940,179)
The later used, perhaps unfairly, as a s entire refutation of Wright theory of evolution (Provine 1986,456). arose A further confusion from the statistical nature of natural selection refutation of these statements was itself. If one defines distinguish between "fitness" as reproductive success, then it is difficult to drift as a sampling error and natural selection.43
was fairly balance theory United States Wright s shifting popular in the was before 1950, partlybecause it adopted byDobzhansky and partlybe
cause his were found to be useful in improving agri inbreeding formulas cultural production. For example, poultry breeders were able to increase the
number of eggs laid by hens, an achievement that makes Wright indirectly are in for the fact that the United States eggs responsible relatively cheap s contribution to statistical method was small today.44Although Wright to Fisher's, his compared "path analysis" technique social sciences in the 1970s (Crow 1990a). became popular
in the
a series of three papers reporting experiments done byWar published a visitor from Brazil supported by a Rockefeller Foundation wick E. Kerr, of males fellowship. Very small populations Drosophila melanogaster (four more were or and four females in each) isolated and followed through ten generations. Selection was assumed strong enough to overwhelm
The careerof genetic driftreached a high point in 1954whenWright
to be present but estimated not to be the effects of drift in such small populations.
43. This comprehensive
and other theoretical and mathematically
problems
were arising fromWright's work Gustav Malecot rigorous viewpoint by
treated from a more in France (Malecot
1966,1969;Nagylaki 1989). 44. Lerner andHazen (1947); Lerner (1958, 117, 223-54); and Snyder (1940, 305) For
Wright's views on progress in relation to evolution, see Ruse (1996a, 367-85,401-6).
22
a Wright:Random GeneticDrift, Concept Out ofControl In each of the three experiments, the frequencies of a specified mutation (forked, bar, aristopedia, and spineless) competing with its al?ele were de termined for about one hundred lines. The results (including the eventual fixation of one mutation) confirmed the theoretical predictions.45 According to James Crow:
Kerr s experiments were done while he was a postdoc inmy lab. I chose the markers, we planned the experiments together, and he carried them out here. Because of his interest in bees we emphasized theX chromo some to mimic a X-linked balanced course moved
chose one nearly neutral species.We haploid-diploid gene, one strongly selected X-linked gene, and one autosomal sure how to polymorphism. I wasn't analyze the data and of
thought of Wright. By the timeWright got involved Kerr had s lab at Columbia. toDobzhansky I think Dobzhansky also sug ex were gestedWright. Actually, when Sewall and I talking about these came toMadison not I he had known after he found that periments
where
of this book,June2005)
The
the experiments were done. (Crow, in review of an earlier version
were an to important contribution Kerr-Wright papers population s not correct the did of but genetics Wright theory by misunderstanding effects of selection supporters tive to natural in designing and analyzing the experiments, even his to assume that he believed drift to be an alterna
otherbiologists.46 Despite the fact that Wright had explicitlyincluded the

continued
selection. For those who read the Kerr-Wright papers more the carefully, extremely artificial conditions of this laboratory experiment probably reinforced the criticism that genetic drift,while theoretically pres ent in any is unlikely to play a significant role in evolutionary population, processes in nature.
attempt to clarify and perhaps revise his position, have been treated Wright's in detail by Provine, and the continuing controversy has been analyzed from a of biology still philosophical viewpoint by several scholars.47 Philosophers can drift defined each other about whether be with argue unambiguously
of genetic drift, and The attacksby Fisher and Ford on Wright's theory
45. Kerr and Wright andKerr (1954). Wright (1954a, 1954b) and 46. Lewontin (1967, 64) and Provine (1986,456). 47. Provine (1985,1986); Hodge (1987,1992); Skipper (2000,2002);Millstein (2002,2005, on the some useful insights 2007a, 2007b); and Plutynski (2005, 2007b). Turner (1987) offers
subject.
23
CHOOSING
SELECTION
and distinguished from natural sidered causes or effects.48
selection and whether
con they should be
driftand/or the Sewall Wright effect:

1. Evolution
In surveying the response to this controversy by other biologists, I found it necessary to distinguish between two hypotheses, both known as genetic
then are incorporated into others by natural selection. In this case, genetic drift has evolutionary significance only in combinationwith
natural selection.
large populations, divided into several partially isolated smaller populations, inwhich certain characters first appear in one subgroup as a result of genetic drift and
occurs fastest inmedium-size
or
2. A nonadaptive character may be explained as the result of genetic drift rather than natural selection.
meant (1) and rejectedthe ascriptiontohimselfof (2), even Wright usually were supporting Motoo Kimura) thoughtthey who (like Wright's by those
was his analogy ofMexican theory.49 Perhaps his best explanation of (1) on a surface with several hollows of varying beans depths (Wright jumping hollows correspond to adaptive peaks, but gravitational force re 1960).The can move from a shallow place places selective force. The only way the bean
to of the kinetic theory of gases (or more precisely of statistical mechanics) formulate this physical analogy, with thermal fluctuations playing the role of genetic drift inmaking possible the transition from a shallow potential well his idea to Fisher more deeper one, he could have communicated still have objected that in a multidimensional effectively. But Fisher would to a
to a deeper place is by randomlyjumping around; gravity(selectionpres will just keep it where it is.Perhaps ifhe had used the terminology sure)
s is vague and incoher Wright shifting balance theory literally, or as a it is understood heuristic best ent;50 qualitative metaphor perhaps are even its of But "some 2004). underlying assumptions though (Skipper were and Charles J.Goodnight viewed as contradictory," Michael Wade Taken
all gene ratios (Fisher 1941,58).
one space of gene frequencies, there is no reason to think there is even posi an maximum for tion that represents absolute minimum fitness) (meaning
48. Brandon (2005);Millstein (2006, 2007a); Plutynski (2007b); and Reisman and Forber (2005). 49. Moore (1953,228) and Wright (1951,1955). 50. Fisher (1941,1953); Coyne et al. (1997); Provine (2001); andRuse (2004).
24
a Wright: Random GeneticDrift, Concept Out ofControl able to design an experimental theory with favorable results.51 test of the interdemic s
phase
ofWright
most ing difficulties of natural selection theory: the fact that the prominent visual differences between closely related species seem to have no adaptive value. How then could natural selection explain the differences between objection G. C. Robson and was them? This was made most
(2) was much more popular among biologists, not only be Hypothesis cause it was to solve one of the outstand simpler, but also because it seemed
must
and effectively by O. W. Richards a reason for the reluctance of some major probably to accept natural selection as an in the biologists explanation for speciation andWright 1930s.52 Since the theory of Fisher, Haldane, implied that there be genetic drift in sufficiently small populations, those who did not understand the various ramifications of the theory assumed that itwas le character.53 But gitimate to invoke genetic drift to explain any nonadaptive since his theory lost credibility when the result was often unfair to Wright,
(and probably would not have made). The popularity of genetic drift in version (2) was widespread publicity about randomness in quantum play dice with atoms
was judged by the outcome of predictionsthathe himselfdid notmake it

also due in part to the could physics. If God
(an assertion thatAlbert Einstein helped to publicize it),why not with genes and chromosomes, which must be by denouncing made of atoms? A corollary of the unpredictability of atomic motion was
the unpredictability of biological phenomena, which was translated into a must make justification for relaxing the requirement that scientific theories
testable predictions.54 As Wright and others pointed out, he was not the inventor of the concept itwith a handy name at an opportune of genetic drift; he just proposed moment. and decline of Somehow the phrases "accidental multiplication mutant and (Dubinin genes" (Muller) and "genetic-automatic processes" on. as not the "simul did But Mark B. catch Adams Romashov) suggested, taneous
in 1931 could reflect broader and Dubinin discovery" byWright such "intellectual currents of thought" at the time (Adams 1968, 26). One is the includes the indeterminism of quantum mechanics, current, which revival of holism and antireductionism in science in the in
neoromantic
51.Wade ing that paper 53. There a particular
52.Richards andRobson (1926);Robson (1928); andRobson andRichards (1936).

was also some and confusion those to between an characters environment that give in that are adaptive for an organism to one organism over another; see advantage (1954,4) makes itmore explicit.
see also Crow's and Goodnight (1991,1015); "perspective" review by and the comprehensive Wade (1992).
(1991,973)
accompany
Millstein (2007b).
54. Scriven (1959) seems
imply this; Villee
25
CHOOSING
SELECTION
know, drift was
controversy only up to 1970. Looking back from 1970, it appears that selec as those familiar with more recent tion had triumphed, but developments revived by Kimura and others in the 1970s significant part of the modern theory of evolution. and
return to this issue in period.551 chapter 22, but here I should again remind the reader that in this I follow the selection versus drift monograph,
terwar
is still a
55. Brush (1980);Harrington (1996); andLawrence and Weisz
(1998).
26
Haldane:
A Mathematical-Philosophical
Biologist Weighs in
first establishing that natural selection could account for the evolutionary was on now.56 to Sahotra Sarkar (2004), Haldane change going According the first to propose a true synthesis of population genetics with classical and genetics, cytology, paleontology, evolutionary theory (see also Carson
biolo 1932) seem to have had the greatest effect in persuading ear gists that natural selection could account for evolutionary history, the in lier papers of J. B. S. Haldane, with have been credited 1924, beginning
works of Fisher (1930a) and Whilethe fundamental Wright (1931,
should be one that attempts future prediction, fies."57 In particular, Haldane
theories were retrospective in intent; Hal 1980). "Fishers and Wrights dane swas purely prospective [intended to predict processes in the future] extent that an though it allowed retrodiction. To the evolutionary theory only Haldane swork quali
s analysis of gene frequencies inBis ton betularia (Haldane 1924) ... itsmost famous paradigm case_Haldane gave genetic Darwinism was being pumped proved that the black pepper moth then proliferating to in selection that matched data model, and hence ways up by pressure gave empirical support to treating natural selection in terms of changing
gene frequencies.58 Haldane mathematical s 1932 book, reader with of natural The Causes a ofEvolution, addressed the non
logical examples
a lively style, providing large number of bio same At the selection. time, he was anxious to
57. Sarkar 58. Depew
56. Sarkar (2004) andDepew and Weber (1995).

(2004, 1223);
Ewens (1979,1990).
he credits the retrospective/prospective
distinction
to Warren
John
andWeber
(1995,237-8).
27
CHOOSING
SELECTION
can take advantage of his own prestige as a mathematician, asserting, "I write of natural selection with authority because I am one of the three most about its mathematical 1932b, theory" (Haldane people who know 33; see also his earlier technical papers included in the appendix). But his many other activities in science and politics kept him from devoting his full attention to
a evolutionary theory after 1932.59 At times he seemed somewhat lukewarm supporter of the natural selection hypothesis, suggest more fit and an not organism ing that natural selection does always make a small was even a to to concede he drive extinction; may species willing Lamarckian ary momentum" Haldane
drift will be noted in chapter18.

was clearly more and theological osophical monism materialistic and as hostile whose
effect over long time periods and an orthogenetic "evolution in a few cases.60 His changing views on the importance of or in phil Wright as issues. He characterized his own position attacked the holistic doctrine of "emergence" interested than Fisher
erties of the hydrogen atom seemed much more complex than those of its parts (electrons and protons), yet physicists did not abandon the attempt atom in terms of the properties of its parts. "Wave mechan explain the ics thus represents the most serious inroad yet made on the doctrine of emergence, which attempts to set up barriers to the progress of scientific to interpretation" and "gives hope standing of evolution."61 As for the creationists who to biologists who seek a rational under
new entities to scientific progress. Some biologists postulated cannot be its from those of parts. The fallacy properties predicted of this approach was shown by the example of quantum physics: the prop
believe in (what is now called) intelligent their Haldane doctrine, asserted, is refuted by two hard facts: first, design, the extinction of many species; and second, the evolution of parasites that
reserves his most result that 59. Haldane (1932b, 13). Haldane emphatic praise for Fisher's as a result cannot be in stable selection the char distributed for of normally population equilibrium acters distributed. This vitiates a large number of the arguments rather sensational fact normally which are commonly used both for and against eugenics and Darwinism" 1932b, 196). (Haldane the importance of domi results are important, Haldane writes, but he underestimates Wright's "a
nance (Haldane 1932b,212).

On Haldane's sometimes
was
He M. life and personality, see R. W. Clark Smith (1988,1992). (1969) and J. confused with his father, J. S. Haldane, whose views on many subjects were quite to Gordon McOuat Winsor and Mary P. different (J. B. S. Haldane (1995), 1932a, vi). According ers took up the study of evolution because of his interest in religion and annoyed assertion "Darwinism is dead" to all theologians. by attributing the 60. Haldane (1932,12,23,108,136-7). some read
Haldane
61.Haldane (1932b, 156-7; 1932a,262).
28
Haldane: A Mathematical-Philosophical Biologist Weighs in lost their faculties and inflict pain on other animals. Could a
have
God havemade the tapeworm (Haldane 1932b, 159)?The common perfect

statement (still in the twenty-first century) frequently made by creationists that "natural selection cannot account for the evolution of a highly complex character" iswrong, and such "common-sense" "are often very conclusions need to use mathematics to get correct answers, Haldane
morally
doubtful"?we believed.62
62.Haldane (1932b,213); see McOuat and Winsor (1995).
29
Fisher-Haldane
Early Reception of the

-WrightTheory
The
a except among in Britain and America, most notably E. B. small group of their colleagues to Fisher, this was because of the unfortunate Ford at Oxford.63 According of the 1930s attracted little attention before 1935 influence ofWilliam tion.64 Fisher was
works ofFisher, Wright, andHaldane published at thebeginning
response came from the Bishop of Birmingham, who asserted that Dar win's theory was now completely confirmed: "Such statistical investigations as those inR. A. Fisher's recent volume are a triumphant vindication of the
ideas about evolu Bateson and his wrongheaded could not see that his also annoyed that T. H. Morgan on selection of own work had made a genetic theory of evolution based not but small mutations irresistible.65 only possible Ironically (in view of the supposed incompatibility of evolution and religion), the first positive
Shull
at the Sixth Fisher's
and articles by Babcock, L. C. Dunn and Landaur, R. R. Gates, R. Gold and E. Moore statement in the text is based on my surveys o? Ameri and H. F. Osborn.The schmidt, Harland, canNaturalist, Genetics, and Journal At the suggestion of Will Provine, I also examined ofGenetics. and Wright) of the sixteen invited speakers (other than Fisher, Haldane, selected publications International Congress of Genetics (1932). For a discussion of this congress,
63. Ford (1931) and Poulton (1931); see books published in theyears 1932-1934 byA. F.
see
"Of the 1500 copiesprinted [of Crow (1992b). According toBennett (inFisher 1999,foreword),
over one-third were sold in the first year. However, sales soon declined Genetical Theory], at the time not read and the last copies were not sold until 1947. Many biologists presumably did or understand Fisher's book." 64. Fisher, elaide.
of Ad letter to E. B. Poulton, undated (ca. 1930), in Fisher papers at University archive. I thank Nancy Hall for copies of this and other letters from the Adelaide October November 65. Fisher, letter to L. Darwin, 15, 1932, and letter to T. H. Morgan, was see still later Fisher Ten 11, 1932, in Fisher papers; (1932). years annoyed about Morgan evolution in the first quarter of geneticists who "discredited themselves" by opposing Darwinian on the century; he wrote to K. Mather February in in embalmed amber mind" nently Morgan's and Powell (1987). His eventual acceptance 5,1942, (quoted that "the ideas of this period are perma in Bennett 1983, 235-6). On Morgan's is discussed by Muller
seeAllen (1968, 1978,1980); Bowler (1978);Dobzhansky (1959a, 254); views about evolution,
of natural selection (1946).
30
Fisher-Haldane- Wright Theory Early Reception of the
potency of natural selection."66 Also ironically reluctance of the French to accept Darwinism), cussion of the mathematical
(in view of the well-known the most substantive dis published in Paris by large (and still preva
Philippe L'H?ritier (1934).

But the standard
theory
was
in a booklet
argument that natural selection without mutations could not originate new species was perhaps directed) that H. Muller's J. experiments
on the by production of small mutations on nature evidence the of such mutations.68 x-rays might provide important Mathematical derivations were admired but did not convince many readers not that they entailed convincing real biological consequences. Evidence from genetics was to, or even known by, naturalists who considered the field
lent among biologists throughthe 1930s.67Itwas only dimly recognized
more
important than the laboratory.69
ments
theory had become part of the discourse of evolutionists about mutations,70 was somewhat distracted to the although their attention theory by argu about Fisher Four major tion were in a state of flux. (East, 1935). theory of dominance in views on the causes of evolu 1936 that show publications s
Gradual Acceptance of Natural Selection In 1935we find a few indicationsthattheFisher-Haldane-Wright (FHW)
A. F. Shull, in his textbook Evolution, declared that after years of neglect, "the theory of natural selection is coming back," but he acknowledged that
Allen (1979) provides a tabulation of views on evolutionary mechanisms the early twentieth century. 66. Barnes (1931); see more extended comments in his book (1933). that Fisher of biologists active in
father, Joan Fisher Box writes Eugenics daughter"
2007,131). See alsoBowler (1998,2004).

67. Goldschmidt nest B. Babcock
In her biography of her gave a copy of his book to Barnes, "an old friend in the notes that Barnes was Society" (Box 1978, 193) and later "godfather of his youngest to Barnes was Fisher's "old tutor" (Moore (Box 1978, 279). According James Moore,
and Robson and Richards (1934,171) (1936). The prominent botanist Er is to argued that the Fisher-Haldane-Wright theory inadequate explain the evolu tion of new species in the genus be because that theory is based on gene mutation. Babcock Crepis lieved that one must postulate transformation of entire chromosomes, such as the reduction from ten to as well as and polyploidy 1934). (Babcock eight chromosomes, interspecific hybridization 68. Muller textbooks (1927) and Crow this work, and Abrahamson forwhich Muller (1997). Muller also published Prize a readable com
prehensivesurveyin the popularmagazine Scientific Monthly (1929). One of thefirstbiology

to mention received the Nobel
andRobbins (1934,577).
69. Kingsland (1995)
in 1946, was Holman
tions the validity of this dichotomy.
ists,see the articles byGarland Allen (1979) andKohler (2002); butHagen (1982,1999) ques 70. E. R. Dunn (1935); Muller (1935); and Plough and Ives (1935).
and Mayr
(1980c).
On
the split between
experimentalists
and natural
31
CHOOSING
SELECTION
traditional
and more
seems thatwe Robson
still have some validity. It almost objections are forced to believe in natural selection because all the other recent are even less credible.71 Animals of for natural about
known alternatives
theories of Fisher, to future work, were Haldane, based on unrealistic assumptions and "[did] not in fact provide any proof of the efficacy of selection," other than the alleged absence of any other and Wright, while valuable as a stimulus
in Nature, selection was weak, but they were much less optimistic the prospects for rescuing the theory.The mathematical
in their and Richards, The Variation monograph, also asserted that most of the traditional evidence
than Shull
"for the spread of variants that occur as single or few indi explanation viduals." But, taken as a whole, "the direct evidence for the occurrence of [njatural [sjelection is very meagre and carries little conviction."72 Accord andWilliam Provine, this book was very influen ing to Stephen JayGould tial and represented the view, widely held at that time, that the differences between
are on the other hand, said species nonadaptive.73 Ernst Mayr, were those who held that view "verymuch in the minority" (1980a, 132). "Present State of the Theory of Natural Watson: ing statement by D. M. S. The Selection" began with the follow
A discussion at the Royal Society of London (also in 1936) on the
a theoretical explanation of any hypotheses it becomes capable of giving conceivable occurrence, and that the scope and indeed the validity of its basal assumption of a selective death-rate determined by a favorable varia tion have not yet been established. (Watson et al. 1936,45) Despite natural vigorous protest (Fisher 1936), Watsons selection set the tone for the discussion. Gates selection, favoring instead mutations Fishers
theory of natural selection is the only explanation of the production of adaptations which is consonant with modern work on heredity, but it suffers from the drawback that by the introduction of related subsidiary
skepticism about at and MacBride re
tacked natural
and Lamarckism,
McAtee's
is probably worthless because of asserted that evidence from protective coloration the contents of stomachs of birds) and other criticisms; the experiment (examining cant yet theory explain how complex organs like the eye arise by small steps, since the early steps no on pure lines still count give negative experiments against the NSH; advantage. Johannsens since "the observed differences between varieties of a species appear not to be adaptive," we must 71. He rely primarily 72. Robson on chance rather than selection to explain their origin" (Shull 1936, 142, 161, 166-83,205,207,212). and Richards
73.Gould (1983, 89) and Provine (1986,295).
(1936,218,314,310).
32
Fisher-Haldane- Wright Theory Early Reception of the
and Hal (Timofeeff-Ressovsky, Carpenter, spectively. Other participants out in favor of NSH. evidence and the dane) pointed arguments Finally, a to for the the British Association Julian Huxley gave pro-selection speech Advancement of Science called "Natural Selection and Evolutionary Prog ress" (Huxley 1936), the first draft of the manifesto 1942 book, Evolution: The Modern Synthesis. that later became his
In 1935, most publications that had discussed natural selection expressed an unfavorable view of it.74 In 1936 and 1937, the pros and cons were about of Fisher, Haldane, counting the publications equally balanced, without in 1938 and Dobzhansky.75 and continuing through the 1940s, Beginning were a majority of the articles and books that discussed natural selection
favorable.76
but argued that it could not explain macroevolution?in particular, the formation of new species. For this, he invoked his own theory of macro mutations, using the notorious phrase "hopeful monsters" (Goldschmidt idea may have been attractive to a few botanists, since form new species in one generation by polyploidy plants occasionally as noted chromosome Weatherwax sets), (1947, 391-2) by (combining 1940, 390). This
was the 1940s, the most prominent of the NSH opponent During ac Goldschmidt the respected German Richard Goldschmidt. biologist as an for microevolution cepted natural selection adequate explanation
and laterby Mayr

league at Berkeley
(1980a, 130). But E. B. Babcock, Goldschmidt s col

firmly rejected
minority process" in plants. s Goldschmidt synthe theory became the punching bag for the modern sis (Gould 2002,68). As he realized, "all theories of evolution tend to reflect
74. Books McAtee, and Sumner. 75. Pro-Natural Selection: 1936 book by Shull; articles by Carpenter, East, Gordon, 1937 book by Sturtevant and Beadle; articles by Ford, Just, Kemp. 1936 books by.Mavor, Mercier, and Richards; articles by Robson by Nordenskiold Robb, Wheeler. and by andWood; Whitefield articles by Balfour, Blum, B?ker, were those Articles expressing favorable views by E. R. Dunn
view (Smocovitis 2008), and the saltationist Mayr called polyploidy "the
and a leader in research on polyploidy,
Nilsson,
Turrill; Timofeff-Ressovsky, ytoz-Natural Selection: MacBride, Watson; 76.1 give the detailed
Scott. byMcKerrow, list for only one more year. Pro-Natural Selection: 1938 books by Allee, et al.; 1938 articles and Mar Elton, Gates, Har?an Dendy, Ford, Shull, Young by Carr-Saunders, tini, Muller, Quayle, Shull, Sturtevant. 1938 book by Bradley. The ^//-Natural Selection: textbook by Young, Stebbins, and Hy so and is to consider the lander, counted as "pro" in the above list, is not enthusiastically willing possibility I am of orthogenesis. See tables 2 and 3 for publications after 1940. and books I have been able to look at are representative assuming that the periodicals at least of those not in the United States and Britain and that a larger sample would published text. in the the trend described change qualitative
1937 books
33
CHOOSING
SELECTION
for him, his own theory was a reflection of the holistic, neoromantic of the 1930s, which was about to be displaced by the mechanistic
the scientific trendsof theirtime"(Goldschmidt 1940, 397); unfortunately

biology reduc
TheModern there was
tionistic biology of the 1940s and 1950s.77 Beginning in 1942 with the publication of JulianHuxley sEvolution:
s and the Synthesis and Ernst Mayr Systematics Origin of Species, a stream of books and articles by leading biologists, articulating
and strongly supporting the evolutionary synthesis. It is now time to ask, what were the reasons for this groundswell of support, and to what extent was it focused on a as the natural selection single coherent theory such
hypothesis?
77. On Barash Stebbins based
Goldschmidt's
(2003). The
(1969) on the Watson-Crick
and reputation, see Dietrich (1995); Schmitt (2000); and is discussed by Gould of his macromutation (2002, 453ff). reception theory was killed new molecular declared that Goldschmidt's by the biology theory life,work, double biology helix model of DNA. But even before as a had rejected macromutation that, leaders in paleon significant factor in evolution:
seeBabcock et al. (1942);Huxley (1942); Sumner (1942); White (1945);Allee et al. (1949); Pat terson and Stone (1952); Watson (1952); Mather (1953); and Sheppard (1954). Mayr, inresponse
toDietrich (1995), wrote that Goldschmidt's (1940) book was felt sorry for him?a few years earlier he was "the pope of German had to make his living by giving undergraduate courses" (1997, 33). reviewed gently because everybody ... now in exile and biology
tology and evolutionary
34
The Faraday ofBiology?

was born and educated, there was much Russia, where Dobzhansky more less support for Darwin's theory of natural selection and much In in between and Amer natural than Britain and separation genetics history ica.78 When he came to the United States in 1927 to work with Morgan's and then at the California Institute of Technol group, first at Columbia enhanced his already-substantial of genet ogy, Dobzhansky knowledge ics and used it to continue his field research in entomology (Dobzhansky 1933). His established famous series of papers on the genetics of natural populations a strong link between the established about Droso knowledge
Dobzhansky:
new in the in the wild.79 about Drosophila laboratory and knowledge to Francisco most "The According important empirical fact Ayala, single is the ubiquity of genetic variation" established and argued by Dobzhansky phila provided strong indirect sup that speciation may proceed in small steps. by showing a He became interested in Donald Lancefield's discovery that newly dis covered strain of D. later called sterile Drosophila, pseudoohscura,produced some males when crossed with other strains, but also produced fertile fe Dobzhansky's port for the NSH males. This
(Ayala 1976, 5).
research
in the mid-1930s
was
firststep in theoriginof a new species (Lancefield 1929,288). Dobzhansky

a new species inspired by the possibility of creating a feat as He regarded previously impossible.80 proceeded
78. On
"may represent
an earlier step in the evolutionary
process"
as the
in the laboratory, by studying the
see Vucinich the nineteenth-century (1974); reception of evolutionary ideas in Russia, s own Russian see in and his work Adams (1994) and lab, Dobzhansky background Morgans account of his life and work. Allen (1994). Ayala (1990) gives a comprehensive on
79.Dobzhansky (1981); Levine (1995); andAyala andFitch (1997).

80. Later he became made
of it (Dobzhansky 1941b,233-5,273, 325).
cies, Raphanobrassica,
aware of the creation in the 1920s of a new spe by G. D. Karpechencko broccoli with and he radish, gave an extensive discussion by crossing
35
CHOOSING
SELECTION
genetic basis of hybrid sterility (Dobzhansky 1936). He argued that two

were in fact different were species because they pseudoobscura one of them was later renamed D. reproductively isolated; persimilis. His work suggested that speciation was not necessarily a sudden discon strains of D.
s mining Richard Goldschmidt theory. Instead, he postulated that there is no as a difference and macroevolution between microevolution qualitative pragmatic methodological assumption81 and then gathered empirical evi dence for this postulate; thus, research on small changes at the level of genes is directly relevant to the general problem of the origin and chromosomes is not a discrete event but a process that occurs over of species. Speciation
tinuous process involving a macromutation, thereby challenging the views of those evolutionists who still followed de Vries and preemptively under
were
a several generations; species might be considered what mathematicians in his review of Goldschmidt's later to call a "fuzzy set."82 Wright, that we have examples of every conceiv book, agreed with Dobzhansky able intermediate step; therefore, there is no justification for assuming the "'bridgeless gap between species which is crucial" forGoldschmidt's theory
of mathematical but his own
(Wright 1941,166). reliedon Sewall Dobzhansky initially Wright to tellhim the conclusions
population genetics to and was influenced to favor Wright's
views on the importanceof genetic drift (Epling andDobzhansky 1942),

results led him Dobzhansky selectionism s career inAmerica in the 1940s in the 1940s.83 Thus, change those views illustrates in microcosm the shift toward
and 1950s.
1937 may be taken as the first announcement of the evolutionary synthesis. a review of the biological evidence Dobzhansky presented comprehensive on mutations, mechanisms for variation, selection, speciation (including and of and He called at isolation), evolution, patterns species. polyploidy tention to he first evidence Muller's which called the conclusive x-ray work, for an external cause of mutations. second edition (Dobzhansky He 1941b),
The publication ofDobzhansky's Geneticsand theOrigin ofSpecies in
as a tool of the application of these chromosomes genetic research by T. S. Painter (1934)," which "enormously facilitated the comparison of the gene 81.Dobzhansky (1937,12); Powell (1995, 74); andOrr (1996).
and Hill (1997). For debates about the definition
somes in the larval salivary gland of flies byHeitz and Bauer (1933) and
in the and emphasized mentioned, the "rediscovery of the giant chromo
82.Willermet
(1935); Emerson (1945); Krementsov (1994);Mayr (1948,1949a);Winsor (2000); andReydon (2005). paper in Dobzhansky (1981, 305-28); Provine (1986, 389-90); andLewontin et al. (2001).
83. Dobzhansky (1943,1971); see the editorial note on page 303 preceding the reprint of this
of species,
see
Dobzhansky
36
Dobzhansky: The Faraday of Biology?
arrangements in different strains."84 This tool provided the basis for Dob s research in the 1940s on natural ofDrosophila. zhansky populations went Genetics and the through three editions under that Origin ofSpecies under
are gener the title Genetics of the Process (1970). These Evolutionary to have been very influential in to considered ally biologists persuading reason for this influence, accord adopt the evolutionary synthesis.85 One to is Leah Ceccarelli the rhetorical (2001), ing approach its author used. than claim that one field of science should be reduced to another
was published revision title(Dobzhansky 1937,1941b, 1951), and a further
Rather book
more fundamentalfield(as did E. O.Wilson
to cooperate as not done in the past (at least in the they had scientific community). Before 1937, naturalists thought Anglo-American in de Vries s saltation theory,while geneticists did still believed geneticists would have not appreciate the great amount of variation in natural populations; Genetics overcome to and the both their misconcep groups Origin of Species helped tions and recognize how they could profit from each other's work (Cec
it clear that knowledge discussed by Ceccarelli), Dobzhansky made and research methods from several fields must be integrated in order to understand how evolution works. In particular, geneticists and naturalists
inSociobiology [1975], another
genes and chromosomes, populations, and species. "Since genetics can pro account the ceed without level, it retains more taking into populational relative independence than do the other fields in the synthesis"; popula
carelli 2001,28). At the same time, as Lindley Darden has pointed out, the relations between fields in the evolutionary synthesis were not symmetrical, because the synthetic theory had to deal with three levels in a hierarchy:
tion genetics depends on "new findings ofmutational processes in genetics," and population processes "can continue whether or not isolation [producing new species] has occurred."86
toM. J.D. White, 84. Dobzhansky in other (1941b, 117). According giant chromosomes were discovered as as 1880. flies in have been much faster early dipterous genetics might Progress if researchers had been familiar with "the work of Balbiani, Carnoy, Alverdes and oth Drosophiia
ers on the of Chironomus" and had looked at salivary glands twenty years salivary chromosomes earlier (White 1945, vii, 3). While the "rediscovery" came too late to play an important role in to accept it did chromosome persuading biologists Morgan's theory of heredity (Brush 2002), come to just in time change Dobzhansky's and isolation. As Muller who confirmed noted views on the relative importance Dobzhansky of natural was selection
and Dunn were
85.Gregory (1946); Gould (1982); Provine (1994); Krimbas (1995); Ceccarelli (2001, 26);
(1961 reminiscences 86. Darden (1986,121,122). assumed in 1937 tended level] may influence [1994,100-1]). quoted by Provine She also notes that the asymmetrical of gene mutation hierarchical "Selection (Darden relations that
in his own survey of research on mutations, the effects of x-rays (Muller 1929,495).
one of those
to break down the evolution
as a result of later research: mechanisms"
[on the pop 1986, 122).
ulation
37
CHOOSING
SELECTION
In thefirstchapterof the thirdedition ofGeneticsand the Origin ofSpe

cies,Dobzhansky (1) "unraveling distinguished and describing two to evolutionary problems: approaches the actual course which the evolutionary
which happened in thepast" (Dobzhansky 1951,11).This is an elaboration

of a brief statement in the 1941 edition (6), where he attributes the identi fication of these two approaches as "generalizing" and "exact induction" to Hartmann (1933). In the spirit of the evolutionary synthesis, the two are rather than antagonistic.87 approaches complementary
that bring about evolution, causal rather than historical that can be studied experimentally rather than events problems, phenomena
and (2) "studies process took in thehistoryof the earth"(this isphytogeny);

on the mechanisms
M.
For other
recent discussion two
of the scientific
issues raised by Dobzhansky's
book,
see Lewontin
(1997).
87. The Sarkar to approaches correspond the work of Haldane distinguish to the and retrospective prospective from that of Fisher and Wright; categories used by see note 57. Dob
use of thedistinctionisdiscussedby zhansky's Ayala (1977) andProut (1995).
38
Evidence
forNatural Selection before 1941
For
our purposes, the most relevant part of Genetics and the Origin of is the survey of evidence for natural selection, much of which Species to support the more could provide reasons for biologists specific natural on mutations. I selection hypothesis based small have used the more ex tensive list found inDobzhansky the evidence into six categories. s second edition,88 inwhich he organized
Laboratory Experiments
In the Origin discussed the "remarkable fact," of Species, Charles Darwin a few years earlier, that many of discovered by Thomas Vernon Wollaston are the beetles inhabiting theMadeira Islands in theAtlantic Ocean wing less, unlike related beetles elsewhere. Darwin suggested that
thewingless condition ... ismainly due to the action of natural selection, but combined probably with disuse. For during thousands of successive generations each individual beetle which flew least, either from itswings so little less or from indolent habit, having [been] ever perfectly developed will have had the best chance of surviving from not being blown out to sea; and, on the other hand, those beetles which most readily took to flightwill oftenest have been blown to sea and thus have been destroyed.89 suggestion, elaborated in Darwin's to a
This
1975, 291), is equivalent an environ population of beetles with varying wing sizes and put them in are most ment where those that have the to be blown largest wings likely (Darwin
on natural selection "big book" start with a prediction that if you
[and later editions], chapter 5 on "effects of the increased use and disuse and information relating to this suggestion, see Darwin (1989, correspondence I thank Sandra Herbert for this reference. 267-74,279-80,283-5). 89. Darwin (1859 of parts"). For
88.Dobzhansky (1941b, 188-214).
39
CHOOSING
SELECTION
away by strong winds, then (assuming wing size is an inherited character) aftermany generations, the population will consist of a much larger propor tion of beetles with small or absent wings. Madeira In the following decades, Darwin's flies of discussion of the wingless was often as an an observation be of how could given example were blown away; explained by natural selection: the insects with wings the others whether was remained selection and Geddes (Thomson 1931). But itwas not clear on or acted small mutations, or even whether wind large
"wind" is varied while other factors are kept con mechanically produced our over the course of many stant; generations, the hypothesis predicts that is increase of should if the wind flies strong but not if proportion wingless it isweak A or absent. at this proposal, biology might scoff a because "everyone knows" that evolution is such slow process that it cant less in a laboratory be directly observed within a human lifetime, much a few weeks or months. But the critic has experiment lasting only ignored critic unfamiliar with modern
to the sci important selective factor. According (Popperian) we an in entific method, should do which the intensity of a experiment the most
a "model the fact that geneticists in the early twentieth century developed so which reproduces organism," theDrosophila fly, rapidly that such experi ments can be done in the in time. real By 1930, any good geneti laboratory even cist could do the classical experiments research and assign Drosophila as exercises for students.90 practical
a favorable environment for In 1937, in France?not usually considered test was done. research on natural selection at that time91?the Philippe L'H?ritier, Yvette Neefs, and Georges Teissier carried out an explicit test of
Darwins
byL'H?ritier andTeissier (1934),which allowed one to studyan isolated

population Nous under controlled conditions.92
prediction. They
used
"population
cage," previously
developed
nous sommes propose detudier exp?rimentalement leprobleme pos? par nous avons avons et Darwin utilis?pour celaDrosophila melanogaster. Nous
a r?alis? les conditions que Darwin suppose Vorigine de I evolution des Insects en normaux ail?s et une constituant insulaires population mixte d'individus
90.Kohler (1994) and Brush (2002,519).

91. Lamarckism more moral dominated French
thistoCatholicswho thinkit is (1983,1989) andGrimoult (2000). L'H?ritier (1981) attributes

than Darwinism. year 1931-1932 in the United States at Iowa State University), where
biology
until
after World
War
II, according
to Buican
spent the academic
92. L'H?ritier (1981); Grimoult (2000, 151-63); andGayon andVeuille (2001). L'Heritier
(mostly
40
Evidence for Natural Selection before1941 une se d'individus portant le car?ctere r?cessif vestigial (vg) qui manifest? par au vol. (L'H?ritier et al. 1937, tout ? des ailes devenues fait impropres atrophie
907-8)
[We decided to study experimentally the problem posed by Darwin and We have realized the have, for this purpose, used Drosophila melanogaster. conditions thatDarwin assumed to exist originally: a mixed population of
some insectswith normal wings and others having the recessive character an so that they atrophy of thewings, vestigial (vg),which ismanifested by have become inadequate for flight.] And were able indeed, in a trial of less than two months, they s firm the results predicted from Darwin ("exactement hypothesis to con celle que himself
L'H?ritier Although permettait de pr?voir Fhypo these de Darwin1). later characterized this as an arnusante" (1981, 338), and Hal "experience dane called it a "rather trivial but still illuminating experiment" (1939,127), identified it as a confirmed prediction and wrote that C. H. Waddington
it is "an example of a type of work which is of considerable importance for the experimental study of evolution but which has been surprisingly little
who gave this as his first example of natural selection in Dobzhansky, the laboratory, did not mention Darwin's role, nor did he use the language of hypothetico-deductivism and prediction-testing.93 Instead, he quickly moved on to the next category.
takenup" (1939,302).
Historical Changes in the Populations Compositionof

that in the case of major changes, such as the evo conceded Dobzhansky lution of the horse or the anthropoid ape, the generic critic of evolution
andWright. He thought to beach. strolling According Gayon ment "provided its authors with lasting fame in the polemical tion. .. .This experiment was an effective reply to adversaries the work on an American he learned about of Fisher of the idea of the population cage and Veuille, the Darwinian experi context of French studies on evolu of Darwinism. It as much seemed
while
an artist's as a scientific to ridicule the its purpose investigation, performance probably being numerous French of 'transformism.' Denying natural selection was like flying in the opponents
was
was rather standards; while the wind strength experimental technique sloppy by modern to was no not there similar control "controlled," group (a exposed apparently population I thank David A. O'Brochta communication for a critique of the experiment wind). (personal The with author, March 4,2005). omits this ex 93. Dobzhansky (1941b, 189). In the 1970 revision of Genetics, Dobzhansky not found any statements in the literature up to that time other thanWad I have ample entirely. (quoted in the text), claiming that the experiment was a confirmation of a prediction.
faceof the wind" (Gayon andVeuille 2001, 89).
dingtons
41
CHOOSING
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(whom I invoked above) would be correct: evolution was supposed to be so slow that no changes could be noticed within a human lifetime inwild spe cies. So, Darwin and his followers had to use the results of human selection and breeding for the theory of natural selection: "The relatively rapid changes brought about in domestic animals and cultivated by artificial selection were considered a model of the evolution in the wild state, rather as evidence
the beginning of the twentieth century, fumigation attempts with to eradicate red scale insects that attack citrus trees inCali hydrocyanic gas fornia were
than evolutionas such" (Dobzhansky 1941b, 190).

At
that there were
no longer worked. ExperimentsbyH. J. fumigation Quayle (1938) showed

two races of red scale?one of which is resistant to the
a few years initially successful, but after
itwas
observed
that
gas, the other not. By eliminating the nonresistant insects, the fumigation called allowed the resistant race to take over the population. Dobzhansky
obtained" (1937,161).
But where zhansky ing "introduced academic"?a
this "probably
the best proof of the effectiveness
of natural
selection
yet
insects come from? Dob did the hydrocyanic gas-resistant suggested that they resulted from mutations, rather than from be
from elsewhere," but he considered this question "largely rather inappropriate pejorative for a professor to use! In a later revision of the book, he seemed unsure whether the mutations "are
before the insecticide is applied, or arise after present in pest populations the application. This question is usually insoluble, and it is of no particular 1970,215). The context for these state importance anyway" (Dobzhansky ments is the shift in the views of many evolutionists toward the position s own is so much hid (supported by Dobzhansky experiments) that there den genetic variability in any present-day population that new mutations are not to drive evolution (Stebbins 1970,182). really needed After mentioning
areas near industrial seemed
other examples of the evolution of agricultural pests dis (citricola scale, codling moth, and stem rust of wheat), Dobzhansky cussed "industrial melanism": the appearance of darker forms of moths in cities in England countries. He and other European to accept E. B. Ford's that explanation (1937,1940b)
melanics
... their are superior to the light-colored types in vigor, and are not protec spread in populations is normally prevented because they areas this tively colored. In industrial disability is removed by the general
darkening of the landscape.94

94. Dobzhansky (1941b, 196); similar statement inDobzhansky (1951,132-3).
42
Evidence for Natural Selection before1941
Experimental Study of Adaptation inPlants

noted the earlier work on plant races adapted to a specific Dobzhansky ... who must habitat by "the Swedish botanist and geneticist Turesson 1941b,
197). But he gave much more space to the recent research of Jens in California M. Hiesey (Clausen Clausen, David D. Keck, and William et al. 1940). In order to show that the differences in of plants of phenotype the same species growing in different habitats to natural selection) reditary (and therefore due (called "ecotypes") are he rather than environmental
basis" (Dobzhansky be credited with having placed it on a trulyscientific
used (directly respondingto thathabitat), they

of reciprocal transplants [:] ... Representatives of the popula tion of the habitat A are transplanted in the habitat B, and those from B are planted inA. Comparison of the native and transplanted strains themethod permits the observer to discriminate between the changes induced directly to the strains themselves. (Dob by the environment and those intrinsic
1941b,197-8) zhansky
there are three ecotypes of the cinquefoil (five-petaled rose, in the Coast Ranges (1) the "coastal ecotype,"growing Potentillaglandulosa): of California, a region which "has a rainless summer and an equable climate For example,
the coastal ecotype does not survive very Although to the much harsher conditions of the foothills, in transplanted the ecotypes tend to carry over theirmost distinctive characteristics general, when transplanted to a different habitat, with minor modifications.95 other in the meadows. long when
to grow almost throughout the year"; and (2) ecotypes permitting the plants in the dry slopes, the in the foothills of the Sierra Nevada Mountains?one
given by Dobzhansky rough summary of the rather confusing discussion et al. are described in more detail in and subsequent work of Clausen see also the discussion Smocovitis (1988,214-33). (1970,292-5); Dobzhansky by Dobzhansky of several strains of the discussed the experiments of W. Sukatschew (1928) on the competition (1941b, 197-9). This near {Taraxacum Leningrad qfficinale) grown in the in the north, and Askania-Nova Vologda strains better the and did densities, Vologda Archangel not when at lower densities. (The one from the planted dandelion angel, conclusion case.) The on additional factors was such as the presence that survival depends of competing and brought south). When southern from other places planted together (Arch at high strains, but did
95. This
is my
than the locally grown not survive in either region did not just on environmental conditions but also plants. This example was omitted from
thefourth edition (Dobzhansky 1970).
43
CHOOSING
SELECTION
Animals Adaptation in Experimental Studies of

Dobzhansky (1941b, 203-5) discussed the experiments ofN. W.Timofeeff on the survival values of strains of Ressovsky in the 1930s Drosophila funebris areas in at three different tempera from twenty-four geographical Europe tures. Each was forced to compete against a standard strain of Drosophila ofD.
with 150 eggs ofD. funebris by putting150 eggs of one strain melanogaster
in a culture bottle and counting the number that hatched. melanogaster the (unstated) hypothesis was that since "D. melanogaster is na Presumably to tive the Tropics while D. funebris occurs in the Temperate Zone," strains ofD. funebris from colder regions (Russia, Sweden) would survive relatively
better at the lowest temperature (15?C) than at thehighest (29?C) (Dob
in this reported the results of several other studies on the gypsy moth, and on the pocket mouse, Goldschmidt chapter?Dice seem on to consist mainly of qualitative these studies Fox crustaceans?but rather than effects of environmental variations with ad hoc explanations, Dobzhansky also systematic experiments. Most purposes in the 1970 edition. of this material was gone or used for other
seem to confirm this hypothesis, zhansky, 1941b, 203-204). The numbers a statistical no to of their made attempt give analysis although Dobzhansky not use the of he did and again language prediction-testing. significance,
Regularities
in
Geographical
Variation
the emphasis is on the similarities of different species in the same Here, area and on the similar variations found as one goes from one area to an is a counterpart on the supraspecies level of the other. "This phenomenon is far outdistanced by botany ecotypic differentiation with species. Zoology in the formulation of regularities of this kind." For example, Gloger s rule literature as Allen's (frequently referred to in theAmerican races rule) states that inmammals and birds, inhabiting warm and humid races of the same species more melanin regions have pigmentation than in cooler and drier regions; arid desert regions are characterized by ac of yellow and reddish-brown phaeomelanin pigmentation.
cumulation
207) (Dobzhansky1941b,
What do these rules have to do with natural selection?
as itmay seem, the correlations between race formation and en Strange vironment revealed by the "rules"... were repeatedly quoted as arguments
44
Evidence forNatural Selection before1941
against the natural selection theory.This amazing confusion of thought was due in the past to the almost universal acceptance among biologists of the belief in inheritance of acquired characteristics.... To suppose that geographical races were originally modifications that have subsequently been fixed by heredity is contrary to thewhole sum of our knowledge_ Therefore, the regularities observed in the process of geographical race formation cannot be due to direct effects of the secular environment, al beit the results of this process can in part be imitated by some phenotypic modifications. (Dobzhansky 1941b, 208,210)
was as sup In other words, all of this empirical evidence previously regarded is false, ithas to be port for Lamarckism, but sincewe now know Lamarckism
as support for the only plausible alternative that accounts for regarded instead correlations between adaptations and environment: natural selection.
Protective and
As
Warning Coloration
as shown was still not spectable, by the treatise of H. B. Cott (1940). There much careful experimental work; mentioned Dobzhansky only the research of Francis Sumner (1935) showing that "fishes whose color contrasts with more their surroundings are caught by predators easily than those with s for colors." As McAtee "refutation" (showing that the stom harmonizing achs of birds contain supposedly poisonous insects in the same proportion as their relative abundance
to the whole bringing only disrepute theory.... Some of the alleged protections and warnings have been shown to be armchair protection and museum mimicry." But despite heavy criticism in the 1930s and McAtee, the theory was revived and became quite re by Heikertinger ueless speculation
for protective and warning coloration, including mimicry?evidence for natural selection going back to the nineteenth century (Kimler 1983a, that much of it is "uncritical and val acknowledged 1983b)?Dobzhansky ...
in their environment), Dobzhansky argued that one should not assume that an absolute from "only immunity predators can make natural selection effective." All that was needed, to according the modern theories of mathematical of one part in one familiar with a population genetics, is selective ad cause a to thousand significant change in the
vantage
compositionof the species (Dobzhansky 1941b,212-4).

Readers thatDobzhansky
the history of evolutionary biology will recognize s survey of the evidence was very incomplete, but my goal is not to evaluate or as a historian; rather, it is to supplement Dobzhansky suggest how the presentation of the evidence in his very influential treatise set the agenda for others.
may have
45
Huxley: A New Synthesis

Is Proclaimed
Julian
Huxley, and "the most
Oxford One
mentor to E. B. Ford, was an inspirational at in force the selectionist attitude powerful developing in the 1920s."96 Huxley himself recalled: according
of the greatest changes in biological outlook during my active life has been the re-erection of natural selection as themain and perhaps the sole agency of significant evolutionary change and adjustment.... Perhaps owing to a certain familiarity with natural history, I clung firmly to a belief in the principle of natural selection and its efficacy during the period up to
the early or middle 1920s, when itwas being neglected or attacked by as advanced, and when most of the biologists who thought of themselves even eminent geneticists could seriously assert thatwithout natural selec
tion all existing types could have come into being, and a vast number of others as well!97 some time in the early 1920s, Huxley series illustrating mimicry in butter
When
T. H. Morgan visited Oxford "showed him some of the wonderful Poulton enthusiasm; had amassed.. and
flies which his vivid

existed.'"98
is extraordinary!' he said, with then, T just didn't know that things like this
. . 'This
of specialists such as Darlington, In his introduction he wrote: the work

(1980, 337). "J. S. Huxley view of evolution in England" synthetic On his worldview, (1982,1984). Hagen 96. Ford ...
(1940), helped to publicize Huxley's edited book, TheNew Systematics

Ford, and Timofeeff-Ressovsky.
played
a central
97.Huxley (1951,593; emphasisadded); cf. Huxley (1926). 98.Huxley (1951,593); alsoDronamraju (1993,237).
(Provine 1980, see Greene (1990).
331). On
role in creating a neo-Darwinian his relation to systematics, see
46
Huxley: A New SynthesisIs Proclaimed reluctance, especially among some of the younger experimental biologists, to recognize the prevalence of adaptation and the power of selection. This is doubtless in large part a natural reaction against the facile arm-chair reasoning of a certain school of evolutionists. is still a widespread
There
While
endorsing the importance of natural selection, he also argued that "isolation is the essential factor in bringing about taxonomic (geographical) and stated that "where isolation is relatively or quite complete
divergence"
will produce a certain degree of random, nonadaptive change." But "isola new combinations tion of genes al fixing of (apart from the accidental to effect differentiation without mutation, and, is ready present) powerless inmost cases, without selection" (Huxley 1940,12). Huxley introduced the term "synthesis" and popularized praised the work of Fisher, who his own version
and the isolatedpopulation small, the Sewall Wright effect(1931, 1939)
The Modern
Evolution: of it with the titleand readable style ofhis influential 1942 book,
Synthesis. He
has radically transformed our outlook on the subject, notably by pointing out how the effect of a mutation can be altered by new combinations and mutations of other genes. Any originality which this book may possess lies to generalize this idea still further, stressing the partly in its attempting by fact that a study of the effects of genes during development is as essential for an understanding of evolution as are the study ofmutation and that of
selection."
He
asserted we
that thanks to the mathematical can now make "quantitative
Wright,
in his book, whose pages are remarkably free of find any such prophecies numerical data.100 Huxley's book is nevertheless a comprehensive qualita tive survey of the arguments and evidence for the selectionist view. As such,
ness than was possible toDarwin" (Huxley 1942, 21). But it is difficult to
and of Fisher, Haldane, prophecies with much greater full work
argues that it is a constriction that eliminated most believed to affect evolution.
theory(Waters and vanHelden 1993, 24-25). Will Provine (1988,1992)

of the factors previously
it is not a synthesis of different theories but of observations in support of one
99.Huxley (1942, 8).

100. Ruse Hardy), Oxford.
that Huxley, (1996a, 329) writes "did not understand enough mathematics, See also R. W. Clark (1968) andWaters
own admission (in a 1921 letter to A. by his to at physics, and chemistry" study physiology and van Helden (1993).
47
CHOOSING
SELECTION
a by "the convergence of number of separate lines of evidence, each partial and indirect or incomplete, but severally cumulative and demonstrative.... These various lines of evidence all converge to support a neo-[M]endelian
induction of melanism (as claimed Huxley noted that the (Lamarckian) W. H. Harrison) had not yet been disproved, so the validity of the by J. natural selection hypothesis, in this and other cases, must be established
view, some showing that small mutations occur, others that selection is ac are tive, that some mutations potentially beneficial, that through selection can be to the needs of the or of the gene-complex, mutations adjusted are ganism, and that adaptations genetically determined and vary in type and accuracy with 122-3). But what direction and intensity of selection" (Huxley 1942, 116,
kind of evidence
to accept it as the dominant process in evolution? answers this work of question. First, the mathematical Huxley explicitly Fisher and Haldane, combined with modern genetics, definitively refutes are much more the three major alternatives to the NSH. Small mutations to likely produce adaptive improvements than large ones, overwhelmingly so when "the harmonious (as now seems probable) adjustment of many the de Vries large independently varying characters is required"?hence
for a biologist who was selection but reluctant
did Huxley himself consider most persuasive already familiar with the older evidence for natural
mutation
operates (Huxley 1942,123). But
ef theory is untenable. Similarly, orthogenesis and Lamarckian even as as cannot in ifpresent, be important selection evolution fects, long
E. B. Ford, the effect of a mutation was assumed to be constant. A given mutation, we may say,made an offer to the germplasm of the species, which had to be accepted or declined as it stood.... To-day we are able
perhaps the most important single concept of recent years is that of the Be adjustment of mutations through changes in the gene-complex. fore this had been developed by R. A. Fisher and his followers, notably
... the offermade a to look at the matter in a by wholly differentway mutation may be merely a preliminary proposal, subject to negotiation. Biologically, this negotiation is effected in the first instance by recombi nation and secondarily by mutation in the residual gene-complex. (Hux
ley1942,124)
also recognized Huxley small populations" (Huxley of drift in "the Sewall Wright phenomenon 1942, 59). This effect is involved in speciation,
andHuxley called it
48
Huxley: A New one of themost
SynthesisIs Proclaimed
important results ofmathematical analysis applied to the facts of neo-mendelism. It gives accident as well as adaptation a place in evolution, and at one stroke explains many facts which puzzled earlier selectionists, notably themuch greater degree of divergence shown by is land than mainland forms, by forms in isolated lakes than in continuous
river-systems.101
became Huxley increasingly skeptical about on G. evolutionary significance of genetic drift. In 1945, commenting s use of to in fossil the record, Simpsons Wright theory explain gaps could persist for wrote, "Geneticists may doubt if such populations millions of years needed for non-adaptive evolution" (Huxley 1945a,
In later publications,
the G. he the 4).
But he was still willing to give drift a role in speciation (Huxley 1946, by driftin small populations ismuch less frequent than Wright thought;
5). In 1951 he asserted that the establishment of nonadaptive characters
and others had shown that "truly non-adaptive Ford, Muller, genes and was not are gene combinations exceedingly rare" (Huxley 1951, 597). He s rather that that made incorrect but asserting Wright theory predictions, the theory was not relevant because adaptation is operating. (Wright
would say thathis theory does apply in this case ifthepopulation is small
enough.)
In 1952Huxley published an article in the American pictorial magazine

a sensational bears what example of natural selection: the crab Dorippe looks like the face of a Samurai warrior. Ac
Life, featuring jap?nica, which cording to a
mass suicideby throwingthemselves Genji in 1155AD, they"committed

into the sea." Later, fishermen noticed crabs that bore a slight resemblance to the face of awarrior and threw them back, since not want to eat they did the supposed reincarnation looked more like a warrior's of a Heike warrior. As face were more
Japanese
legend, after the Heike
warriors were defeated by the
a result, the crabs that to survive and reproduce, likely ar so the resemblance became in later generations. Huxley nearly perfect one to curi is "more that such this be common; mimicry just happens gued ous and than others."102 surprising I find it also curious and surprising that Huxley's example of the Heike crab was not used in any of the books I examined from the period 1953
s inclusion of drift. to objected Huxley 102. Huxley The Life article (1957,141).
101. [quotation Huxley (1942,155,186,194,197,208,229,232,242,259-60,265,326,363 on 200]). SeeMuller (1929) fora good explanation Ford of this point. Olby (1992,70) notes that
is reprinted therein.
49
CHOOSING
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ora of one "just so" stories invented by earlier Darwinians, they thought this was was was to it be true.103Or perhaps it excluded because just too good considered an example of artificial rather than natural selection.
1970. Perhaps afterskeptical biologistshad become fedup with thepleth
103. But Carl tion in his popular Huxley used
as a illustrations of fish whose heads frontispiece not but did make an explicit connection authors, Waddington
Haldane and 164-5, note 202). Waddington (1969, 113) pointed out that in a 1927 textbook,
looked with remarkably like those of the the Heike crab.
was not reluctant to use the Heike crab as an example of natural selec Sagan television and Smocovitis series, Cosmos; see Sagan (1996, (1980, 25-26)
50
Mayr: Systematics and the Founder Principle
Modern
biologists resisted using it.How
most
genetics is a highlymathematical subject,yet evolutionary

in the twentieth century disliked mathematics and did the subject become established? One might emergence of three mathematical biol a very unlikely event, which three founders, These century.
conjecture
that the simultaneous
were were
and Wright?was Haldane, ogists?Fisher, to occur in the happened early twentieth were able to interact with each other separated, although geographically thanks tomodern communications and transportation technology and thus allowed to start a new
able to team up with two biologists who were not expert mathemati cians but were very good at doing their empirical work and publicizing at at results: E. B. Ford and Theodosius Oxford Columbia, Dobzhansky collaborators Those attracted students who carried on respectively. bright s new their research programs and thereby sustained the specialty. (Haldane influence was as well somewhat more diffuse, in part because he worked on several as evolutionary theory, but he did inspire another theo
specialty might have died out ifnot for the fact thatFisher and Wright
specialty within
the discipline
of biology. That
other topics
scenario conveniently illustrates what Ernst Mayr called the This founder principle. It resembles a special case of random genetic drift, in which a new species is formed by a very small and very nonrepresentative subset of a larger population. is known as one of the leaders of the Mayr evolutionary synthesis. His
rist, John Maynard Smith; see Smith [1988,126-9].)
and theOrigin ofSpecies, book, Systematics published in 1942 with an in
formulated the synthesis from troductory endorsement by Dobzhansky, the viewpoint of taxonomy and ornithology.104 In 1963 Huxley called him 104. and analysis of thebook (1985,44-59). Mayr (1942).Eldredge gives a detailed summary
51
CHOOSING
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"undoubtedly
the best all-round
was
to be retrospectively interpreted.106 In the opening pages of the new "mutual un Mayr welcomed Systematics, derstanding between geneticists and systematicts," exemplified by "Rensch and Kinsey among the taxonomists,Timofeeff-Ressovsky and Dobzhansky
age 100, he was "widely considered the father of modern evolutionary biol a he had considerable distinguished biologist-turned-historian, ogy."105As on on not influence how its development only evolutionary theory but also
the world" (Dronamraju 1993, 236). At the time of his death in 2005 at
evolutionary
geneticist
we
now have
in
among the general biolo Fisher only briefly and did s not cite Haldane at all,while it he foundWright clear that making theory the most congenial.107 His rejection of "essentialism" in favor of "population to Paul (2000), associated thinking" was, according Thompson (perhaps s view for with distaste the mathematical of the world. Mayr paradoxically) con own empirical work and general knowledge of the subject had already vinced him that geographic isolation is the key factor in the speciation of birds. Now he had a theory to support that view: thanks to Dobzhansky, "and Sewall Wright said evolution should be faster in small populations, this is exactly what we find"?in the Gala West Solomon Indies, Islands,
and Diver among the geneticists, and Huxley he mentioned gists." As for the mathematicians,
pagos Islands, and Hawaii. Alfred Kinsey found this true for gall wasps, W. F. Reinig for bumblebees.108 But now Mayr put his own stamp on the idea with a further twist: The reduced variability of small populations ... [is due] sometimes to the fact that the entire population was started by a single pair or by a single fertilized female. These "founders" of the population carried with them a very small proportion of the variability of the parent population.
only
On
(1984);Greene andRuse (1994); and Provine (2004). 105.Guterman (2005, A17) andYoon (2005). 106. Mayr and Provine (1980); Mayr (1992); Sloan (1985);Greene (1994);Greene andRuse and Winsor (1994); Johnson (2005); (2005).
107. Mayr calculations of Sewall Wright and the theo (1942, chapter 1). "The mathematical retical analysis that preceded them have been of particular value" (Mayr 1942, 217). He was and lectures (Mayr 1980b). He strongly influenced by Dobzhansky's publications rejected the selectionist view on polymorphisms of Fisher and Ford (Mayr 1942, 75). Later he criticized as used, for in his papers on the "cost" of natural "beanbag genetics" example, by Haldane 1991 hindsight view of drift is quoted next. tion; see Ewens (1993). Mayr's genetics and speciation, see Provine (2004). selec
his early career and environment,
see Mayr
(1980a,
1992a,
1999); Hagen
(1982); Hapgood
on
108. of Mayr (1942,236); Kinsey (1936,1937); andReinig (1939). For a survey Mayr's views
52
Founder Principle Mayr: Systematicsand the "founder" principle sometimes explains even the uniformity of rather
This
large populations.109 As cited his own work on the reef examples of the founder principle, Mayr heron whose color is always gray on some islands but (Demigretta sacra), both gray and white on others. s support for isolation as the major cause of speciation in birds Mayr did not prevent him from endorsing the importance of natural selection in other cases, while cautioning readers to be skeptical about the adaptive some differences invoked extreme selectionists.110 of significance by took a firm stand in favor of another postulate of the natural selec Mayr tion hypothesis: there is no qualitative difference between macroevolution and microevolution. He wrote: All the properties and phenomena of macroevolution and of the origin of the higher categories can be traced back to intraspecific variation, even though the first steps of such processes are usually very minute. (Mayr 1942,298; see also 291)
was convinced s as the cause argument for isolation major Huxley byMayr s "definitive refutation of Goldschmidt s of speciation and praised Mayr heterodox views on speciation" (1943,348). was much less firm, But Mayr s early support for Wright's general theory to his recollections half a century later. In a letter toMichael according Ruse dated November 20,1991, Mayr wrote: In my 1942 book, in order to be "modern," I quote Sewall Wright copi was very much op ously.However, inmy actual thinking and working I
to the mathematical approach, he credited Wright can size the same effect. later credited population produce Mayr B. Rensch with an earlier recognition of the Founder 1980d, 26). Principle (Mayr see 110. For caution against extreme selectionism, (1942, 75, 96). As good examples Mayr of selection he mentions the Palearctic larks (Alaudida?), "the first birds in which correlation 109. Mayr (1942, 236). for the idea that fluctuations In another in nod between soil and coloration 1932); was discovered" (Sumner and Benson's (by Hartert) study of melanistic on (Mayr 1942, 86); Sumner's work rodents in lava flows (Benson 1933). argues that the color can't be due to cli
Peromyscus
the research of Dice and Blossom, Mayr Concerning so it "is obvious that selection mate because the blackest races sometimes occur near the palest, an must have role" rules, sometimes important by predators played (Mayr 1942, 87). Rensch's as evidence for Lamarckism, can be as a result because he first used them rejected interpreted of natural selection "clines" for character correlated with 89). 1942, (Mayr Huxley's gradients climatic factors are a consequence of natural selection, though in some cases there seems to be no
adaptationalsignificance (Mayr 1942, 95).
53
CHOOSING
SELECTION
all along when he wanted to be posed to him. And I fought Dobzhansky lieve in the neutrality of the human blood group genes and theDrosophila gene arrangements. (Ruse 1996,414) the 1940s, Mayr During natural selection hypothesis. rangements a prediction in for the expressed increasingly strong support on In 1945, discussing Epling's data gene ar (see chapter 12), he made what he later called of these arrangements would be found
Drosophila that the evolution
to
marized
be influenced by natural selection (Mayr 1963, 207). A 1949 paper sum
the "impressive body of evidence" for selection s effects in both even those like Goldschmidt who denied that it can zoology and botany; its adaptive role in local populations admitted explain macroevolution
a 1950 paper turning point toward selectionism in in the position abandoned and Stresemann, which Systematics by Mayr in color polymorphisms and the Origin of Species that "most conspicuous and that they are "accidents of birds and other animals were nonadaptive" variation and without selective significance" (Provine 1986, 453). Instead, Will Provine sees a and Streseman wrote, Mayr al?eles involved in a balanced "Since recent genetic evidence indicates that polymorphism have different selective values,
(Mayr 1949b, 519).
it seems probable that many subspecies and species characters that have heretofore been considered as neutral' are controlled by genes which differ in their selective values" (Mayr and Streseman 1950,299). In his a 1963
on the use of the genetic drift concept in evolutionary vigorous attack term is used in so many contradictory ways that it "has been The biology. rather discredited": was fashionable to attribute the period from about 1935 to 1955 it
was attestedby Stephen Jay Gould (2002, 535),Mayr included naturalists
book, Animal
Species
and Evolution,
whose
influence
on
During
to "drift"or to the Sewall Wright effect in puzzling evolutionary changes the same manner inwhich the preceding generation of evolutionists had as due to "mutation." (Mayr 1963,204) explained similar changes selective neutrality, but such
Drift
is often postulated in cases of supposed rare: cases are actually quite
was
Selective neutrality can be excluded almost automatically wherever poly or character clines are found in natural populations. This clue morphism
used to predict the adaptive significance (previously denied) of the distribution pattern of the gene arrangements inDrosophila pseudoobscura
54
Founder Principle Mayr: Systematicsand the
as caused by genetic drift due to errors of sampling quoted in the past has more recently been interpreted in terms of selection pressures. (Mayr
case
Virtuallyevery (Mayr1945) and of thehumanblood groups (Ford1945).
1963,207) We have at least two pos Why didMayr change his position on drift?
want
to be scientific orthodoxy; later, when challenge what appeared as in he had established his reputation, he could afford to do so. Here, other cases, we cannot prove why a particular scientist expressed a particular view?at least not without much fuller evidence. his own founder principle, which could be regarded as a special case of ge on the not netic drift, assumption of selective perhaps because it did depend neutrality.
to his sible answers. According remarks, itwas because of the published to his letter to Ruse, he had evidence. However, always rejected according drift, so we may conjecture that he went along in 1942 because he did not to
Paradoxically, Mayr (1980d, 26) continued tomaintain thevalidity of
55
10
Simpson: No Straight and Narrow Path for Paleontology
In
as a presented the evolutionary synthesis theory that is compatible not if derivable the of with, from, findings paleontology.111 In particular, he showed that those findings did not require the invocation of orthoge netic rectilinear trends. This kind of trend "is a product of the tendency of of scientists to move in straight lines [rather] than of the ten to do so" nature for dency only specific evidence (Simpson 1944,166).The was the for natural selection, admittedly indirect, example of the horse. were Evidence that paleontologists in favor of abandoning orthogenesis natural selection may be found in the contributions of D. M. S.Watson the minds and Alfred Romer to the 1947 Conference on Genetics, Paleontology, and
son
his 1944 book, Tempo and Mode in Evolution,George Gaylord Simp
Evolution (Jepsen et al. 1949, 49, 58, 107-9). Simpson favored Wright's
a was drift over a purely selectionist explanation, preference that in his later book The Major Features Evolution (1953a).112 of reversed
111. Simpson, answers inMayr This (1980c, 452-63). article, based on Simpsons quoted to questions from or write an could not attend the workshops (because Simpson Mayr original article for it), seems to me to remember influences it forces especially valuable because Simpson and to articulate opinions that he might not have included if left to prepare a paper on his own. Of course that does not make itmore accurate or objective. s 1936 book, s considered Schindewolf Material Basis Simpson along with Goldschmidt of Evolution theoretical views were untenable. The (1940), to be "seriously misleading_Their necessity
thetictheory" (Mayr 1980c,457).

In his introduction
of considering
such opposing
views was
a definite factor in the formulation
of the syn
to the 1993
translation
says that in 1965, his advisor, Norman paleontologist,"
cated (Gould 1993).
despite
his opposition
of Schindewolf's 1950 book, Stephen Jay Gould called Schindewolf "the world's Newell, greatest living to the himself advo evolutionary synthesis which Newell completed overseas in
112. was in continues: "My (1980c, 456). He Simpson, quoted Mayr manuscript before Mayr's book appeared, and thereafter I could make no I was changes because the army until late in 1944, after my own book was (457). published"
56
Simpson:No Straight andNarrow Pathfor Paleontology
Gould and others,Simpson singlehandedly According to Stephen Jay

brought paleontology
Evolution (1949b), followedby amore popular exposition, The Meaning of

and a series of articles.113 At sis itself by adding laboratory and field studies of natural But Gould, a the same time, he strengthened the synthe a quantitative temporal dimension for evolution, which selection could not supply. had deviated from the evolutionary
into the evolutionary
synthesis with Tempo andMode,
wrote
was tologist, Gould particularly concerned with the role of Simpson, who had argued that fossil evidence, especially for horses, that had been used to support orthogenesis, could be better explained by natural selection. Gould that Simpsons 1953 book, The Major Features Evolution, of
paleontologist who a that synthesis by proposing theory of "punctuated evolution," complained the synthesis had evolved into a rigid dogma that stressed natural selection a to the exclusion of all other alternative or paleon contributory factors. As
differs from Tempo andMode_It...
some prestige, and (unfortunately in respects) hardened. In particular, in on reliance exclusive (for Simp creasingly selection-toward-adaptation son, in the gradual, phyletic mode), coupled with a greater willingness to reject alternatives more firmly than the evidence warranted, marks both Simpsons new book and the growing confidence of the synthetic theory in general_Though he chides others (quite properly) for assuming that
some subtle but important displays shifts in theoretical emphasis and content. These shiftsmirror some gen eral trends in the modern synthesis, as its theorywon adherents, gained
have some immediate use. (Gould
structures are inadaptive because they cannot imagine a use for them, he often constructs adaptive scenarios, in the speculative mode and on the must opposite (and equally invalid) assumption that prominent features 1980,166-7)
Moreover, Simpson had rejected Wright s genetic drift(whichhe had fa vored inTempoand words) "itcould not Mode), concluding that (inGould s
trigger any major evolutionary event."114 For Gould, devaluing the profession of paleontology: Simpson thiswas tantamount to had
in the period 1920-1940 of speciation were widely held [by German planations paleontologists] in 1952 (Rensch 1983, 35). On the diversity of responses to (and sometimes even later)"?even see Cain andMode, (2003). Tempo statement he quotes from is less dogmatic: 114. Gould "Genetic (1980, 168). The Simpson drift is certainly not involved in all or inmost origins of higher categories, even of very high cate
of paleontolo (2000). For details of one important effort toward collaboration Laporte see Cain in a new direction was a slow and (2004). But moving paleontology gists systematists, ex to Bernhard Rensch, "Lamarckian process, as Simpson himself pointed out (1950). According 113.
57
CHOOSING
SELECTION
unified paleontology with evolutionary theory, but at a high price in the price of admitting that no fundamental theory can arise deed?at from the study ofmajor events and patterns in the history of life. Why be a paleontologist if all fundamental theory must arise elsewhere? (Gould
1980,170)
in the history of extremely interested and well read was not an unbiased interpreter of primary certainly evolutionary biology, own sources. The of of his acceptance equilibrium theory punctuated extreme selectionism. Reinstating random of would the require rejection Gould, although genetic drift as a significant factor in evolution might help his case, al to face scrutiny by popula quantitative though itwould have been risky theories of drift developed by tion geneticists versed in the mathematical
Fisher
mind
Goldschmidt's theoreticalspeculations(Gould 2002,451-66). With

let us look at two of Simpsons and Goldschmidt) views not quoted by Gould.
a resurrection s to Gould and Wright. More advantage would be as without but of Goldschmidt's macromutations, empirical phenomena
this in
of sudden In 1967 Simpson statedthatthe theory major changes (Schin

dewolf, Beurlen, was
now almost no support for conclusively refuted by Rensch. In fact there is that view except by a few philosophers not sufficiently acquainted with scientific data on evolution. In particular, Goldschmidt s "systemic mutations" have
never been observed and ... need not be taken seriously if,as is the case, the phenomena that theywere postulated to explain can be explained in terms of known processes and forces.... In some of his latestwork Gold schmidt implicitly retreated from his position. (Simpson 1967,231-2) But at the same time, he was rejecting macromutations a role for (to Gould genetic drift.Having again allowing and orthogenesis by s dismay) hardened
his view against drift in the 1950s, he restored it as a secondary factor in the s founder 1960s. Drift "plays a role, perhaps a minor one," although Mayr more influence on evolution (Simpson and Beck 1965, 431). principle had
gories
such as classes known
positively Structure are made
next sentence, not or is: "It is not quoted by Gould, phyla" (1953a, 355).The to have been involved in any instance." Michael Ruse, in his review of Gould's metaphor?"Gould's facts
criticized his "notorious" hardening Theory, ofEvolutionary no matter what" (Ruse 2003,397). to fit the metaphor,
58
No Straight andNarrow Pathfor Paleontology Simpson: It provided a useful weapon contaminated the writings In the 1960s
pose or the existence of a goal in evolution. It equally renders untenable all the other theories that attempted to explain evolution by the dominant or exclusive action of one or another, such as the [n]eo single principle insistence on natural selection as essentially the whole Darwinian story....
Simpson ing of the synthesis that Gould protested: "The existence and persistence of apparently random as well as clearly oriented features in evolution [is] an unanswered the reality of pur argument against theories demanding
against the orthogenesis and teleology that still of some of his fellow paleontologists. was also rejecting in another way the harden
and random changes have ... a bearing on changes in broad Nonadaptive the appearance of new phyla, classes, or other major types of organization,
groups" (Simpson 1967,230-1).
59
11
Stebbins: Plants Are Also Selected
The
in botany had been responsible for yard Stebbins Jr. in 1950.115 Research two to the acceptance of the modern natural selection hy major obstacles in the twentieth century: de Vries's mutation pothesis early theory, based on his experiments with the evening primrose (Oenotherd), and Johannsens on research with beans.116 Yet "with the line pure exception theory, based of the fact that genes about genetics needed ment lie on chromosomes...
was tionary synthesis subject, by the publication
process of bringingthe major biological disciplines into the evolu
to most historians of the completed, according inPlants by G. Led of Variation and Evolution
about evolutionary theory from a botanist was long overdue. the assertion (see chapter 1) that natural selection is only Addressing cannot create and invoked a persuasive negative anything new, Stebbins a a sense it is negative only in the that sculptor's creation of metaphor: a statue by removing chips from block of marble is negative.117
searchon higherplants" (Stebbins 1980,139).Thus by 1950, amajor state
every other significant fact for the synthetic theory was worked out from re
are see Anderson (1999). (1952) and Kleinman polyploidy, which especially important for plants, on 116. "De Vries's theory that new species result from single mutations?a theory based a roadblock for botanists, as was Johannsens two kinds of varia claim that Oenothera?was quite
and 115. Stebbins (1950) and Smocovitis (1988, 1997, 2006). On the role of hybridization
tions exist and that all that selection did was to separate pure lines. I used to tell my students that at the Danish its nice neat Johannsen was a Dane looking landscape with gardens full of pure lines of peas and beans. His world included no conception of the confusion that exists in a tropi are cal rain forest or in the waste lots of the sub tropics, where weeds coming in and the entire
in Oenothera and other plants of polyploidy theories of de Vries and Goldschmidt. 117. Stebbins (1950,102). The
See also Stebbins (1950, 101-2) andDarlington (1980, 76-79). Stebbins (1980) notes the role
but does not admit was also that it gives any support to the
ecosystem
is disturbed. He
thought
that his world was
the natural world"
(Stebbins
1980,
146).
for example, Lerner (1959) and Mettler andGregg (1969).Holmes (1948b) argued at length
sculptor metaphor
invoked
by many
other authors;
60
Stebbins:Plants Are Also Selected
for natural selection was still meager. two experiments demonstrated it: Syl Among cross-fertilizing plants, only v?n showed in 1937 that clover adapts to a colder climate by elimination of But the direct botanical evidence on adaptation of ecotypes of the coast tarweed of California (Hemizonia
the lesshardyplants, andClausen, Keck, andHiesey (1947) presenteddata
on a few other rust (Potentilla Stebbins mentioned experiments gladulosd). in the he cited research self-fertilized Later book, by homozygous species. Har?an and Martini (1938) on barley as "the most extensive experiment yet on natural selection in the performed higher plants," which confirmed the prediction
and of of a sunflower of the tidy-tip(Layiaplatyglossd), hybrid, augustifolia),
for the "survival of al?eles in a heterozy by Fisher and Haldane acts Selection gous population. very rapidly at intermediate gene frequen cies, but more slowly at very high and very low ones."118 In the absence of
s research on the resis extensive data on plants, Stebbins included Demerec aureus to tance of the bacterial species penicillin (see chapter Staphylococcus on Gershensons and annual variations in seasonal 14), (1945) experiments and Dubinin swork on forms of hamsters, Dobzhansky Drosophilapseudoobscura, s results for The only com and Tiniakov Drosophila funebris.
melanic
was thatof mix W. B. Kemp (1937) for parable evidence forhigherplants

tures of grass and clover, separated into nearby fields subjected conditions (used for grazing or for hay).
to different
While
for it. In most cases, he believed, the popu lation is not small enough for it to be significant. It is likely, but not proved, that it applies to some plants inHawaii (Gouldea, Cyrtandia, Bidens), and 1950 he had lost his enthusiasm to the writer inwhich the action of random fixation or drift seems to have of small isolated populations
Stebbins had defended the Sewall by Wright effectin 1944,119
it is very likelythat it affects land snails."The only plant example known is thatof the complex ofPapaver alpinum in theSwissAlps, as describedby
part in the differentiation
taken a prominent
Faberg? (1943)" (Stebbins 1950,145-7).
selection is not purely negative since it includes variation. The assertion that natural is only a negative factor, so it cannot explain the adaptation of the organism that does evolve, is no longer taken seriously by biologists since there is good evidence of ample variation in to allow progress natural populations and enough favorable new mutations by selection. But the that natural selection
to cite Fisher and Haldane, but they also give no explanation ofwhere only do the authors fail towhich they got the "theoretical curve" they compare their data. Stebbins had also cited Har?an et al. (1947), and his own research on the competition and Martini, along with Sylv?n, Clausen between diploid species of grass and the autotetraploid derived from it, in Stebbins (1949).
not
issue is still debated byphilosophers(Nanay2005). 118. Stebbins (1950,109) andHar?an and Martini (1938).This is a rather paper: mystifying
119. Stebbins (1944); seeCain (2004, 68-69).
61
12
Chromosome
Inversions
inDrosophila
During
ered nonadaptive and therefore attributable to the effects of random ge netic drift,were instead primarily controlled by selection. First and perhaps on the views of evolutionists was the series most in its impact important of observations conducted by Dobzhansky and his colleagues (including Drosophilapseudoobscura.The originally wanted Epling background and motivation for this research of Wright
and 1950s, several biologists obtained empirical re sults that seemed to show that certain characters, previously consid the 1940s
Carl Epling and SewallWright) on the genetics ofwild populations of are described indetail byProvine (1986); Provine shows that Dobzhansky
to find a definitive detailed
but ended up doing just the opposite. theoryof genetic drift

s (1944) observations
empirical
confirmation
on the third chromosome
of D.
pseudoobscura suggested that different gene arrangements had evolved from a ancestor results were single by inversion of chromosome Chapters. His discussed symposium by Ernst Mayr, G. G. Simpson, and G. L. Stebbins.120 Epling had assumed that the gene arrangements were equal in selective value, but Mayr argued that "each gene arrangement... may have that the reduction in a 1945
at each is a consequence of the fact locality. This of crossing over in the inverted Chapters prevents the free recombination of genes" (Mayr 1945, 74). Stebbins agreed that some combinations would have a gene different selective values selective advantage under certain ecological conditions, an assumption ... has obtained some indirect evidence. I forwhich Dobzhansky agree
120. For 1944)
of the Committee
reprinted byCain (2004, 87-107).
further details, see the correspondence published on Common Problems of Genetics,
in Bulletin
no. 4 (November and
12,
Paleontology,
Systematics,
62
ChromosomeInversions in Drosophila of any distribution pattern over with Mayr that ... the maintenance or millions of years would be of thousands improbable unless periods the dispersal of the various chromosomal types were restricted by the
belief.(Stebbins1945)
selective activity of the environment....
Epling
... now holds the same
himself had already come to the same conclusion, as he ex Dobzhansky inversions inD. later The chromosome 1951,1971). plained (Dobzhansky are can be observed examples of adaptive polymorphism, which pseudoobscura in field studies but also under conditions where the laboratory produced relevant variables Two can be controlled:
A may be better adapted than B in summer,while B is superior toA dur ing thewinter season. (Dobzhansky 1951,109)
species, A and B, can be sympatric [occupy the same territory] only a provided that the environment in territorywhich they inhabit is het erogeneous. The heterogeneity may be spatial or itmay be temporal....
By shiftingthe balance back and forthbetweenA and B in response to

in the environment, natural selection allows the organism changes resources more one of the two to exploit the available efficiently than ifonly or B, were allowed to survive. A genotypes, seasonal These observations glands, which chromosomes. ous possible in the salivary made use of the giant chromosomes have a visible pattern reflecting the gene arrangement in the was observed. There are vari Usually the third chromosome
A and B and betweenE and F, giving A BCDE FGHI. be brokenbetween Then the middle sequenceBCDE is inverted, AEDCB FGHI. Fi giving to is the DCBFGH inverted, sequence nally leading AEHGFBCDI.
The first can arise from the second or give rise to the second through a same is true for the second and the third. But the single inversion. The
producing
or inversions, of the chromosome segments, rearrangements, a For ABCDEFGHI the may sequence polymorphism. example,
find in natural populations of some species only the first and the third arrangements, it is probable that the second remains to be discovered, or existence of previously unknown azteca was predicted gene arrangements in Drosophila pseudoobscura and D. with the aid of the theory of overlapping inversions, and most of these pre at least that it existed in the past_The
third can arise from the first,or vice versa, only through the second ar rangement as the probable intermediate step in the line of descent. Ifwe
63
CHOOSING
SELECTION
dictions were subsequently verified by discovery of the requisite inversions in nature. (Dobzhansky and Sturtevant 1938; Dobzhansky 1941a).121 that we have here an example of a confirmed novel prediction, on based the hypothesis that the chromosomes have evolved but not neces and Sturtevant 1938,33). It is com sarily by natural selection (Dobzhansky to the parable prediction of "missing links" from the general hypothesis that Note later organisms edly confirmed some (by process) from earlier ones, repeat in the late nineteenth and twentieth centuries by discoveries have evolved
of fossils. recalls, "It seemed at first that the chromosomal Dobzhansky polymor no now known that the contrary is the phism had adaptive significance. It is re case" (1951,114). The importance of this realization is reflected in the the inversion experiments are described
in a chapter now titled "Adaptive in "Chromosomal (instead of the noncommittal Polymorphisms" Changes" on now the second edition), which is placed immediately after the chapter "Selection," rather than before it. and experiments, I need present only a brief summary with population cages showed a seasonal variation of the
and the organizationof the thirdedition (1951) ofGenetics Origin ofSpecies'.
crucial observations
SinceDavid Wyss Rudge (2000a) has given a detailed account of these
here. Experiments an frequencies of the different patterns. Starting with experimental popu lation containing 11 percent of "Standard" (ST) and 89 percent of Chiri cahua (CH) chromosome, within four months 70 percent. the frequency of ST
rose to
was caused to elimi by natural selection. But if so, why has selection failed nate CH chromosomes altogether? The establishment of the equilibrium, are present in the gene pool with definite frequencies, is due to the chromosomal polymorphism being bal anced, because the heterozygotes (individuals having one ST and one CH atwhich both ST and CH chromosomes third chromosome) are superior in adaptive value to both chromosomal
It is clear, that the carriers of ST chromosomes had some adaptive ad vantages under the conditions of the experiment over the carriers of CH chromosomes, and that the rapid increase of the incidence of the former
(ST/ST andCH/CH). homozygotes
1951,114-5) (Dobzhansky
121. The quote including the two citations at the end is from Dobzhansky (1951,110-2); of the ori his figure 3 (111) shows the proposed mechanism for the inversions. For description of Natural Populations series, see Provine (1981) and Lewontin gin and results of the Genetics
(1981).
64
Chromosome
Inversions
in
Drosophila
What
to do with evolution? "Since the gene arrangement is a hereditary trait, we are dealing here with genetic a are evolutionary changes changes in the constitution of population. These se are definition. these Furthermore, by brought about by natural changes does this have in a chromosome lection." Moreover, studied under controlled it is an evolutionary process that can be quantitatively conditions: "The speed of the changes observed the hot season can be accounted for if the chromosomal at 25?C" they are observed to have in experimental s shift
in nature during types have adaptive values which
laboratory populations to Provine, According toward the NSH was in 1943, where
1951,118). (Dobzhansky the first public indication of Dobzhansky he
breaks rather abruptlywith the past: because of cyclic (seasonal) and year
to-year changes in the frequencies of... gene arrangements ... he con
cludes that these gene arrangements affect the adaptive values of their car riers.Cyclic changes ruled out chance, and mass migration was ruled out on by the studies dispersal. Thus the proportions of these gene arrange ments in a population were said to be subject to a surprisingly intense natural selection. (Provine, inDobzhansky But Provine under notes 1981, 303)
does not always keep up with changing views expressed in private. For our purposes, the public position as well as the private view is important, since for a scientist as influential as Dobzhansky, the public position is likely to persuade others. In a 1946 paper, Dobzhansky periments that natural selection microand macroevolution: asserted on the basis of his Drosophila could cross the supposed ex
that the inversions were "had concluded Dobzhansky two years before the of this paper, as is shown appearance" to Sewall dated May 4,1941.122 As sometimes happens by his letter Wright a scientist's in a research position fast-moving project, the public record of selection
barrier between
Some of the chromosomes
obtained by crossing over between the three ancestral wild chromosomes have properties very different from the latter. It is, therefore, possible to "select" products of recombination of the gene
complexes that deviate greatly from the ancestral types, being completely outside the limits of variability of these ancestors. 122.Provine (1986, 389ff). The discussion towardthe end of thepaper byDobzhansky and
to be an intermediate (1938, 61) seems with hindsight to inversions. adaptive description of chromosome stage in the transition from
Sturtevant
nonadaptive
65
CHOOSING
SELECTION
This
Dobzhansky
that his own work, along with that of Timofeeff in 1940?and first discovered cyclic seasonal changes Ressovsky?who of Dubinin the more recent publications andTiniakov, disproved the pre claimed vious view that "adaptive observed within a human evolution in nature is too slow a process lifetime."123
view published as the first article in the new journalEvolution (1947),
"great advances vored by artificial or natural
allows
in rebuilding the organism in directions fa selection" (Dobzhansky 1946, 288). In a re
to be
N. W. A brief note on these precursors acknowledged by Dobzhansky: was a for known best Russian geneticist prominent Timofeeff-Ressovsky on the reversibility of mutations.124 In his study of popula his research tions of the lady-beetle, he found a polymorphism maintained by climate selection pressure that favored the red form inwinter and the dependent in summer (Timofeeff-Ressovsky 1940). N. R Dubinin and G. G.
black
of an energetic selection which has changed funebris and found "evidence in response to changes in habitats, the genetic structure of populations contrary to the prevailing view that evolution proceeds very slowly" (1945, 572). All these results clearly pointed to natural selection as a powerful process that could produce observable
Tiniakov (1945, 1946a, 1946b, 1946c) studiedpopulations ofDrosophila
had proposed Dubinin himself help fromrandomgeneticdrift.(Recall that a formof driftin 1931.)
s was the to testimony, it surprising result of Dobzhansky According his own experiments that led him to change his opinion about the relative importance of natural selection and genetic drift: in these experiments, drift a clearly had negligible effect, contrary to what was previously believed.125
effects in just a few months, with no
On the otherhand, JohnBeatty suggeststhat Dobzhansky did not have

123. Dobzhansky in the "GNP" of the results. See was a coauthor of some of papers Dobzhansky's Wright on the not that suggests interpretation completely agree they did the editorial note in Dobzhansky (1981, 470). Burian (1994) suggests that (1947,1). While
series, Provine
as a not s natural selection rejection of macromutation phenomenon explicable by Dobzhansky should be seen in the context of his relations with his mentor Iurii Filipchenko, who coined the term "macromutation." 124. On
(1998); andReindel (2001). Junker

isDobzhansky 125. Here and drift: "In 1939, there was
the career of Timofeeff-Ressovsky, s later account initiated
see Glass of why
(1990);
Paul
and Krimbas
(1992);
he changed his views on natural selection at intervals repeated sampling approximately monthly in three ecologically of of the season, Drosophila pseudoobscura during populations breeding San Jacinto, in California. The purpose of this work was a rather different localities on Mount The results of allelism of recessive lethals in the third chromosomes.... study of the frequencies of the three localities, about 15 miles obtained seemed startling: not only were the populations but in two of the localities the chromo apart, clearly different in the chromosome frequencies, some were to month. That these changes could changing significantly from month frequencies
66
Chromosome
Inversions
in
Drosophila
to be full of variation" on which natural in particular the human species] selection could act. "Human variation might play a part in enabling the to to environmental various sorts." species respond adaptively upheavals of s "classical" This would help him make a stronger case against H. J. Muller in the human about the "genetic load" of mutations to its it him and would allow significance, judge "the fu population ture of the human species to be quite rosy."126Vassiliki Betty Smocovitis for Dobzhansky, this that supports interpretation, arguing "evolutionary position and in the debate
to to selection, enough empirical evidence justify his change from drift hence one must invoke Dobzhansky's "values": "He wanted [theworld, and
models
a stochastic view of favoring random genetic drift,which enforced not be favored in a postwar frame ofmind evolution?and culture?would
was out that random drift might be thought this plausible but also pointed a of evolutionary biologists: "A reassessment important in small population ... a few a by key people might trigger general response." (Dobzhansky, were all inNew York at that time.) and Simpson, Mayr City In the absence
world" (Smocovitis 1996,131). Gould (2002, 543) seekingto improvethe
to support the of documentary evidence Beatty s own statement to accept I Smocovitis interpretation, prefer Dobzhansky that the empirical evidence justifies the choice of the selectionist over the
drift hypothesis.
be caused by natural selection seemed hard to believe_The that the changes may possibility more have resulted from random genetic drift A de coup grace to this surmise plausible. appeared was administered are at least some of the the that regularly cyclic, follow by finding changes
seasons.... not have made But even this would the selectional ing the succession of the years some of the to many not been to believable explanation biologists, had it reproduce possible in artificial in and 1946).... Dobzhansky changes laboratory experiments (Wright populations ... evidence ... not all, natural The of D. clearly shows that many, though possibly populations ... in successive months. to be recorded pseudoobscura undergo genetic changes rapid enough It is certain that these changes are brought about by natural selection; random processes may contribute at most minor local fluctuations. The nature of the selective forces, the results ofwhich are so more than plainly visible, is still conjectural?despite thirty years of study" (Dobzhansky 1971,110-1,127-8). For the extension the paper by Levine of Dobzhansky et al. in Levine s research (1995,120-39). toMexican populations of D. pseudoobscura, see
126.Beatty (1987a, 300,303,305)
67
13
Ford:Unlucky Blood Groups

leader of theOxford School of Ecological Genetics, E. B. Ford,
The
tions: the prediction, based on Fisher s theory, that human would be found to be correlated with certain diseases. In 1940 Ford pointed that variations
and sponsored many experiments, testing the roles of nat the concept of balanced ural selection and genetic drift and developing 1940 he that this research confirmed the could claim polymorphism.127 By I discuss just one of his contribu importance of natural selection. Here conducted blood groups
theorem") caused by changes in its variability. This dardanus: we may
can be observed, for example, in the butterfly Papilio if fitness depends on mimicry of another species.128 Conversely, of a polymorphism, we find changes in the distribution of members infer that fitness is due to some as yet unknown a connection between the frequencies environmental factor.
out (as a consequence of Fisher s "fundamental in the fitness of one member of a polymorphism in the environment should be correlated with changes
Ford proposed to apply this idea to thehuman blood group polymorphism

by predicting of different groups and to susceptibility specific diseases. There had already been hundreds of reports claiming to find statistical conditions or correlations between the ABO groups and various medical
to 1921. characteristics psychological (including criminality), going back But these reports were not considered reliable by modern (i.e., 1940s) stan not did realize that because the dards, very large samples must investigators be used to obtain significant correlations.129Within the community of ex
Turner (1987);Ford (1957b; 1980,341-2); andRuse (1996a,457). In a balancedpolymor 127.

phism?for and nonmelanic moths?the example, between melanic so on not it is both obvious how strong they are sides, equally strong reduces the pressure against one form and allows it to predominate. text. For additional references to experiments whose results disproved selection pressures are often until changing environment See the quote in following the Sewall Wright effect, see
on Ford, seeB. C. Clarke (1995) andKimler (1983a). information Skipper (2000).For further 128.Ford (1940a, 505); see also Ford (1937). 129.Hardin (1952,418) andGarratty (1994,1996).
68
Ford: Unlucky Blood Groups on no such correlations were believed to have perts genetics, population or to be established.130 Ford made his been demonstrated likely prediction more in 1945: explicit Individuals belonging to the different blood groups are not equally viable, and we may expect elimination to fall upon theAB class.... A valuable line of enquiry which does not yet seem to have been pursued in any detail
would be to study the blood group distributions in patients sufferingfrom a wide some conditions infectious variety of diseases. It is possible that in
or otherwise, theywould depart from their normal frequencies, indicat ing that persons of a particular blood group are unduly susceptible to the (Ford 1945, 85)
disease in question. Fords
was confirmed in the 1950s.131 The initial report by prediction London medical researchers Aird, Bentall, and Roberts (1953) stated that are more with blood A persons group likely than those with significantly O to have cancer of the stomach. They did not mention Ford s predic tion. In a more comprehensive article the next year, which also reported an association with peptic ulceration, these authors, together with Mehigan, a not precise prediction but did provide citation (Aird et al. 1954, 318). A team at the State University of Florida a survey of a published early results, noting that Ford (1945) "hinted at" relationship between blood groups and diseases "in suggesting the possible did credit Fisher and Ford with a to natural selection."132 relationship of blood groups concluded that firm correlations J.A. Fraser Roberts lished between duodenal ulceration had been estab
and group O and between pernicious anemia, as well as stomach cancer, and group A; he also found "fairly strong evidence for an association between group A and diabetes mellitus." In a very brief theoretical comment, he did not mention Ford but asserted: "Fol
groups, see Box (1978, chapter 13). to Ford (1957b), 131. According the first confirmation of his prediction was by Struthers in babies and found the incidence higher with group A (1951), who studied bronchopneumonia than group O. A. E. Mourant the success of (1959), an expert on blood groups, acknowledged Ford's prediction and admitted that "it is unlikely that any genes are completely unaffected by natural selection"(Mourant 1959, 60); yet he insisted that we still don't know "the mechanism to the diseases mentioned, if indeed the relation is one of whereby blood groups affect liability cause and effect" (59). a successful he would have traded for a plausible Apparently prediction explanation. 132. Buckwalter et al. (1956b, 1215); see also Buckwalter et al. (1956a, 1956).
130. Aird, Bentall, Roberts (1953), and Kitchin et al. (1959). Of course the prediction was on Fisher's to Michael Ruse (1996a), such successful predictions helped theory. According in research Ford to get grants to support further research. On Fisher's interest and participation based on blood
69
CHOOSING
SELECTION
con a neutral gene is almost unthinkable-A long lowing Fisher [1930a] on an must must be dynamic, not static. It tinued polymorphism depend (Roberts 1957,124). interplay of selective advantages and disadvantages" and disease connection, along with Dobzhansky's covery of the blood-group in "rather dramatic turnaround" (triggered by his results on chromosome versions) and thework of Cain and Sheppard on Cepaea, as part of a turning of the tide against drift and "overwhelming support" for selectionism.133 Ford was of course entitled (Ford 1957b); particular case
A multiauthor 1989 book on thehistoryof probability portraysthedis
to claim victory for his own theory in this in at the Darwin centennial celebrations
His
Wright had alreadychangedhis views by 1951,when he suggestedthatthe

distribution selection of blood group frequencies may be the result of "intergroup ... a certain balance between local isolation and cross breeding"
and colleagues were also quick to point out the new port for selectionism in the ongoing controversy with Sewall Wright. students
1959, he told the audience that his 1945 prediction about the selective controlof human blood groupshad now been confirmed(Ford 1960,194).
sup But
I could not find any evidence that other biologists outside Nevertheless, to accept his selectionist views because of this Ford's group were persuaded was a novel nor it that made confirmation, any difference whether it predic tion rather than a theoretical deduction of known facts.134
(Wright 1951,454).
Genetics, and Man,
blood groups (Dobzhansky andMontagu
as an important role in evolution adjunct, natural selection (Beatty 1987a), particularly an still wanted
Before 1950Dobzhansky had supported Wright s view thatdriftplays

not a substitute, for though in the evolution of human
1947). In his book Evolution,
to in Dobzhansky explain the differences as a result of blood groups among different populations genetic drift, ignor recent connection with diseases, even their the discoveries concerning ing
was aware of Allisons work on the connection between sickle though he cell and malaria 1955, 130, 143). In a survey article in Sci (Dobzhansky ence, he stated, "The functional significance of the blood groups in man is article he still found the evidence "inconclusive."135 group In his book Mankind
stillfull of uncertainty"(Dobzhansky 1958, 1093), and in another survey

he cited the evidence for blood connections with diseases
Evolving,
and smallpox)but triedto explain themby (now including plague, syphilis, drift genetic (Dobzhansky 1962,280).
133. Gigerenzer et al. (1989,157); John Beatty is probably responsible for this chapter.
134. Sheppard (1958,1959a); Dowdeswell (1958); andC. A. Clarke (1961,1971). inBajema (1971; see especially12). 135.Dobzhansky (1961); reprinted
70
Ford: Unlucky Blood Groups
mention
groups as examples of adaptively neutral traits goaded correlates of the blood groups many investigators to discover physiological A relationship among the classical A, B, and AB that are not neutral.... and gastric ulcers is probably the most thor blood groups and duodenal of blood
theOrigin ofSpecies), way: "The frequent Dobzhansky told the storythis
In Genetics of theEvolutionary
Process
(the fourth edition of Genetics and
not give Ford any credit for his prediction oughly established." But he did and downplayed the significance of the discovery that these characters are not thus far discovered are with dis adaptively neutral: "The associations eases ages, so that the Darwin affecting mainly people of postreproductive ian fitness is but little changed." Moreover, he insisted that the connection
with disease
could not explain the observed differences in blood group fre omit the suggestion, among local populations, quencies although he did in differences are a result of genetic Mankind that those made Evolving, drift.136 founder principle, which he now said was supported by the work of his own the "hyperselectionism" of other evolutionists, group.137He denounced new interest in neutral or non-Darwinian although he admitted that the evolution was perhaps just another "swing of the pendulum" was just (Dobzhansky symp
his affection fordrift, Mayr s especially By 1970Dobzhansky had revived
1970,262).
One might suspect that hyperselectionism
a nationalistic
in chapter 9. Ernst Mayr, whose views were discussed While Mayr did as sert that because selective is "it rare, appears probable that random neutrality fixation is of negligible evolutionary importance,"138 his continued support seem to as a for his own founder principle would hyper disqualify him s in quoting Mayr, had left out part ofMayr selectionist.139 Dobzhansky, statement, a sentence that explained why he rejected genetic drift as a major in the past as caused by factor in evolution: "Virtually every case quoted
tom ofBritish distastefor the American drifttheory(Gould 1983;Turner one of But the 1987). by Dobzhansky was hyperselectionistsidentified
errors of more genetic drift due to sampling has recently been interpreted in terms of selection It is hard to believe that pressures" (Mayr 1963,207). this is literally true, since so many authors had invoked drift to explain so many different examples of supposedly nonadaptive illustrate the rhetorical strategy of the selectionists, characters. But followed it does also by Cain
17 herein.
136.Dobzhansky (1970,296-8; 1962,280-3). 137.Dobzhansky and Pavlovsky (1957);Dobzhansky and Spassky (1962); and see chapter
138. Mayr (1963,211), as quoted byDobzhansky (1970,262). 139. Mayr (1970,120). Mayr (1963,204); cf.
71
CHOOSING
SELECTION
drift have been
shown to be really due to selection (and by implication, the converse has not then it's reasonable to suspect that all other happened), ascribed to drift are also really due to selection. Of course the phenomena rhetorical force of the statement was enhanced if some of these reversals
(1951a) and Sheppard (1967): if severalphenomena previouslyascribed to
were
confirmed novel predictions and if,as Provine argued, "the three most chromosome inversions, frequently cited examples" of drift?Drosophila in and all found and blood snails, banding coiling patterns groups?"were to be to substantial or even enormous subject
and other theorists who discussed genetic drift often insisted Wright that the fate of such a very small subpopulation was more likely to be com once a gene to plete extinction of the drifting gene: frequency fluctuates zero, it can't fluctuate back to a finite value. Blood group researchers found that some subpopulations surveys, according to Neel of American and Salzano Indians?sixty-two (1964)?completely out of 109 lacked the B
to this issue in thefinal chapter.) 1983,65). (I return
selection pressures"
(Provine
This isprobably group. why, according to Slatis (1964), thedistributionof Brues (1954) had argued, the fact that four-fifths of all possible combi
nations of frequencies were missing might better be explained by natural selection. et al., "While to selectionist ac Gigerenzer According Dobzhansky's count of chromosome in has stood the test of well pretty shapes Drosophila ... of blood groups in humans have not.... time, the selectionist accounts Italian has successfully accounted for dif [The biologist Cavalli-Sforza] frequencies in this population is often attributed as to drift, even though,
inNorth group frequencies among different populations terms of drift alone."140 to Linda in Stone and Paul F. Italy According in 1964.141 first conclusive evidence was published Lurgin, Cavalli-Sforza's Turner
ferences in blood
play a decisive role in thedebate of the 1940s and 1950s.

The association
(1987) argues that this research provided the missing evidence for drift, but from my viewpoint, it came too late (like Kettlewell's work) to real between blood groups and disease is undoubtedly a were not to in but the 1960s able satisfac give biologists (Garratty 2000), was out association. for the It tory explanation by several experts pointed on that the blood group polymorphism population genetics theoretically, should not be stable, because the heterozygotes do not have a selective ad
140. Gigerenzer see note quotation, 141. Stone
et al. (1989, 133.
157),
citing Cavalli-Sforza
(1969).
On
the authorship (1964).
of this
and Luquin
(2005,59-74,209)
and Cavalli-Sforza
and Edwards
72
Ford: Unlucky Blood Groups over the homozygotes?unlike morphism, where they do.142 vantage Something
in the sickle cell and malaria
poly
similar happened with three of the best known predictions of elementary particles in physics: the positron, the meson, and the Or (omega minus meson). In each case, the theory that predicted the particle was soon until that happened, the success of the prediction Nevertheless, be considered evidence for the theory. could
found to be inadequate andwas replacedby another theory(Brush 1993b).
still
142. Allison (1955,1968); Lewontin (1967; 1974,235);Mettler andGregg (1969); andC. A. Clarke (1977,487).
73
14
Resistance
toAntibiotics
Another the first to discover crobes attacked
to medicine application of natural selection . the same time as Ford s Ren? J.Dubos, prediction.
appeared around who was one of
phenomenon as or "fastness," but its cause was not understood. It might have "training" to the well-known been considered analogous of resistance phenomenon are to insecticides, be there differences although significant biological
an antibiotic that the mi drug (gramicidin), warned to them.143 This resistance such would by drugs develop had already been observed with other drugs and was known
tween the two (Simon 2003).
In a comprehensive treatise on The Bacterial Cell, Dubos reviewed experi occur in small ments made "to establish whether the resistant bacteria always
to historian According Creager, before the 1940s bacteriolo Angela af gists favored the Lamarckian explanation. But the notorious Lysenko on Lamarckian cast fair in the Soviet Union theories and may suspicion have made considers
numbers during normal growth in the presence of the drug, or whether they are as a response to the presence in the medium of the sub produced only stance with reference to which resistance 1945, 322). In develops" (Dubos or a Lamarckian other words, is resistance a Darwinian phenomenon?
some scientists more to selectionist theories. receptive Creager to be the 1943 experiment by Salvador Luria and Max Delbriick "one of the first clear demonstrations that inheritance in bacteria was not Dubos
Lamarckian"
(Creager 2007,166). (1945) concluded that the evidence favors the Darwinian expla nation of resistance: it is due to selection acting on the preexisting variabil
to be resistant to the that happened ity of the microbes. Those drug could survive and pass on this attribute to their descendants.
was "awell established fact that strains of bacteria resistant to By 1944 it various sulfa drugs, as well as strains resistant to penicillin, may readily be obtained by growing bacteria inmedia containing increasingly higher con 143.Dubos (1942,1944) and Moberg (1999).
74
Resistance to Antibiotics
at Cold experiment in 1944 byMilislav Demerec Spring Harbor. Demerec, strains of found that "makes it probable evidence using Staphylococcus aureus, that the second [Darwinian] alternative is correct": resistance of bacteria to antibiotics is a result of selection.144 a published, in ScientificAmerican, good popular explana s as a prominent example of "evolution in the experiment were a little too laboratory." For those readers who recklessly the following neo-Darwinian command "Make love, not war," he noted that "in certain Dobzhansky tion of Demerec cities penicillin-resistant gonorrhea has become more frequent" (Dobzhan view
or was some of the bacte by the action of the antibiotic already present in ria as a result of earlier mutations. This question was answered in a classic
centrations of the respective chemicals." What was not established, despite was whether the resistance was induced the Luria-Delbriick experiment,
sky1950b, 33,35).
Dobzhansky supported
the Darwinian
view over the Lamarckian
cies (1951), which "made antibiotic resistance a crucial?and observable? at work" of natural selection example (Creager 2007,176). Further confirmation came from a classic experiment by Joshua L. Le derberg and Esther M. Lederberg: They showed that bacteria could display an inherited resistance to it.The disposed
in the thirdedition of his influential book, Geneticsand theOrigin ofSpe
already present,
a predictionby don't saytheyare confirming Dubos, JoshuaLederberg they

studied with Dubos This at Columbia and later recalled that Dubos's Bacterial from which work was I can say I learned most of the microbiology the launching pad for my own investigations"
even if never been to directly penicillin, they had mutations were not caused by the penicillin but were thus excluding a Lamarckian interpretation.145 Although
Cell "is the work I know....
who were
interested in practical applications of the theory. Dubos himself seemed to regard it not so much as an important scientific discovery but more as an illustration of his not try to general philosophy: humans should conquer nature (in this case by using powerful drugs) but should try to live in harmony with it.146
to antibiotics cannot be called a novel pre Although bacterial resistance diction from natural selection, its confirmation did provide important sup port for evolutionary theory, especially for those scientists and physicians
(Lederberg 1990, introduction).
144. Demerec the experiment,
145.Lederberg and Lederberg (1952) andLederberg (1989); for a textbook descriptionof

see Curtis (1968, 695-6).
(1945,16,23);
see also Creager
(2007,167?F).
146.Cooper (1998) and Moberg (2005).
75
15
Two Great Debates:
Snails andTigerMoths
One
was
in small partially isolated described byWright's theory, populations"?as or in the same book?and not to pure isolation presented geographical
on his studies of two report by Cyril Diver closely related spe cies of snails, in the same areas nemoralis and Cepaea Cepaea hortensis, living in England and elsewhere in the United States and Europe. He attributed the split of the presumed earlier single species to "random differentiation
of the chaptersinJulian New Systematics (1940) Huxley sbook The

a
in a new investigation of Cepaea nemoralis, designed to testDivers conclusion and perhaps to settle the Ford versusWright de bate of the late 1940s about the relative importance of natural selection and at Oxford collaborated random drift.This was Wright,
natural selection (Diver 1940, 327). Cain and PhilipM. Sheppard After the end of World War II,Arthur J.
the beginning of what Provine, in his biography of it in great calls the "Great Snail Debate." Since he has described
detail, I need only summarize his account and conclusions.147 Cain and Sheppard, in their first paper on C. nemoralis, found (contrary to Diver) that color and banding patterns "have definite selective value" that speciation predation by thrushes. They concluded should be ascribed to natural selection, not to genetic drift, and that other cases ascribed to (Cain and Sheppard genetic drift should be reexamined in connection with
that might have been expected. Since Dob to support their zhansky and Mayr had used Diver 's results pro-drift (non in the early 1940s, Fisher, Ford, and other selectionists views adaptationist) would would
1950, 275). According to Provine, this resultdid not play the role in the
selectionist versus drift debate
tryto defendhis theory Wright would therefore by arguing thatgene fre

147. Provine 1986, chapter12); see also theobituary of SheppardbyC. A. Clarke (1977).
that this evidence against the importance of drift have anticipated be considered evidence againstWright's theory of evolution and that
76
Two Great Debates: Snails and TigerMoths were
quencies he was
. .. were to believe that the gene governed frequencies prepared primarily by natural selection rather than by random genetic drift" (in the case of "conspicuous" a letter from Wright to Provine), as shown according by polymorphisms, to Cain inNovember La 1950. Even afterM?xime
read theCain and Sheppard [1950] paper on polymorphism in Cepaea,
strongly influenced by drift. But
in fact, "even beforeWright
motte
to be nonrandom."149 Carter, who properly studied has been shown voked genetic drift in his recent book on animal evolution (1951a),
can postulate "drift"just because they "personally not see" any in a variation. Then using the rhetorical possible selective value later (see chapter 13), Cain asserted, "So far, ev strategy employed byMayr ery supposed example of random variation [or "genetic drift"] that has been such as G. S. Carter, who had in tried to
tion, except as itmay interact with natural selection; he was not really inter ested in invoking drift to explain neutral characters, which other biologists seemed to think provided itsmost useful application.148 in Nature when he criticized biologists Cain also started a mini-debate
own results on in France, which he published his Cepaea populations as of insisted supporting Wright's genetic drift, theory Wright interpreted it is not important for evolu that while genetic drift is a real phenomenon,
or selection without not claim that there is proof. He did characters (Cain 1951b). Carter replied, "I am of nonadaptive a real factor in evolution.... glad that Cain admits that genetic driftmay be We should ... regard it and selection as equally possible explanations when neither is proved" (Carter 1951c). In the second edition of his book, Carter either randomness no evolution retreated a little by adding the statement "Adaptation differences in their environments may often be more of the demes to small important than drift" and by adding a reference to the 1950 Cain and Sheppard paper.150 In a 1957 book on evolution, Carter omitted genetic drift entirely. to the nat to Sir opposition According Cyril A. Clarke, the American
148. Provine 1952, 1959); (1986,441-9 see also Lamotte [and references cited therein]; quotation on 441); Lamotte he discussed (1950), a less well-known paper in which (1951, selec
turn the tables by accusing Cain of claiming that "selective value should be assumed in all characters until the contrary is proved" (Carter 1951a). Cain denied taking that position, maintaining instead that one should not assume
to at the end of Millstein remark published (1955). According Sheppard a case for most at to other researchers the "was able make unlike time, (2007a), Lamotte, good a that the role for of different he showed sizes; drift, many significant by studying populations as fluctuations were greater in smaller populations, (2007b) gives predicted by theory."Millstein tion, and his discussion a detailed account of the work of Cain 149. Cain (1951a, 424). Turner and Sheppard, as well as of Lamotte. calls this the "fairies in the garden" argument against drift
(1987, 342).
150. Carter (1954,247,262).
77
CHOOSING
SELECTION
ural selection
editor [of thepublished proceedings].This injudiciousact rousedPhilip to (Clarke 1977,477). fury"

Although was cades, it selection the Great Snail Debate continued considered genetic in the 1960s drift. As to have next few de through the a victory for natural produced Gould wrote in 1982, itwas
are in hypothesis, particular the belief that polymorphisms was in still strong 1959. Sheppard's paper, presented at adaptively neutral, in that year, which was "the only con the Cold Spring Harbor Symposium was deleted reasons tribution which gave facts and by the against neutrality,
over
"one of the centerpieces
stillrecalled it as amajor victoryforadaptationism (Gould 2002, 541).
Stephen Jay of the adaptationist program";
twenty years later, he
Fisher
and Ford conducted another test of genetic drift theory Wrights two colonies of the moth Panaxia dominula (scarlet tigermoth), con using near Oxford. They measured fluctuations in the frequency veniently located of a heterozygous form, medionigra, easily recognizable by the colored wing in 1947 that these fluctuations were pattern it produces. They reported
must be larger than estimated from random drift theory and hence com in his caused by changes in the selective forces. reply (1948), Wright, and that furthermore, it plained that his theory had been misrepresented to assume that any effect not due to driftmust be due to was not legitimate were further replies back and forth, leading not to a resolu selection. There much tion of the controversy but to great animosity some stimulus to useful further research.151 among its participants and to
151. Provine
the 1990s. per (2000) follows itthrough
(1986)
gives
a detailed
account
of this controversy
through the 1950s,
and Skip
78
16
Dunkers
and Drift
showing that fluc tuations in the gene frequency of the medionigra gene are "too large The for such shifts to be attributable to genetic drift [and therefore are] due to ... has no direct on the selection[,] question whether the fluctua bearing result of Fisher dorninula, tions at some other locus are attributable
and Ford on Panaxia
a 1952 paper byBentleyGlass, Milton S. Sacks, Elsa F. Jahn and Charles Hess at theBaltimoreRh Typing Laboratory of theJohns Hopkins Uni
"The real difficulty, as Sheppard versity and the University ofMaryland. sure to is of neutral survival value has be that characters [1951] pointed out, ... itself varies are such, especially since survival value undoubtedly actually from year to year and place human populations known selective advantage to
to one or the other," according
to
with
more useful to investigate place." It might be to be derived from very small breeding groups; a an effective size of 200 or fewer members, "an al?ele would require
of 0.01 to avoid the actions of drift, and consequently et al. 1952,145,146) be many genes might effectively neutral." (Glass Such a group is the "religious isolate" known as the Dunkers, living in Franklin County, Pennsylvania. They descended from a sect of the German toAmerica in the early eighteenth cen Baptist Brethren, which migrated into in and then 1881. and his colleagues three Glass groups tury split up traits believed to be nonadaptive, chose to measure including the ABO blood groups. They found variations in gene frequencies, as compared with in Germany and the United States: 59 per typical samples of populations percent inGermany and 40 percent smaller correspondingly frequencies these and other variations "aremost (Glass et al. 1952,158). isolated human groups, Glass in the United in B, AB,
centhad A blood group in the Dunkers, significantly greater than the 45
had States, and Dunkers that and O. They concluded to genetic drift" reasonably attributable In a 1954 review, which included research on other
stated: "It is hard to see how any factor except account drift for such divergent gene frequencies as could genetic possibly in Northern Greenland_The Polar Es those of the Eskimos ofThule
79
CHOOSING
SELECTION
kimos have a veryhigh frequency of the ABO blood group al?ele Io and a low frequency of the al?ele Ia" (Glass 1954,130). correspondingly Neither paper from theGlass group citedFord s 1945 prediction that
groups would be found to have selective advantages or or the 1953 confirmation of that disadvantages prediction by Aird, Ben tall, and Roberts. Yet despite the large amount of research in the 1950s the human blood and 1960s adaptively their assumption that blood group al?eles are as cited the Dunkers neutral,152 biology textbooks occasionally that undermined
an example of genetic drift(Elliott 1963,1968).
McKusick (1978); Falconer (1981, 76). Hopkins; see
152. Glass did acknowledge this later (1962). Another possible effect is in the Amish communities studied by Victor A. McKusick
example of the founder/drift at Johns and his colleagues
80
17
Harden? Gould:Why Did the Synthesis The Changing Views of Wright Dobzhansky and
1983 Stephen JayGould extended his complaint about hardening Wright, the two foundersof the In beyond Simpson toDobzhansky and
synthesis who had tempered natural selection by genetic drift within in their isolated populations of speciation. He quoted explanations small from
on selection and to show Wright, "increasing emphasis adaptation." As for he now denies that he had ever "advocated a radically non-Darwinian ap to in selection and proach evolutionary change by demoting adaptation the contrary, his shifting-balance theory, "which does specify an important role for genetic drift, is strongly adaptationist? but that adaptation arises at a level higher than the traditional Darwinian On focus on individuals." Gould ent favor of accident."
and the thefirst(1937) and third(1951) editionsofGenetics Origin ofSpecies
s pres accepts this interpretation of Wright assumes a it unfamil "hierarchical of type theory, although thinking" to most evolutionists iar and uncongenial But he 1983, 85-86) (Gould notes that in for research his William Provine, forthcoming biography of has significantly changed direction: his own views since
found that Wright, Wright 1930 in a pro-adaptationist The
careful reader in 1932 would
believed nonadaptive of races, subspecies, and perhaps genera. Wright's more recent view that the shifting balance theory should lead to adaptive responses at least by the subspecies level is found nowhere in the 1931 and 1932 papers.153
almost certainly conclude that Wright random driftwas a primary mechanism in the origin
153.Gould, quotingProvine (1983, 86).
81
CHOOSING
SELECTION
Provine
theoryin the same 1983 book (Provine 1983). His biographyof Wright,
published Wright's confusion. The doned in 1986, documented refusal to acknowledge the change of views in more that change, and described detail, noted the resulting
supported Gould
hardening
thesis in his article about Wright
Drosophila
as a to with Dobzhansky had caused Wright major fac reject random drift same time tor inD. at Genetics exactly the Dobzhansky's pseudoobscura while
were able to in which the cyclic they duplicate pseudoobscura, in inversion in found natural changes frequencies populations by using the and Teissier. "Working population box technique pioneered by L'H?ritier
reason aban and Dobzhansky primary why both Wright drift in favor of selection was the result of their joint research on
and the Origin ofSpecies (in itsfirsttwo editions) emphasized randomdrift" (Provine1986,403).
Leaving aside Huxley and Wright and Mayr for the moment, we must ask: why did
move from a selectionist Dobzhansky theory incorporating random drift to a purely or primarily selectionist theory (what I call the in the 1940s? Their switch surely has a lot to natural selection hypothesis) do with explaining why other leaders and supporters of the synthesis began to favor a hard version. Gould discussed Dobzhansky s shift toward strong selectionism in
to the 1982 reprint his introduction of thefirstedition (1937) of Genetics and the with the thirdedition (1951) Comparison of this Origin ofSpecies. showed that,likeSimpson and Wright, Dobzhansky had hardened his the oryby deleting
the two chapters that contained most material on nonadaptive or non selected change (polyploidy and chromosomal changes, though he in cludes theirmaterial, in reduced form,within other chapters). He adds a
on He argues that chapter "adaptive polymorphism" (1951,108-34). or anagenesis, "progressive" evolution, works only through the optimizing, ... But the winnowing agency of selection based on competitive deaths most remarkable addition occurs at the beginning. I label it remark right I doubt that Dobzhansky really believed what he literally sure I said: feel that he would have retracted or modified it had anyone a fascination for pointed out that he had allowed adaptationism to displace the oldest of evolutionary truths. In this addition ... Dobzhansky poses the key question ofwhy mor so a cluster of somany cats, another phological space is "clumped":?why of dogs, a third of bears, and so much unoccupied morphological space able because
new
between?(Gould 1982)
82
Gould: WhyDid
Harden? The Changing Views ofDobzhansky and Wright theSynthesis
s to support and rejects Wright adaptationism "adaptive landscape" model as "skewed" and irrelevant to evolution his explanation of "clumping." After s new argument for selection, two pages devoted to refuting Dobzhansky Gould offered only one paragraph to explain why Dobzhansky, Simpson, and others had become more favorable to the natural selection hypothesis since 1940: I do not fully understand why this hardening occurred, but I regard it as so dominated important topic for historical research since its result the research program of evolutionary biology formany years. In part, the "ecological genetics" of E. B. Ford and his panselectionist school in En
Gould went on to accuse Dobzhansky (who died in 1975) ofmisusing
an
s own Dobzhansky empirical work increased his belief in the power of selection. In 1937, he tended to attribute inversion frequencies in natu ral populations ofDrosophila to genetic drift, but he then discovered that these frequencies fluctuate in a regular and repeatable way from season to season, and decided (with evident justice) that theymust be selective.
colleagues and Schmalhausen, who held that selection and once denied itmost adaptation might triumph in the very realm that had s 1951 argument for continuity between micro- and strongly. Dobzhansky is much the 1937 version. Finally, than macroevolution stronger (p. 17) like Simpson, Rensch,
ficients in nature buoyed strictDarwinian faith. Dobzhansky have gained confidence from the claims of macroevolutionary
a Their commitment to adaptationist gland must have had major effect. their of and everything discovery of strong selection coef explanations must
also
the evolutionary significance of certain peculiar mutations ofDro I infer from Gould's remarks the suggestion that Dobzhansky gave sophila. own area to in his the of his "first love"?entomol results greater weight perceive ogy?than to evidence
In the epilogue to his introduction, Gould stressed Dobzhansky as a "trained taxonomist and ground entomologist," which allowed
s back him to
facie evidence
for this suggestion: there are more than one hundred refer ences to to four to E. B. Ford, twelve to G. G. Simp compared Drosophila, to to 1.1. Schmalhausen. nine But Dobzhansky thirteen B. and son, Rensch, it clear that the theoretical is based ex interpretation of his Drosophila ... on "the of balanced theory developed polymorphism contrary towhat one might expect from Gould
and the indexto the 1951 editiono? Genetics providesprima Origin of Species
from other organisms
(Gould
1982).
Indeed,
the
also makes periments
byHaldane, Fisher,Ford, Wright, and others"(Dobzhansky 1951,116).

By 1957, Dobzhansky,
83
CHOOSING
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account, was
own of the campaign against the hardening launching his a synthesis. In paper with Olga Pavlovsky, he attacked the view that natural can be is selection all-important, while drift ignored. As soon as a gene is shown to have any effectwhatever on fitness, the conclusion is drawn that its distribution in populations must be deter mined solely by selection and cannot be influenced by random drift. But this is a logical non-sequitur. The important work ofAird et al. (1954) and of Clarke et al. (1956) disclosed that the incidence of certain types of
gastro-intestinal ulceration is significantly different in persons with differ ent blood groups. This is, however, far from a convincing demonstration that the observed diversity in the frequencies of the blood group genes
in human populations is governed wholly, or even partially, by selection for resistance to ulcers. To make such a conclusion tenable itwould have to be demonstrated that the environments inwhich human racial differ ences have evolved actually favored greater resistance in certain parts of theworld and lesser resistance in certain other parts. Thus far no evidence
has been adduced
to substantiate any such claim. (Dobzhansky
and Pav
1957,311) lovsky
tested In a carefully designed experiment, Dobzhansky and Pavlovsky the prediction that because of genetic drift, a small population will evolve with greater genetic diversity than a large one starting with the same fre even when both are to strong selective forces. It was also quencies, subject a test of s founder Mayr principle. Ten small populations (twenty founders ten D. and of founders each) each) (4000 large populations pseudoobscura constructed to have a 50:50
were
ratio of PP (Pikes Peak) zndAR (Arrow were in their third and head) gene arrangements chromosome, kept they in population (about cages under uniform conditions for eighteen months nineteen
s earlier outcome of the selection zhansky experiments (chapter 12), "the in the is conditioned by random genetic drift. experimental populations a and The either-selection-or-drift point of view is fallacy" (Dobzhansky
experiment.
had generations). The results showed that the small populations significantly greater variation in the PP andAR frequencies. "Although the trait studied ... is subject to powerful selection pressure," as found inDob
Pavlovsky 1957, 316). Mayr's founderprinciplewas also verifiedby the

The Dobzhansky-Pavlovsky stration of the phenomenon is regarded as a classic demon of genetic drift,154 and it certainly shows that experiment
154.Brousseau (1967); Ruse (1981, 32-33); andReisman and Forber (2005).
84
Gould: Why Did
Harden? The Changing Views ofDobzhansky and Wright theSynthesis
can be tested under theory predictions from the Fisher-Haldane-Wright controlled conditions. But does ithave any relevance to evolution in nature? and Pavlovsky argued that it explains the fact that populations Dobzhansky
Mani?la of the butterfly jurtina, "rather uniform throughout Southern En some obvious environmental gland, despite diversity in different parts of the territory [,] ... show quite appreciable divergence on the islands of the each other and their environments 1957, 317). This appear rather uniform" (Dobzhansky was attributed to selec entirely divergence
Scilly archipelago,although these islands arewithin only a fewmiles of

and Pavlovsky
statements (1937, 1941) that the situation arose skywithdrew his earlier in random drift of continuously small size, or fre "through populations narrow 'bottlenecks.'" Instead, "in the island popu quently passing through lation we observing the emergence of novel genetic systems moulded interaction of random drift with natural selection" (317). are by
tion byDowdeswell and Ford (1953) and Ford (1964, 54-58). Dobzhan
ascribed they
But Ford strongly fromScilly could be rejectedtheview thathis results

are or any other kind of drift, to the founder insisting that principle to natural selection (Ford 1975, 66-75). due completely
to anti-drift has already been dis Dobzhansky's change from pro-drift and Provine cussed at length by Beatty, Smocovitis, Gould, (see chapter 12). It remains to note only that by 1970, he was beginning to suspect that itwas time to reverse his position again, in the light of recent "theories of non-Darwinian ries,which were Jeffrey Powell evolution these theo by random walk." He characterized as new to become more in "a the 1970s, swing of popular as a drifter and then became a selection
thependulum in the opposite direction"(Dobzhansky 1970,262). He told ist. Now, inmy old age, I findmyself becoming a drifteragain" (Powell 1987,34). What is thispendulum? In the 1930s itpointed to "Wrights random
in 1974, "I began
assumed to be products of natural selection" (Dobzhansky 1970, ran as was it illustrated Now 262), (1963) position. suggesting byMayr's domness once more. I will reserve my own speculative answer for the con cluding chapter.
drift idea [which] appeared to offer a ready explanation of adaptively neu tral changes"; twenty-five years later, it swung back so that "hyperselec tionism became fashionable. All differences between populations and spe cies were
to defend his views on the having evolutionary role of isolation and genetic drift against the attacks of Fisher and Ford, while at the same time appear versus or ing to retract drastically modify those views. In the famous Fisher as selectionist and controversy, the Fisher side is often portrayed Wright
During thisperiod, Wright found himself in the awkwardposition of
85
CHOOSING
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the side as anti-selectionist. During the period up to 1970, Fisher Wright seemed to be winning the battle, while Wright was either losing or claim was not anti-selectionist. At the same time, the perception that ing that he there was a battle seemed to stimulate fruitful research and thus advanced
the sciencewhile confusingthehistory (Provine 1985; Skipper 2000). In any case, by 1950 Wright was no longeropposing the hardening of the
evolutionary Beginning synthesis. with M?xime Lamotte's work on C. nemoralis in the 1950s
did not accept this evidence as support for his own genetic drift, yet Wright in fact, he "admitted that two of the prime examples of random theory; drift he had frequently cited were actually cases of adaptations resulting
(Lamotte 1950, 1959;Millstein 2007b), new evidence seemed to support
to argue that genetic drift primarily from selection."155 But he continued could occur in the laboratory as theoretically predicted and that it could act to the in combination with natural selection in evolution. While objecting
cause for evolution," he stated that "in so premise that there is any "single as far my position could be classified as advocacy of any single principle, this
was selection"(Wright 1955, 16). Later he asserted thathe had principle

never "attributed any evolutionary significance to random drift except as a
accurate. always same At the founder principle did not time, the credibility ofMayr's seem to suffer from the refutations of genetic drift by the Ford group. While Wright and others considered the founder principle a special case of genetic drift, it came to be regarded as a distinct hypothesis, perhaps consensus circa I960.156 to the only exception pro-selectionist strongly selectionist view: lineage is the explanation "Our ultimate goal
not
triggerthatmay release selection"(Wright 1967, 255), althoughProvine of his own early (1986) has shown that writingswas Wrights recollection
the
the evolutionary significance of drift. As Timothy Shanahan suggests, "A number of celebrated cases of nonadaptive evolution by random drift'were later successfully reinterpreted as adaptations formed by natural selection."
in the study of a phyletic terms of its of each morphological change in selective advantage" (Gould 1966, 621). But we have ample evidence from the statements of other leading evolutionists to suggest why they dismissed
Gould did not explainwhy he himself, in a 1966 paper, expressed a
(Shanahan 2004,134).
155.Provine (1986,443); see Wright (1951,455). 156.Provine (1986,407-8,484); Mayr (1963,209-11,529-34); and Mather (1973, 84).
86
18
Harden? Why Did the Synthesis The Views ofOther

Founders and Leaders
in the early 1950s, his theory. Haldane, view that drift assists selection.158 Later ... will matter to a whole
o as far as I know, never de begin with the founding fathers: Fisher, JL viated from his strong selectionist position.157 Wright more or less abandoned his drift theory, while others adopted what they thought was seemed to favor Wright's he asserted that "chance earlier effects
case of very small rarely species," except in the the founder principle may come into play or "the si populations where multaneous establishment of several factors which are harmful singly but
In a review of the then pres (Haldane 1958,17). adaptive in combination" ent status of natural selection for one of the many celebrating publications s the centennial ofDarwin mentioned the Haldane briefly Origin ofSpecies, idea that evolution may result from genetic drift in small tribes. He wrote, "The weakest point in argument is that he has not adequately Wright's considered what happens when this tribe starts hybridizing with others"
(Haldane 1959,141).
Ford, no like his colleague Fisher, always insisted that genetic drift has important role in evolution but went further in his public criticism of
of adaptivepeaks Wright: the theory
iswholly unrealistic and has done much harm in biology; assuming, as it are so stable that a apparently does, that ecological conditions given type of genetic adaptation ... is persistent. (Ford 1957a, 86)
157.Bennett (1983,46); see alsoFisher (1950,19) andFisher and Ford (1947,167-71; 1950).
158. Haldane statement emerged (1953, foreword; 1954,117). The first comment simply endorses Waddington's that genetic drift is one of the "very few qualitatively new ideas" that have (1953,186) from the mathematical theory of evolution.
87
CHOOSING
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He
the theory of genetic driftwas proposed, it seemed reasonable because selective advantages were thought to be small; but we now know, Ford insisted, that are in fact com "large selective advantages mon in natural so selection overwhelms drift.159 The verdict: populations," admitted thatwhen experiments
ex by Ford and his colleagues proved that natural selection plains everything, while drift explains nothing.160 Julian Huxley, an influential advocate of the synthesis, did not show sympathy for genetic drift after 1942. Returning from a two-week
much with
visit to theUnion of Soviet Social Republics in 1945,Huxley reported

s influence, Russian evolutionists were approval that despite Lysenko even more selectionist than those in Britain and theUnited States and had
more kept their pure research going effectively, despite wartime pressure to do work on seasonal Nicolai Dubinins applied work. He mentioned s "first selective changes and control of mutations and Gause experimental proof of the efficacy of organic selection" using ciliates.161 Reviewing Simp sons Tempo and Mode inEvolution, Huxley noted that gaps
in the fossil
more
rapid in small isolated populations, could exist for the millions populations evolution
record might be explained away by Wright s hypothesis that evolution is

but geneticists may doubt if such of years needed for nonadaptive
now clear that selective (Huxley 1945a). Later he wrote, "It is so small as to be undetectable in any one generation, are ca advantages on the scale of all time, of producing pable, when operating geological of biological evolution.... change Evolutionary is almost selection" effected always gradual wholly through (Huxley 1954,3). He asserted that the discovery that human blood groups are maintained a balance of selective in a morphism and advantages by is almost phenomena and disadvantages the observed
as a in evolution, until 1963, simply omitted genetic drift significant factor when he again changed his mind and stated that drift "can have definite evolutionary consequences."162 In an abridged edition of his took notice of the proposed Mayr "non-Darwinian cause Lamarckism, evolution" 1963 book
tion that the ratiosoriginatedby drift (Huxley 1955). In later writings he
(chapter
13) confirms Fisher
and refutesWright
s sugges
seven years later, published revival of genetic drift under the name
(inappropriate and misleading, he asserted, be orthogenesis, and other heresies could also be described
159.Ford (1957a); see also Ford (1971,36-38). 160.Dowdeswell andFord (1953) andFord (1955,1960). 161.Huxley (1945b,255); cf. Kolchinsky (2000a). 162.Huxley (1963, introduction) andOlby (1992, 70).
88
Why Did
Harden? The Views ofOtherFounders and Leaders theSynthesis
new ideas in a by that term). He attempted to refute the couple of pages, as the not that should natural selection concluding they primary replace cause of evolution 1970,126-8). (Mayr that the burden of proof lies on the shoulders of any one who wants to invoke differences: genetic drift to explain nonadaptive "Ifwe cannot see why a difference should be connected with differential ad Stebbins asserted
we are not aptation, justified in concluding from this fact that the difference to is that (Stebbins 1966, 74). Contrary non-adaptive" early assumptions more genetic drift should be relatively important in small populations, no can be example of differentiation between populations is known which ascribed solely or principally to [chance]. One reason is that the effects of selection, both direct and indirect, are very strong, particularly in popula tions which are becoming smaller because of a worsening environment.
no gene remains unaffected a sufficient Consequently, by selection for number of generations so that its frequency can be permanently altered by chance alone. (Stebbins 1966, 76) Many phenomena selection. However, can be a combination of chance and natural explained by "Chance factors by themselves probably have little effect on evolutionary processes, but in combination with reduction in population size and the accompanying natural selection, they may play an important
s later view. But three years later, Stebbins is essentially 77). This Wright s restatedWright that concept of adaptive peaks and valleys by proposing one to rather than the from another mutations, drift, trigger jump peak (Stebbins 1969, 124-5). Presumably he meant small mutations. This was a nice act on the part of Stebbins, since he insisted at the same balancing ex time that the Goldschmidt macromutation theory had been definitely cluded by modern molecular biology and developmental genetics (Stebbins
role inguiding the early stagesof adaptive differentiation" (Stebbins 1966,
1969,104).
Stebbins described
also had a notion of fashions
did not (as faras I know) mention in any of his scientific publications but
in a letter to Edward textbook. According characterized dels
in evolutionary
theory,which
he
thoughtaftertheOrigin ofSpecies: (1) a romanticperiod (1860 to 1903), which the rediscovery of Men (2) an agnostic reaction (1903 to 1935), in
by "extreme enthusiasm for Darwinism and adaptationism";
O. Dodson, who summarized it in his 1952 to Stebbins, there were three periods of evolutionary
end because
laws and the invention of the gene concept seemed to lead to a dead of Johannsens conclusion that natural selection has a limited
89
CHOOSING
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effect in improving a pure line and because of de Vries's unwieldy large mutations; and (3) the modern synthesis, beginning around 1935, inwhich was revived.163 Darwinism write:
harden?David J. Why did the synthesis Depew and Bruce H. Weber
Gould
professes not to know the causes of this "hardening." He mentions the prestige of solid adaptationist explanations in the field (Gould 1983, 88). These seemed to confirm selectionist experiments conducted in the
laboratory,where gene frequencies of different phenotypes could slowly be shifted by controlled environmental variation. We would like to sug gest, however, that standing behind this prestige is the deepening influ
mathematically expressible set of laws to laboratory work, and extending laboratory results to the field,was precisely what would be needed to argue that evolutionary theory now stood on a solid, quantifiable, and potentially axiomatic-deductive basis. (Depew and Weber 1985b, 230)
ence of on the empiricist models of science synthesists, their successors, and their philosophical defenders. A clean series of inferences linking a
163.Dodson (1952, 91-94); see alsoCockrum and McCauley (1965, 631).
90
19
Moth The Peppered

addition to the evidence presented byDobzhansky (1941) and sum
in chapter
In marized
vations
7, and the more recent research on chromosome inversions resistance to antibiotics (chapter 12), blood groups (chapter 13), (chapter 15), many other obser (chapter 14), and snails and tiger moths
decades
moths thathad previously had light-colored wings were being replacedby

moths (of the same species) with dark-colored wings. It seemed plausible that this phenomenon was somehow connected with the darkening of tree bark caused by smoke from factories, but there was considerable disagree (1937,1945) was more viable, but its numbers were the form that dark (melanic) argued were more to be seen and eaten likely ordinarily limited because dark moths about the exact nature of the selective factor. E. B. Ford
as a probable example of even natural selection, before the research byH. B. D. Kettlewell inEngland on the was widely noted in the late peppered moth (Biston betularia).164 It areas nineteenth century that in industrial of England and other countries,
quite well known in the 1960s. Industrial melanism was frequently mentioned
and experiments were cited in books published during the four covered by this survey. Iwill mention here only one, which became
ment
a member of Fords group at Oxford, tested Starting in 1953, Kettlewell, were were eaten the hypothesis that the moths by birds and that they being more to be eaten when the color of their wings contrasted with the likely more than they rested. He marked and released 200 moths, mostly the dark form (carbonaria), in a heavily polluted area near and then tried to recapture them. A significantly higher Birmingham were observed of the being proportion light moths disappeared, and several color of the trees on which taken by robins and sparrows. Niko Tinbergen, a Dutch-British biologist
a more direct treeswere a lighter color. Others suggested by birds when the in polluted air and the physi connection between the chemical compounds that determined wing color. ological mechanism
164.On Kettiewell,see Turner (1990) andRudge (2003,2005,2006).
91
CHOOSING
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who laterreceivedthe 1973 Nobel Prize in Medicine orPhysiology forhis

research on animal behavior, took motion pictures of redstart birds taking and eating the moths. As a control, the experiment was repeated in the un as was much polluted countryside ofDorset. There, expected, the dark form more to escape the birds. Kettlewell to have established not claimed likely was the selective agent, but also that human only that predation by birds were more a or dark of which moths judgments conspicuous against light correlated well with the effectiveness of the birds in catching background the moths.
1956 and follow-up can probably made the United
were published in 1955,with a full report in KettlewelTs firstresults

papers in later years. A 1959 article in ScientificAmeri his work better known to biologists and teachers in States. As one might expect, the 1955 report was immediately
to in their own country. lenged the Americans study industrial melanism the next few years, itwas mentioned in books by Ford's colleagues During and and other British (Dowdeswell by Sheppard) biologists.165 Kettlewell's book on evolution, as far as I experiment did not appear in any American and Adrian M. Srb (both at know, until 1961. In that year, Bruce Wallace it briefly in their book. Cornell University) mentioned (Wallace Adaptation and Srb 1961,40). Since this book appeared in a second edition in 1964, we was was thus may assume it widely read and partly responsible for publiciz was noted in four books on evolution ing Kettlewell's work. The next year it authors: Eldon John Gardner atUtah State University, David by American at at theUniver Merrell the Paul Amos Moody J. University ofMinnesota,
citedby Ford (1955) at a Cold Spring Harbor colloquium, where he chal
sity of Vermont, and Herbert H. Ross at the University of Illinois.166 The earliest reference I have found in a zoology text is by Alfred M. Elliott at the University ofMichigan. He and Charles Ray Jr. at Emory seem to have been the first to mention it in a general biology University text (Elliott and a vague reference to the Ray 1965); there is also changes Beck, both atHarvard.167 After that, it became a familiar piece of evidence for natural selection inmany textbooks, most of which reproduced Kettle pair of photographs showing moths against backgrounds.
in populations of lightand darkmoths byG. G. Simpson and William S.
well's
similar and contrasting
165.Lack (1957);Dowdeswell (1958,1959,1963); Sheppard (1959b); Barnett (1958); Wad Beer de and (1962); dington (1961); Huxley (1963). 166.Gardner (1962); Merrell (1962); Moody (1962); andRoss (1962). 167.Elliott (1963); Elliott andRay (1965); and Simpson andBeck (1965).
92
The Peppered Moth evo s research on the to persuade peppered moth help natural selection as the primary cause of evolution?
Did
Kettlewell to accept
lutionists continued
No. Most had already adopted thatposition,or like Moody
(1953, 1962),
to insist on a significant role for genetic drift in evolution. But even a in many textbooks, through its appearance popular articles, and itmay have reinforced the evolutionary beliefs novel byMargaret Drabble, of the next generation of biology students168?although by that time, the evidence for evolution overwhelming. and for the natural selection hypothesis was already
168.Drabble (2001, title page, 252); Hagen (1999); andRudge (2000b).
93
20
Natural Selection?
The Triumph of
A jl
According
to Ernst
Mayr,
at a meeting
in Princeton
in January 1947,
the origin of diversity.... Not all other biologists were completely con verted. This is evident from the great effortsmade by Fisher, Haldane, as late as the late 1940s and 50s to present and Muller again and again
agreement with the conclusions of the synthesis.All participants endorsed the gradualness of evolution, the pre eminent importance of natural selection, and the populational aspect of
therewas universal and unanimous
as gists such Max Hartman.
evidence in favor of the universality of natural selection, and from some a reasonably agnostic statements on evolution made by few leading biolo (Mayr 1982,568-9) synthesis in the
United States andEngland (table1,p. 131) had decided thatrandomgenetic

drift by itself, while theoretically expected to occur under certain conditions, has essentially no significant role in evolution, except perhaps to facilitate natural selection, so that natural selection acting on small variations is all
By the 1960s, the founders and leaders of the evolutionary
in La Jolla,W. at Oxford.
important. The views of founders and leaders have been discussed in earlier now turn to four other was often cited as chapters. I biologists whose work at the Institute of evidence for natural selection: N. P. Dubinin Cytology at in Francis B. Sumner the Moscow, Scripps Institute of Oceanography C. Allee at the University of Chicago, and David L. Lack
Dubinin, who had earlier proposed an idea similar to genetic Wright's drift, started to publish his research inwestern journals in 1945. In a series of papers with G. G. Tiniakov, he reported that natural selection rapidly in of measurable the genetic structure of populations produces changes
94
Natural Selection? The Triumph of near Moscow. of genetic drift (or Dubi No mention Drosophila funebris nins own "genetic automatic appears in these papers.169 The processes") selective factors in them are seasonal changes in climate and industriali
zation.
(Pero Sumner, whose work on protective coloration of the deer mouse was often cited as evi and the mosquito fish myscus) (Gambusiapatruelis) and also contem dence for natural selection, was originally a Lamarckian
evolution by drift. He was persuaded plated the possibility of nonadaptive own results to accept an by his adaptationist interpretation. Since Lamarck ism had been refuted and "inner perfecting tendencies [orthogenesis] do not belong esis worth in the realm of science," natural selection was the only hypoth s (1940) revival of macromutation considering. Goldschmidt Darwin wrote, it amounted to believing inmiracles unacceptable?as
was
with Mendelian
a result of selection in accordance by predators, in science his for theories dislike laws, despite particulate movement and for the faddish character of theMendelian (Sumner 1945,
the color of the mice was
Sumnerwrote thathe decided (Sumner 1942, 437). In his autobiography,
as an "Because he enjoyed a well-deserved 239-43). reputation exceedingly careful experimentalist, his reinterpretation of the adaptive value of the dif ... races carried much ferences between geographical weight with both sys tematists and geneticists."170
the social life of animals, published was theory, it Lack, an now overshadowed
Allee, who followed Wright's theoryof evolution in his 1938 book on

(with four other authors) amassive a huge amount of evidence trea
tiseon animal ecology in 1949.Although he stillseemed to favor Wright s selection (Allee et al. 1949).
by for natural
on Darwin's finches, is ornithologist known for his research a scientist who, like from of drift to changed Dobzhansky, example on natural selection as a result of his own research.171 In his monograph an view that natural selection finches, Lack rejected Darwins would favor different varieties on different islands, because he could find no evidence that the differences have adaptive significance. Instead, like "most the Galapagos forms which have been described geographical tive evolution as "small and isolated populations" Lack wrote: was
Sewall memoir, Wright effect(Lack 1945,116-7). In an autobiographical
in birds," their nonadap best described by the
169.Dubinin andTiniakov (1945,1946a, 1946b, 1946c, 1947a, 1947b). 170.Provine (1986,230; see also 1979) andGerson (1998, 363). 171.Provine (1986,406-8) andGould (1983, 88).
95
CHOOSING
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were
win's finches are adaptive, and that ecological isolation is essential for the new persistence [sic] of species, only when reconsidering my observations five years after I was in theGalapagos. (Lack 1973,429) Finches, Lack stated that, contrary of adaptive differences
as regarded non-adaptive (hence the importance of Sewall Wright's variation of animals in nature" by G. C. theory of genetic drift)-"The I Robson and O. W. Richards in 1936 fairly reflected current opinion.... reached my conclusion, that subspecific and specific differences inDar
asked me, in themid-1950s, why I postulated that various subspecific differences in the finches are nonadaptive, it isworth stressing that,before my work, almost all subspecific differences in animals Since
... Arthur Cain
In the first edition of his book Darwin's to Robson
and Richards, he believed "the absence must exist. related [between species] is only apparent"; adaptive differences no "There is other way of accounting for the ecological isolation of each was more He selectionist than Hux species" (Lack 1947,134,142). already who in his 1942 book had stated that "ecological isolation is a cause of ley,
species-formation," whereas Lack asserted that speciation is "equally expli cable through competition between species" (Lack 1947,142). When Darwins Finches was
so he could a not let him publisher would change the text, only add preface, was now on inwhich he that the who burden of anyone explained proof claimed that species differences are nonadaptive: This In the interval, views on species completed in 1944.... formation have advanced. In particular, itwas generally believed when I wrote the book that, in animals, nearly all of the differences between textwas
torof theEdward Grey InstituteofField Ornithology atOxford, but the
reprinted in 1961, Lack
had become
direc
a adaptive, change of view which the present book may have helped to it now seems probable that at leastmost of the seem bring about. Hence ingly non-adaptive differences inDarwin's known, prove to be adaptive. (Lack, 1961) In his book Ecological Adaptations, Lack finches... would, more were if
same species, and between closely related species in the subspecies of the same genus, were without years later, it is adaptive significance_Sixteen or almost all, subspecific and specific differences generally believed that all, are
did not mention
drift, but wrote:
who
This book, like every otherwork of natural history, shows, at least for those have eyes to see, the immense power of natural selection and the towhich it can give rise. (Lack 1968, 310) complex and subtle adaptations
96
Natural Selection? The Triumph of influence on one biologist. Alis a historian of science) wrote in later became (who and Richards the doctrine of Robson Ecology that an immediate
Lacks the Journal
1947 book Crombie Animal of
had
tair Cameron
now be are (1936)?that species differences rejected, nonadaptive?could since Lack had shown the differences between bird species to be adaptive
(Crombie 1947).
the research by Lack, and later by Peter and Rosemary Grant, to the evolution, their influ general acceptance of Darwinian was ence came already established. primarily after 1965, when the NSH on this states that "Lacks Edward J. Larsons book subject comprehensive Although contributed cites only one book published before 1965 to support that statement.172
was textbookdogma by the 1960s" but view of theprocess [of speciation]
Results ofa Survey of Biological Publications

Given and leaders of the evolutionary syn thesis in favor of natural selection?denying any significant role for genetic of the founders about the "followers"? ?-That is, what review articles this consensus
drift or macromutations?what on evolution
about other biologists who published books and technical and related topics during the 1940s, 1950s, and 1960s? Here, the picture is not as clear-cut. In the decade 1941-1950,
less than 40 percent of those publications as the (natural selection acting on small mutations) pri accepted the NSH or did not is in factor evolution and indicated that drift mary insignificant
mention it at all (see table 3 [p. 133] fordetails).Almost 30 percent, while
as an important factor, stipulated that drift may accepting natural selection also allow a also have a significant influence on evolution. Some would and/or role for other factors: large mutations (13 (13 percent) orthogenesis more one to because than hundred numbers add up percent percent).These
the categories are not exclusive; for example, Breder (1944) was willing to 13 percent rejected and orthogenesis. About accept both large mutations natural selection entirely.173
172. This Sciences was a high school textbook school drift (American quote texts Institute Sciences, Biological references are in his (2001,198); evolution before 1961 did not mention 1942: 1943: Ashby, et al., of Biological
Curriculum
note 6 (295-6). Most 173. Natural Emerson, White; Natural 1950: Birdsell, Hubbs 1946:
at all (Grabinerand Miller 1974).

selection without (2), Mather, Willier; none; 1947: Lull, Luria;
Study of the high
[1963]). The
is from Larson published
(publications): 1944: Conklin, none; 1942:
1941: Hubbs; 1945:
none;
1948:
Wigan; 1949: Colbert, none;
Burnet,
Clausen
selection with
1949:Allee et al.,Boyd, none; 1946: Lush; 1947:Crombie; 1948: Li,Muller, Schaeffer; Muller;
Boyd, Montagu.
drift: 1941: Dice;
Jepsen; 1950: Summerhoff. 1943: Amadon, 1944-1945: Hazel;
97
CHOOSING
SELECTION
just below 30 percent.174 In the last decade surveyed, 1961-1970, without drift dropped slightly but remained
about 60 percent accepted natural se lection acting on small mutations, without drift, as the primary or only fac tor in evolution. Support for drift (along with natural selection) remained In the next decade (1951-1960), support for natural selection about 50 percent, while sup 35 percent. By the second half of this
port for drift increased to about decade (1966-1970), and orthogenesis had large mutations, Lamarckism, completely disappeared.175 So, the views of these other evolutionary biolo gists in the late 1960s reflect the views held by the founders and
leaders
Large schmidt;
mutations: 1947-1948:
1941: none;
none;
1942: Breder; Lamy;
Lamarckian:
1941-1950:
1949: Davis, none.
1943: Westall; 1950: none.
1944:
none;
1945-1946:
Gold
1941: none; 1942: Breder; 1943: none; 1944: Conklin; Orthogenesis: 1947: Lull; 1948: Holmes; 1949: none; 1950: Summerhoff.
1945: Lillie;
1946: none;
1944: Conklin; Rejects natural selection: 1941: none; 1942: Breder; 1943: McAtee, Willis; none. 1945: Lillie; 1946: Errington; 1947-1950: 174. Natural selection without drift (publications): de Beer, Gregory, 1951: Blum, Clausen, Rice and Andrews, Watson; 1952: Mathews; 1953: none; 1954: Cain, Lerner, Sheppard, White; 1955: none; 1956: Harvey; 1957: Carter; 1958: Clausen and Hiesey, Darlington, Dowdeswell, Kettlewell, Birch, Sheppard; 1959: Anfinsen, Clark, Dowdeswell, Shull; Lerner, Morley, Sheppard (2); 1960: and Stone; Sheppard, Waddington. Natural selection with drift: 1951: Carter, Lamotte; 1960: Van Valen.
1953: M. Smith; 1959: Moody; 1954:Carter; 1955:Li; 1956: none; 1957: Waddington; 1958: J.
Dunn, 1951: none; 1952: Dodson; 1953-1954: Large mutations: 1958-1959: none; 1957: Waddington; none; 1960: Dodson. Lamarckian: 1955-1956: none; 1951: none; none; 1955: Goldschmidt; 1956:
1952: Dodson,
Lindsey, Patterson
Orthogenesis: none. 1958: Brough, de Beer; 1959-1960: natural selection: 1951-1952: Rejects none; 1958: Cannon; 1959-1960: none. 175. Natural 1963: Cain, selection without
1952: Lindsey; 1953: none; 1954: Carter, Russell (published none. 1957: Carter, Waddington; 1958: Cannon; 1959-1960: 1951: Carter, de Beer; 1955-1957: 1952-1953: none; 1954: Carter; none; 1953: none; 1954: none; Heuts;
1962); none;
1954-1957:
drift: 1961: Lack, Wallace 1964: none;
and Srb; 1962: Hardy,
de Beer, Gardner;
Dowdeswell; Darlington, 1966: MacArthur and Connell, Ross, lison, D. L. Clark, Gottlieb, Workman; Natural selection with
1965: Blacklith,
1967: Hamilton, Solbrig, Williams; 1969: Ewens; 1970: none. Moody; 1963: Ehrlich
Levins, MacArthur; 1968: Al Sheppard; 1965: Car
and Kimura, Moody, Large mutations: Lamarckian: 1970: none. Orthogenesis: Reject natural nott; 1964-1966:
M. Smith; 1967:Dunn; 1968: none; 1969: Mettler andGregg, Weller; 1970:Crow lquist;1966:J.
Volpe. 1951-1960: none; 1962: Ross; Waddington; Sinnott; 1963-1970: none. none; 1965: Hardy; 1965
drift: 1962: Merrell,
and Holm;
1951-1960: 1951-1962: selection: none;
none; none;
1961:
1962-1964: 1964-1970: 1969-1970: none.
1963:
1951-1960:
none;
1961: Waddington;
1967: Carter;
1968: Cronquist;
1962: Blackwelder; none.
1963:
Sin
98
Natural Selection? The Triumph of
in the 1940s Another
not cover the latest results, by active researchers and do historians have recently begun to recognize their value in reflecting the of a scientific community (Brush 2005, Bertomeu accepted knowledge It is important to learn what the leading scientists S?nchez et al. 2006). this book have been devoted come established to that question. But ideas do not really be a scientific are in taught knowledge discipline until they to the next not think what the leaders Textbooks may report generation.
to consider here includes textbooks, group of publications written by founders and and articles popular books, (again excluding those textbooks are often dismissed by scientists because they leaders). Although are not written
of drift). synthesis (rejection
after rejecting "other factors" but before the hardening
of the
thinkabout thevalidityof a theory, andmost of thepreceding chaptersof
of textbooks over time reflect,with some time lag, changes in the views of the leaders. Conversely, if an idea never appears in a textbook, it can hardly be said to be established knowledge. is succinctly expressed in a statement by profes sional scientist and amateur historian Stephen Jay Gould: The value of textbooks To learn the unvarnished commitments of an age, one must turn to the textbooks that provide "straight stuff" for introductory students.... Sur veys of textbooks provide our best guide to the central convictions of any era_This field of vernacular expression has been neglected by scholars, though the subject would yield great insight. (Gould 2002,576-7) and articles in the category a more large sample; comprehen reveal significant differences. books
us are since many college today, but they do tell what students learning, and textbooks are written by college professors, they give an indication of what in the content is accepted by academic scientists. Furthermore, the changes
In my survey, I have included popular of textbooks in order to get a reasonably sive study would probably
find that whereas natural
we we first In thedecade 1941-1950, if disregardthequestion of drift,

selection about two-thirds of the evolutionary biologists accepted on small mutations in their books and technical acting
reviews, about half of the textbooks did so (see table 2, p. 132). Fewer than 15 percent of evolutionary biologists supported large mutations, compared decade as the time when evolutionary biologists as a community rejected
to nearly45 percentof authorsof texts (see table 2). I would identify this
on small mutations, while many by natural selection acting to support evolution biologists continued by large mutations. proceeds
de Vries (and Goldschmidt) and accepted the hypothesis that evolution

other
99
CHOOSING
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In the same decade, while driftwas gaining support among evolutionary itwas gaining a foothold in textbooks but was not biologists (see table 3), as as (see table 2).176 quite popular orthogenesis In the next decade, 1951-1960, the proportion of textbooks that en or dorsed natural selection (with without drift) jumped from half to about rose from 13 percent to about three-quarters,177 while the support for drift 20 percent.178 Large mutations received the support of about 20 percent, in less than the previous decade.179 considerably
the textbooks were similar to the 1961-1970, Finally, in the decade books and technical reviews in that they had almost 60 percent support for
176. Natural et al.; 1943: 1946: Shull and Mather, Moment, Natural Hogben, Shull. none;
selection without 1944:
S. F. Cain, Howells;
et al., J.A. Thomson;
Pauli; James, Nelson, Robbins andWeir. selection with Sinnott; 1947: Curtis
1942: Rogers and Goddard, Marsland; 1948: Alexander, Holmes; 1949: Darlington 1947: Emerson; 1950: Baitsell, Bates, Etkin, Fuller, Gibbs, Grove and Newell, 1945: Borradaile, Kenoyer Shull et al.; 1942-1944: none; 1945: Altenberg; none; 1949: Hardin; 1946: 1950: 1942: 1944: 1947: and
drift: 1941: Colin,
Fasten, Gerould
and Poole;
drift: 1941:
and Guthrie,
Daubenmire;
1948:
1941: Fasten, Fuller, Guyer,Moon and Mann, O'Hanlon, Weymouth; Large mutations: Hunter and Hunter, Weatherwax; 1943: Baker and Mills, MacDougall and Hegner, Storer; Grove and Newell, and Weimer; 1945: Lowson; 1946: Haupt, E. G. White; Strausbaugh Emerson, Hunter Mavor, Weatherwax; 1948: Guyer, Potter; 1949: de Laubenfels, Fuller, Hunter (2), Stauffer,Wheat
and Fitzpatrick, Winchester; 1950: Villee. 1941: Fasten; Lamarckism: 1942-1943: and Weimer; none; 1944: Howells, Strausbaugh none. 1945: Lowson; 1946-1947: none; 1948: Guyer; 1949-1950: 1941: Guyer; 1942: Swingle; 1943: MacDougall and Hegner, Storer; 1944: Orthogenesis: Grove and Newell, 1949: Hunter and andWeimer; 1945-1947: none; 1948: Guyer; Strausbaugh 1950: none. selection: none. drift: 1951: Hegner and Stiles; 1952: Elliott, Gardiner, Hardin, et al.; et al.,Transeau 1953: Darlington, Grove and Newell, Kenoyer and 1955: A. 1957: Grove 1956: Mainz, Allison, Brown; Whaley; Morgan; et and Anderson, A. Milne and 1958: al.; Moore, Winchester; Milne, J. Whaley selection without 1941-1942: none; 1943: de Laubenfels; 1944-1947: none; 1948: Rejects 1949-1950: 177. Natural natural
Hunter, Mavor; Carter; Milne
and Milne, Wilson; Guthrie
1954: Breneman, Newell, 1959: Breneman, 178. Natural 1955: Hall Villee; Natural
and Milne. and Stiles, Kettlewell Brouwer, Hegner (article), Mavor, Milne selection with drift: 1951: none; 1952: Rogers et al; 1953: Moore; 1954: none; and Moog; andWells; 1956: Alexander, Wells 1957: Snyder and David, Johnson et al., 1959: Hardin, Weisz, 1955: Sinnott; Zirkle; 1960: none. 1958: Beaver.
1958: Yapp/Borradaile; selection rejected:
179. Large mutations: 1951: Fuller, Stanford, Weimer; and Milne; 1952: Milne 1953: Kenoyer et al.,Transeau et al.; 1954: Fuller 1956: Haupt, Miller and Haub; 1957-1960: Villee; andTippo, none. Lamarckian: 1951-1953: 1958: Yapp/Borradaile. none; 1954: Fuller and Tippo; 1960: Elliott and Ray. 1953: Grove and Newell; 1957: Grove and Newell; Orthogenesis: Natural selection rejected: 1951-1954: none; 1955: Sinnott; 1956-1957: none; 1958: Beaver; none. 1959-1960:
100
Natural Selection? The Triumph of
natural selection without drift180 and about 35 percent for natural selection with drift.181 The major difference is that about 10 percent of the textbooks still endorsed large mutations,182 which had almost disappeared from the reviews. and technical monographs
did not translate
rarily) valid. The
lishedknowledge in the 1960s?is
So, our conclusion?that
the natural selection hypothesis
became
estab
other biologists miliar fact that knowledge presented research frontier by a decade or so. One
consensus of the founders and leaders strong pro-selection into equally strong support for natural selection among a of the fa during the period of this study, consequence in textbooks generally lags behind
just barely (and probably only tempo
the
we have four cases: persuasive.183 Here, lege, changed from the de Vries
at textbooks to be advantage of looking frequently popular enough revised is that one can sometimes see an author changing his or her mind an idea of what reasons were in successive editions and can thereby get
1. James Watt Mavor (1883-1963), professor of biology atUnion Col

large-mutation theory to the NSH (small
mutations) between the second (1941) and third (1947) editions of his
et al., Grove selection without drift: 1961: Bower, Brown, Core and Newell, et al.; 1963: and Lape, Winchester and Lowell; 1962: Beaver, Bonner, Buffaloe, Goodnight Institute of Biological and American Sciences/BSCS, Buddeke, Grogan, Hausman, Braungart et al., StaufFer et al.; 1964: W. B. Crow, Dillon and Dillon, Eisman and Tanzer, Good Kenoyer 1965: Miller and Vance, Nason, Penny andWaern, night and Gray, Guyer and Lane, Marsland; 180. Natural Heiss Romer, Storer and Usinger, Winchester; StaufFer et al., B. Wallace; and Flemister, Keeton, 181. Natural selection with 1966: Avis, Berrill, Cockrum et al., Grove and Newell, Hardin, and Throneberry, 1967: Ashton, Baker and Allen, Buffaloe et al., Platt and Reid, Wilson and Loomis. Nelson drift: 1961: Hickman, Villee; Johnson et al.; 1962: Alexander,
Gardiner
Cockrum Johnson
and Ray, 1966: Beaver and Noland, Berrill, Elliott 1968: 1967: Moment, Villee, Weisz; Winchester; et al., 1969: Johnson and Palmer, Storer et al., Strickberger, Villee Elliott, Lerner, Nason Weisz; et al. et al. et al., Parse 1970: Jessop, Nelson (general), Johnson (zoology), Keeton, Orians, Weisz; et al., Winchester. gian and MacCauley, Dillon, Weisz; et al., Mavor and Manner, Weisz, Mavor et al.; 1964: Winchester; 1966: Avis, 182. Large mutations: 1961-1962: none; 1963: Villee and Manner, Stauffer et al.; 1967: Villee; 1968: Taylor andWeber, Villee et al; 1969: Kroe ber et al., Nason and Goldstein; 1970: none. Lamarckian: Orthogenesis: 183. According 1961-1968: 1961-1965: to "Planck's none; none; 1969: Orians; 1966: Mavor 1970: none. and Manner; 1967-1970: none.
1963:Elliott, Villee et al., Weisz, Yapp/Borradaile;1964:Dillon andDillon, Whaley et al.; 1965:
scientists (especially older, well-established ones) do Principle," a new idea; to not accept they just retire and die, while the younger genera change their beliefs tion, which has learned the new idea, takes over. This principle was first tested experimentally by as David Hull, Peter D. Tessner, and Arthur M. Diamond, using Darwins theory of evolution the new idea (Hull et al. 1978). We have already seen that in the mid-twentieth century, several leaders did change their views about the importance of genetic drift in evolution.
101
CHOOSING
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General
making this revision (Mavor 1947, 864, 880). 2. AlfredM. Elliott (1905-?), professor of zoology at the Universityof
Michigan, was
Biology
text but to remove his endorsement forgot
of de Vries
in
Dunkers zoology text (1952, 1957), but in latereditions,he discussed the an as Glass and his of drift (studiedby (Elliott example genetic colleagues)
1963,1968). thogenesis, line evolution, almost as if the horse wanted'to a shy, seclusive forest dweller the same time, he gradually abandoned his support for or to be a case of exemplified by the horse, which "seems straight At
completely
selectionist
in the first and second editions of his
In later editions, as it not as seem (Elliott 1968). The successive editions straight-line might state inconsistent show signs of revision, with contradictory piecemeal ments on different pages. 3. William Beaver versity, presented
change itsway of life from to a swift running prairie animal" (1952,658). influenced by Simpson, he stated that the evolution was
his general biology textbook (1958),which he dropped in the sixthedition

the "modern synthesis" (628). The seventh (1962), where he also mentions of theUniversity ofDayton, gives edition, coauthored by George B. Noland
Uni (1896-?), professor of biology at Wittenberg a view in of natural the fifth edition of selection negative
a fuller treatment of evolution, are stating that the major factors (presum in order of importance) mutation, genetic drift, natural selection, and ably 1966,506). No examples of natural selection migration (Beaver and Noland are author presented one view in a popular book but then presented a few years later. Garrett Hardin in a textbook published at the (1915-2003), professor University of California, Santa Barbara, dis cases in cussed several which genetic drift is important in Nature andMans (Hardin 1966), he presented tion of drift.184 the theory of natural selection with no men given. 4. One
a different view
Fate (Hardin 1959). But in the second edition of his textbookon biology
184. This 1952). Hardin
is in effect the same as his position in the 1952 edition of the textbook (Hardin is best known for his influential paper, "The (1968). Tragedy of the Commons"
102
21
Is Evolutionary Theory Scientific?

Those to revive the biologists who attempted theory of natural selec tion in the 1930s were acutely aware that the theory had acquired a its earlier supporters were too eager to explain ev
bad reputation because
that4 extremely few attempts have ever been Raymond Pearl complained ... to test the or even made principle of natural selection experimentally, even though it observationally, by precise and critical specific observations," ever enunci most important abstract biological principle "perhaps the ated. Tons of books and papers have been written about it" (Pearl 1930, that evolution 176). A. F. Shull, in agreement with Pearl, recommended ists, in their renewed efforts to revive the theory of natural selection, should is
like thefairytales ofRudyard Kipling (1902). erything by a "just so story"
colors, its mimicry and its forget about "its sexual selection, itswarning a more (Shull 1936,212). signal colors" in favor of experimental approach in in that the D. M. S. the Watson, passage quoted chapter 5, complained is so flexible, it can explain anything. But two decades after Pearls NSH critique, substantial progress had been made in satisfying his requirements for "a proof that natural selection has altered a race" (Pearl 1930, 175), as
and biologists began to debate the question, is evo Species, philosophers itmake lutionary theory scientific? By this the philosophers meant, does testable predictions? The biologists had to decide whether to answer that had already done in the quotation at the beginning question (as Haldane or to argue that the criterion should not of this monograph) philosophers' at the especially relevant beginning of the twenty-first century: creationists it is supposedly not testable, and their attacks attack Darwinism because to nonscientists who have learned a naive version in high school
shownby the detailed survey ofAllee et al. (1949, 641-55). Startingaround the time of the 1959 centennialofDarwins Origin of
be applied to evolutionary theory(or tobiology ingeneral).The question is
may seem plausible the scientific method Cooper 2001) may
of
try to defend
science courses. Biology teachers (e.g., evolution by arguing that reconstructing
103
CHOOSING
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past events, when no human was present to observe them, is a legitimate astronomers do it all the time), without in science method (geologists and realizing that evolutionists have in fact made testable (and confirmed) pre In fact, one of the most prominent creationists mentions Darwin's insects as an example of an untestable tautologi hypothesis about wingless cal proposition he is apparently unaware of the experi (Johnson 1993,21); dictions. described Even
ment
in chapter 7. before 1959, therewere two important experiments that confirmed predictions of the Fisher-Haldane-Wright theory under controlled condi tions, where both natural ence the evolution of a
and Spassky reminded readers that the Dobzhansky-Pavlovsky experiment showed that large populations have smaller variations in genotype frequen cies, as predicted, and reported another experiment confirming the predic tion that "if the sizes of the foundation of the outcomes
chapter 3) and ofDobzhansky and Pavlovsky (chapter 17). Dobzhansky
selection and genetic drift were expected to influ the research of Kerr andWright (see population:
constant, the variations tion stocks came from a source with higher genetic variability than from a more uniform source" In and Spassky 1962,149). genetically (Dobzhansky varia the meantime, LaMotte had shown that the (1959) expected greater tion of smaller populations was also present in natural populations. Fishers an associa theory of balanced polymorphisms. He suggested that to certain diseases would be tion between blood groups and susceptibility was et al. (1953,1954) and others found in humans.This confirmed byAird in the 1950s
are stocks of the populations kept should be greater if these founda
was made byFord (1940a, 1945), based on A different kind ofprediction
were aware of this evolutionary biologists (chapter 13). Many s and by Dobzhansky success, yet like the experiments by Kerr andWright in the discussions with philosophers. group, itwas rarelymentioned
Popper
of the Darwinian laws?in
versus Darwin
stated that "the recent vogue of histori for the evils of fascism and communism, "might be as regarded merely part of the vogue of evolutionism" (Popper 1945, 69). It was therefore necessary to discuss first the use of evolutionism in the bio no as a scientific such thing logical sciences. He asserted that there could be the social sciences. He cism," which he blamed law of evolution, because
an attack was theory of evolution. His article primarily on "historicism"?the assumption that history is determined by inescapable
In 1945, thephilosopherKarl Popper published thefirst of severalcritiques
104
Is Evolutionary Theory Scientific? can the evolution of life on earth ... is a unique historical process_We or to find a natural law not hope to test a universal acceptable hypothesis we are confined to the observation of one to science if Nor process. unique can the observation of one us to foresee future devel unique process help
opments.(Popper1945,70)
Yet at the same time, he admitted, "I see in modern Darwinism the most successful explanation of the relevant facts" (Popper 1945, 69). s and then as a critique, published first in the journal Econ?mica Popper not attract attention from evolu did much The book, Poverty ofHistoricism,
translation of his Logik der Forschung had appeared, con time, an English a detailed on scientific method and on the taining exposition of his views use of as a criterion to falsifiability distinguish real science from pseudosci 1959). Explanation one must be enough: willing unknown facts or events. not (Popper The American ence of the facts, no matter how successful, was to make testable predictions of previously
was printed in 1957,with a second edition in 1960. By that tionists until it
Scriven then published an article in philosopher Michael Science inwhich he argued, against Popper (and also against R. B. Braith as as well C. G. and P. that evolutionary the waite, Hempel Oppenheim), indeed offer "satisfactory explanation of the past ... even when Scrivens article prediction of the future is impossible" (Scriven 1959,477). elicited reactions from biologists and became part of the debate about the ory does
scientific legitimacy of evolutionary theory in the 1960s. I see two serious problems with Scrivens argument. First, he defines a category of subjects, including "a great part of biology, psychology, anthro
and quantum phys pology, history, cosmogony, engineering, economics, errors are in which he calls these ics," "irregular subjects"; subjects, "serious known to arise in the application of the available regularities to individual cases," and he implies that "prediction is excluded" in these subjects (Scriven 1959,477). Aside from the derogatory connotation of the word "irregular," most cer his conception of prediction is impoverished. Quantum physics can be tested in the some tainly does make predictions that laboratory, and Scriven incredible accuracy. Apparently has in mind the fact that the present position or quantum state of an in dividual subatomic particle is indeterminate until it ismeasured, hence its future position or quantum state is unpredictable without a measurement, and the and presumed fact that cosmogony deals only with unobservable events in the distant past. (He had hence unknowable argued in 1954 that the age of the universe of them have been confirmed with
we will never be able to determine whether
is finite
105
CHOOSING
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or infinite.) Moreover, can or no science, strictly speaking, irregular, regular offer an accurate "prediction of the future," except in the sense that it can more or less a measurement we make in the predict accurately the result of and extrapolations about fu present under controlled conditions. Guesses so tureweather, forth may be useful in planning our lives earthquakes, and but become increasingly unreliable after a few days or weeks. Yet meteorol ogy and seismology are respectable sciences. to the mathematical Thanks theories of Fisher, Haldane, and Wright itwas pos and the availability of fast-breeding organisms like Drosophila, sible by the 1940s to test the predictions of evolutionary theory in precisely the same way that one could test the predictions cosmogonical theories185?though seemed unaware of this fact.
of quantum mechanics and even some and biologists, philosophers,
had already clearly distinguished between the historical Dobzhansky to evolution approach (phylogeny) and the causal-mechanistic approach the second approach
does use the hypothetico-deductive method; Popper and other philosophers who asserted that evolutionary theory is not falsifi able simply ignored the entire corpus of work forwhich Dobzhansky and his colleagues had become famous by 1959.187 and others have attempted "to en Second, Scriven asserts that Darwin capsulate the principles of evolution in the form of universal laws and base
As was subsequently (populationgenetics).186 pointed out byAyala (1977),
on them" but have failed (Scriven 1959, 477). On the contrary, predictions while I do not claim that biologists have established universal laws of evo lution, I have shown that they have used the principles of natural selection to generate several that have been tested and confirmed (or predictions refuted, as in one case).188 A theory that governs the future behavior of a system, like Newtonian or quantum mechanics or can also general relativity, yield predictions about its present behavior.189 This is a crucial fact often ignored by those who
say that evolutionary theory cannot make testable predictions. It is ironic that Scriven lumps Darwinism with cosmology in the category of irregular can not six years of the publication theories that explain but predict.Within 185.Berry (1982,57); see alsoHull (1974) and Williams (1981).
Dobzhansky andAyala ofbeing too Popperian (Lloyd 1988,10)!

188. Other from Fisher's
see also Prout (1995). in 186. Dobzhansky (1951, quotation chapter 6); 187. Ironically, a prominent member of a later generation of philosophers were discussed early testable predictions are discussed (1986). theory by Leigh by Lewis (1986);
of biology
accused
some
later predictions
189.Ruse (1969) andProut (1995).
106
Is Evolutionary Theory Scientific?
to the of Scrivensarticle,thepredictionfrom thebig bang theory(contrary

state radiation steady theory) that space is presently filled with microwave was at a few degrees above absolute temperature spectacularly confirmed; this confirmation played some role in the decision of proponents of steady state to switch to was not big bang, although that decision greatly influ
related example, closer to the domain of biological evolution, is the stars. to in of the of elements In order origin theory explain the synthesis from hydrogen of elements heavier than helium, Fred Hoyle predicted a nuclear reaction, the tri-alpha reaction (fusion of previously unobserved A three helium nuclei confirmed to form a carbon-12 nucleus), which was subsequently in a laboratory experiment. Even though the entire process of the evolution of elements from hydrogen takes too long to follow in the can be laboratory, almost every individual stage explained theoretically and
enced by knowledge that the radiationhad been predicted (Brush 1993a).
Although many scientific theories do make successful predictions, that success is not a necessarily requirement for accepting those theories.190 It does raise the barrier for anyone who wants to replace a successfully tested an as yet untested one. theory by
testedexperimentally (Brush 1996b, chapter2.3).
How
The
Evolutionists
Replied
to and Scriven Popper
were too to concede Scrivens biologists quick charge that evolutionary not make testable it does I remarkable that Ernst find theory predictions. a confirmed Mayr, who had himself made prediction from evolutionary was theory (as he
with Scriven: 1963,207]), agreed

The
pleased
to remind
the readers of his next book
[Mayr
has emphasized quite correctly that one of themost important contribu tions to philosophy made by the evolutionary theory is that ithas demon strated the independence of explanation and prediction.
can describe and theory of natural selection explain phenomena with considerable precision, but it cannot make reliable predictions_Scriven
But, he continued, we are happy if "our causal explanations [also] have pre dictive value" (Mayr 1961, 1504). Apparently what Mayr meant was that can be scientific as as it evolutionary theory long provides good explana even it does it if doesn't make testable tions, predictions, but sometimes
190. See note 5.
107
CHOOSING
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make
logical prediction (defined a or scientific law") theory to from the future). The lat temporal prediction (inference from the present ter (what in call advance" call "novel and physicists "prediction philosophers is "much more in the biological sciences" (Mayr prediction") rarely possible
testable predictions. Later he distinguished as "conformance of individual observations with
1982, 57-59). But three years later, he argued that "in principle there is no sciences with respect to difference between the physical and the biological experiment and observation": neither makes "absolute" predictions (Mayr he agreed with Scriven that "the ability to predict 1985, 50). Nevertheless, a is not a requirement for the validity of biological theory" and stated: "The can describe and explain phenomena with con theory of natural selection siderable precision, but it cannot make reliable predictions, except through
or not make tionary biology geology does sciences do, but argued that physical
statements as, for instance, 'the fitter individu such trivial and meaningless als will on the average leave more offspring'" (Mayr 1988,19, 31). a historical science like evolu Simpson also agreed with Scriven that predictions in the same way
the
or the testing of hypothetical generalizations proposed explanations a some historical record of has the aspects of predictive testing_ against ... A conspicuous example has been the theory of orthogenesis, which maintains that once an evolutionary trend begins it is inherently forced to continue_That fossil record. As that record_A plainly has consequences thatwould be reflected in the a matter of observation, the theory is inconsistent with or event crucial historical fact may be deduced from a
or ac theory and search may subsequently produce evidence for against its occurrence. is tual prior That has been called "prediction"... what actually occurrence not is the antecedent but the predicted subsequent discovery. (Simpson 1964,144-5,147) A few years later he wrote, "Claims that the hypothetico deductive method one allowable in science are almost absurdly (e.g., Popper 1959) is the only an allowable method.... Retrodictive extreme, but obviously it is interpre tation and explanation are almost unique to the historical sciences."191 The most elaborate refutation of the philosophical critics in the period Ghiselin accepted Popper theory satisfies it: s
Method.
before 1970 was Michael Ghiselins book, The Triumph of the Darwinian
falsifiability criterion and insisted that
Darwin's
191. Simpson (1970, 85, 90); see alsoAyala (1968b) andLewontin (1972).
108
Is Evolutionary Theory Scientific? It seems almost unreal that, among all theories of science, theDarwinian von Bertallanfy] theory of natural selection has been singled out [e.g., by as is if it of hence refutation_A scientific, refutable, theory incapable is possible to given even one conceivable state of affairs incompatible with its truth. Such conditions were specified byDarwin himself,who observed
that the existence of an organ in one species, solely "for" the benefit of another species, would be totally destructive of his theory.That such an never been found is a most natu adaptation has compelling argument for ral selection. (Ghiselin 1969,63)
But Ghiselin
were
did not discuss the more recent predictions by Dobzhansky, and Ford, Mayr, Wright. several evolutionists believed they had already satisfied the Although demand for testable predictions and could do so in the future,192 others not satisfied that their theories were sufficiently predictive.193 Rud
wick (1964) argued thatorthogenesiscould be tested but thatthe synthetic

re not (at least in the domain of theory could paleontology). Waddington s criterion, a cant "in be that arguing jected Popper hypothesis practice," to deal with the ob disproved because it "can always be suitably amended was in biology as in it raised" (1969, 110). Conversely, jections possible, other sciences, for a theory to make a confirmed prediction and yet still be
wrong.194
disagreed among themselves. E. Manser agreed with Pop Philosophers not is therefore fails "to explain that and per evolutionary theory predictive in the normal scientific sense" (1965, 31). Lehman asserted, con Hugh can be a form as to fit the tra Scriven, that "Darwins in such put theory using covering-law model [Hempel] rather than the deductive-nomological "We also try to show that Darwins the 'statistical-probabilistic pattern continued by saying, pattern. He can be used to make predic principles
in this and the previous note to three and (Lewontin, biologists Simpson, was also an "un on this issue in different took different There Waddington) positions places. dercurrent of perplexed and subtle dissatisfaction with the concept" of natural selection, perhaps Note the references who to be resolved by using the semantic "theory of signs" of C. W. Morris (Mason and Langenheim
192. Bonner (1962); Dowdeswell (1963); Simpson (1964, 136); Simpson and Beck (1965); Wallace (1966); Baker andAllen (1967, 503); Carlson (1967, 305); Lewontin (1967); Shep Moorhead andKaplan [1967,98]);Dobzhansky (1968); Slobodkin pard (1967); Waddington (in and Smith (1969). (1968); 193.Lewontin (1965,1968b, 1974); Medawar (in Moorhead andKaplan [1967,"Remarks by theChairman ]); Simpson (1964,147);Waddington (in Moorhead andKaplan [1967,13, 98]).
1961,158). 194.Dunn andLandauer (1934,239) andLaudan (1981).
109
CHOOSING
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tions,"but he does not give any examples of facts not already known (1966, wrote that not 14-16). Dudley Shapere prediction is impossible in biology, just difficult because of the number of interacting factors, not because of the explanation' "essentially historical character of biological (Shapere 1969, can make testable pre 14). A. G. N. Flew (1966) asserted that Darwinism dictions, but the examples he gave were not impressive and not supported to one kind of antibiotic, by specific evidence: if bacilli develop resistance thenwe may confidently expect that new antibiotics will eventually produce similar immunity.195 Thomas Anderson theory does make genetic drift predicts in particular, the theory of random that a population of only one hundred animals will extinct, while a population of one hundred thousand sub of one hundred each will prob subpopulations relatively rapid and progressive evolution" (Goudge seemed Goudge testable predictions; to be satisfied that natural selection
probably become divided into one thousand ably enjoy "sustained,
1961,126).
Michael Ruse areas that have pointed introduced to the return of to "those light-colored moths smokeless zones" as a fact that was "clearly ex is also evidence for natural selection, views on Darwinism (Ruse 1977).
insects the case ofwingless 1969,339); he noted. Ruse later criticized Popper's in the first of a Mary B.Williams, an axiomatization of Darwinian posed tion for a truly predictive
pected" (and thus could have been predicted)by the synthetic theory(Ruse
series of papers on this subject, pro theory,which "provides the founda (Wil
to biology is qualitatively different from physics with respect predictions. But with the exception of Ruse's brief remarks about wingless insects, not discuss the confirmed did of natural selection philosophers predictions mentioned in this monograph, have been able to examine.196 at least in the literature of the 1960s
liams 1970, 370). David Hull (1974) cast doubt on thebasic premise that
(and therefore testable) theory of evolution"
that I
criticism of In my opinion, the most satisfactory response to Popper's was the book Darwinism who does not by philosopher Stephen Toulmin, a at asserts mention it all. Toulmin that it is futile to seek simple criterion such as predictive success to determine whether a theory or activity is "sci entific."
195. See comments 196. References by Barker (1969). for philosophical discussions
fromsourcessuchas the Index and thebooks by Hull (2001);Hull andRuse (1998); Philosophers and Sober For further discussion of the testability of evolu (1984a, 1984b, 1993). Lloyd (1988); see Williams and Zachos (2002). (1973,1982,1985) tionary theory,
of predictiveness
in
biology
can be retrieved
110
Is Evolutionary Theory Scientific? critical questions which a philosopher brings to science must be co ordinated with the factual studies of history. We must be prepared to scru
The
tinize in detail a representative selection of classic theories, analysing criti came to be cally themerits forwhich they accepted, and the standards by which they established their claims to superiority. (Toulmin 1961,16) ... is ... an Forecasting application of science, rather than the kernel of a science itself... novel and successful theorymay lead to no increase in a our forecasting skill;while, alternatively, successful forecasting technique may remain for centuries without any scientific basis. (Toulmin 1961,36) cannot "foretell the of Thus, Darwins coming-into-existence theory creatures of a novel Yet small Darwinism did forecast "short-term, species." scale events": When Australians itwas used myxomatosis [a viral disease] to control the rabbit forecast on the best Darwinian principles that a new
population, strain of rabbits would
cases re involving melanism inmoths, the happened in other small-scale on. actions of infectivemicro-organisms to anti-biotics, and so (Toulmin
become dominant, whose constitutions were more correctness of this prediction has helped resistant to the disease_The to confirm the merits of the Darwinian theory.And the same thing has
1961,25)
Mathematicians
and
Challenge Natural
Dissatisfaction with
Physicists Selection
the supposedly nonpredictive nature of evolutionary use to learn theory and the belief that biologists might be able something ful from physicists and mathematicians led some of them to organize a on "Mathematical to the Neo-Darwinian symposium Challenges tation of Evolution, "held at the Wistar Institute of Anatomy Interpre and Biology
in Philadelphia, April 25-26, 1966.197 As it turnedout, the symposium

197. Martin sown in Geneva was in its coeditor of the proceedings, wrote Kaplan, preface that "the seed in the summer of 1965 ... when of the Wistar [director [Hilary] Koprowski ... on math confronted by a rather weird discussion between four mathematicians and Kaplan the Darwinian 1967). concerning theory of evolution" (Moorhead is a chairman of the symposium, said "the immediate cause of this conference sense of dissatisfaction about... the so-called neo-Darwinian [t]heory." Apart
Institute] was ematical Peter Medawar,
doubts
pretty widespread
Ill
CHOOSING
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was not devoted had
primarily
physicists apparently andWright Haldane, (as Ernst Mayr pointed out in one of the discussions) but instead tried, rather clumsily, to reinvent the theoretical wheel. What was more
never
to the and topic of its title.The mathematicians looked at the canonical works of Fisher,
at Los Alamos and coinventor of the hydrogen bomb; Ulam, mathematician at William and Marcel Bossert, computer expert Harvard; Sch?tzenberger, at the of mathematics of Paris. the more active professor University Among was V. F. a well-known in the nuclear discussion participants Weisskopf, at theMassachusetts Institute ofTechnology physicist nal instigators of the meeting. The biologists at the symposium?especially Mayr and one of the origi
was what Ernst called "Evolu interesting (at least to me) Mayr as to the Mathematical of Evolution," tionary Challenges Interpretation^] those interpretations were presented byMurray Eden, professor of electri cal engineering at theMassachusetts Institute of Technology; Stanislaw
andWaddington? over were that the models the mathematicians argued proposed by grossly out such essential factors as mutation rates and simplified, leaving popula tion size. Ulam's model, for example, led to an intractable equation that did a very not to sexually reproducing organisms, but by making apply rough was able to arrive at he the conclusion that in approximation, qualitatively cluding
reproduction speeds up evolution enormously?" Ulam replied, "Obviously a common sense tells us [this] immediately; the whole point of pursuing mathematical model is to get some quantitative estimates." Waddington and was Niels Barricelli then remarked that this what Fisher, geneticist exactly Haldane, more accurately and with simpler earlier. other biologists repeated and years mathematics?thirty-five Mayr that they had already arrived at the same conclusions without knowing any mathematics and without using (as Ulam did) a model that failed to make andWright had done?much sense (Moorhead and Kaplan 1967,21-33). biological I suspect the net result of the symposium was to convince that they already had all the math
from religious "philosophical
"How [does] this [differ] from the standardbiological theorythat sexual
sexual reproduction would
speed up evolution. Waddington
asked,
they needed
the biologists from Fisher, Haldane, and
Popper?that or an too much. It is too difficult to could not be envisage imagine evolutionary episode which In addition, "there are objections made by fellow explained by the formulae of neo-Darwinism." scientists who very widely feel that, in the current theory, something among biologists generally" (Medawar is missing.... 1967, p. xi). These objections held
the such as "the kind of pious bunk written by Teilhard de Chardin," objections, ... have been and methodological very well voiced by Professor Karl objections the current neo-Darwinian defect of explaining [tjheory has the methodological
... are
112
Is Evolutionary Theory Scientific? to feel inferior because
and they did not need Wright, the equations themselves.
they couldn't derive
The outcome of Popper's critique of evolutionary theory,mentioned by several of the mathematicians and physicists at the symposium, was less fortunate. Popper apparently never learned about the successful predictions had actually made; he repeated his claim that Darwinian not scientific, and even theory is though he eventually retracted it (Popper the had done: another weapon had been handed to the been 1978), damage creationists in their battle against evolution.198 that evolutionists
a law (Act 590) state time the Arkansas demanding legislature passed equal The law included the and evolution, although itwas later ruled unconstitutional. or cannot be logically falsified" (chapter phrase, "evolution experimentally observed, fully verified, 7); the last word clearly indicates that the creationist writers of the law had exploited Popper's 198. In 1981 for creationism Assembly used a similar phrase.
bill proposed in (butnot passed by) the Maryland General critique(La Follette 1983).A similar
113
22
Context and Conclusions
In
the first part of this chapter, I discuss some speculations about pos influences on evolutionary theory; then sible cultural and philosophical I return to a survey of the scientific reasons for accepting the natural selec hypothesis. has called attention towhat he calls the "metaphysi Jonathan Harwood cal foundations of the evolutionary synthesis," by which he means attitudes and other concepts that may toward holism, reductionism, materialism, tion
have
war
or the formation and reception of the syn discouraged encouraged thesis at specific times and places. He suggests that the ideology of Bildung inter (cultivation) and the calls for holistic anti-reductionist synthesis in "might have prompted Dobzhansky, Mayr, Germany a to in tackling the problems synthetic framework adopt to have broad "romantic cultural as well
by this traditiontend theory"(Harwood 1994, 5). Intellectuals influenced

as scientific interests and are attracted to the also in biology," as exemplified by Goethe. Harwood some the suggests that there may be similarity between these influences and as he described in an earlier work ones that acted on German geneticists, (Harwood 1993), and on physicists in the first part of the twentieth century, tradition that Julian Huxley, to the philosophical a was proponent of the "synthesis" label, sympathetic and Arthur Eddington, idealism ofA. N. Whitehead,
et al. Simpson, of evolutionary
as discussed by Paul Forman (1971). Similarly, Colin Divall (1992) argues
James Jeans and to the "emergent evolution" concept of C. Lloyd Morgan, as a reaction Har against (Herbert) Spencerian mechanistic materialism. to characterize these metaphysical wood used the phrase "styles of thought" assumptions, and that phrase has become ence in the last two decades.199 popular among historians
of sci
in Context was devoted to "Style in Science" (Das see Gerson and Bowler (1998) biology, evolutionary the usefulness of the concept of "epistemic has also endorsed (2004). Richard Lewontin style," as Maienschein 1991). A related term is (Lewontin 1995, 96; Maienschein presented by Jane 199. An ton and Otte entire issue of the journal 1991). For applications Science to
114
Context
and Conclusions
to to support a uni synthesize evidence from different fields of biology fied evolutionary theory based primarily on natural selection. When they invok concluded that most of this evidence could be explained without was an were almost ing drift, that large mutations always fatal while there to drive evolution, that Lamarckism was ample supply of small mutations not reliable evidence, and that orthogenesis was not a respect supported by able scientific concept, they ended up with a hardened synthesis that could be considered reductionist and materialist. As Provine pointed out, itwas not really a was synthesis but actually a constriction. But such a theory was
here. In the 1930s, an evolutionary "synthesis" is misleading and that selection with Lamarckism, drift, mutations, theory synthesized was clear by orthogenesis might have suited the spirit of the time. But it the 1950s that the leaders of the neo-Darwinian synthesis really wanted The word
War II with the styleof thoughtpopular in post-World quite compatible

science and culture: not romantic but realist, sometimes known as "midcen It was the age of physics based on elementary particles, turymodernism." orbitals, geology based on plate quantum chemistry based on molecular on and analogies between brains molecular based tectonics, DNA, biology
and computers.200
the present account, it seems most pertinent that the idea of ran domness (the basis of the genetic drift concept) was strongly associated with a holistic, anti-reductionist, anti-mechanist metaphysics, which arose For in countries after Weimar World War I, especially in the German-speaking a to to It cultural Paul Forman was, according (1971), response Republic. criticism of scientists and mathematicians who claimed to know everything. Itwas also associated with the radical anti-realist "Copenhagen interpreta
German biology in the 1930s.Garland Allen (1983) portraysI.Michael

Manchuria Lerner, who was born in in theUnited States, as an example and worked in Canada
same movement, which may tion" of quantum mechanics (Brush 1980).The of be regarded as a revival of the romantic and neo-romantic movements the nineteenth century (Brush 1978), has been shown by Anne Harrington (1996) and other historians of science to have had a strong influence on before settling of the influence of "holistic material there is no reason to think it views had no discern
on Sewall had any influence United States; in fact, Wright s Wright in the
biographer argues that Wright s own philosophical
ism" in the form of "genetic homeostasis."But
introduced by Gerald Holton (1988) to identify concepts like "force" or "atom" or "evo whose observation and mathematical lution," go beyond meanings analysis. On the relation be see Holton tween and and Brush themata, (2001, epistemic styles chapter 33). "thema,"
200. Brush (1980).
115
CHOOSING
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and Fisher
(1934) recognized a possible connection between indetermin ism at the atomic level and indeterminism in evolution. But I would argue between the holistic anti-reductionist in biology
on his scientific ible effect work (Provine 1986, 96-97). Wright did not invoke Muller (1929,498) Heisenberg's indeterminacy principle,although
that one should look for connections
as and the eager acceptance of genetic drift?not part viewpoint a as of himself proposed, but rather catalyst for natural selection, Wright as an for nonadaptive (and somewhat vacuous) all-purpose explanation
characters and species differences. At the same time, one should allow the new are also driven in the by possibility that oscillations popularity of drift
as Similarly, Simpson pointed out, the idealist philosophy of German to support a saltationist (macromutation) view biologists often led them of evolution. But he seems surprised that the same biologists (especially could also support an empirical approach: Rensch) To me ... it came as a really unpleasant shock to find Rensch avowing himself an idealist after a long discussion of evolution that could be en dorsed, word forword, by the most rigid materialist ... [but] almost all of us [Americans?] share with Rensch evolution. (Simpson 1949a, 184) a basically inductive approach
empirical findings (Futuyma1979,272-6).
to
it is not at all surprising to find empiricism com From my perspective, in late bined with idealism in science, since that is just what happened nor to is it surprising find that em neo-romanticism; nineteenth-century
can facilitate a transition from romanticism to realism (in this piricism to the case, the transformation of evolutionary theory from the 1930s in the establishment of the mod 1950s), since that is just what happened ern energy concept earlier in the nineteenth century (Brush 1978). Indeed, in some ways the unification of biology by the evolutionary synthesis is to similar the unification of physics by the law of conservation and trans formation of energy. Other
classification schemes may be useful here. For ex philosophical to John Greene, "The champions of the modern synthe ample, according a sis ... the reductionists between polemical position midway occupied and the vitalist-finalists." its existence.201 Fisher s theory was are shuffled randomly They used the language of purpose but denied
of course during
on the that genes assumption a the reproduction, but in way that makes also based
201. Greene (1986,220); see alsoGreene (1999).
116
Context
and Conclusions
more as in the kinetic predictable. Just theory of gases, for population genetics, there are always so many particles that fluctuations are relatively small, and most properties of the system appear to be determined a few parameters. The kinetic a by theory is legacy of outcome much his model century and was revived in the twentieth Jeans) century, just asMendel's early realist-atomist theory of heredity was revived. One might guess that the as reflected the metaphysical evolutionary theories of Fisher and Haldane the realist-atomist smentor (by Fisher period of the mid-nineteenth
I) and therefore sumptions of this "neorealist period" (beforeWorld War were not very most with the 1940s until and 1950s, when popular biologists to the mechanistic-reductionist biology returned approach, perhaps best
Watson-Crick double helixmodel ofDNA.202 exemplified by the

As Richard Lewontin wrote in 1968,
II have seen a vindication in biology of ... an as away of analytic doing science ... the belief that into down their compo technique by breaking systems nent parts, by or using simpler organisms, one can learn simplifying them about more complex systems-It is not the case thatmolecular biology the twenty years since World War our faith in the Cartesian method and analytic while population biology is holistic. Population ... biology is properly analytic and operates by the process of simplifica and tion, analysis, resynthesis. (Lewontin 1968a, 1-2) But of course no movement?holism or reductionism, or romanticism support at any time; there is always an op is Cartesian
realism?ever position
gains unanimous party to be denounced:
complex that they cannot be treated except by holistic statements. The influence of these people has held up progress in population biology for many years and, in addition, has tended to degrade population biology as a science. They are the stamp collectors of biology who, because they them selves are unable to analyze the complex problems of ecology and evolu tion, try to convince the rest of us that nothing but "objective description" of nature is possible. (Lewontin 1968a, 2)
It is unhappily true that there are population biologists who reject the analytic method and insist that the problems of ecology and evolution are so
based wave
See Depew and Weber idea of a "genetic code" (1985b, 228-9 [and elsewhere]). The on the structure of a was in part molecule the book What isLife? by Erwin large inspired by a realist the who advocated (1944), Schr?dinger physicist (unsuccessfully) interpretation of his 202. equation.
117
CHOOSING
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And
War II, according to publicationsby thepsychologist Donald T. Campbell

in that period.
have been a popular
if they wait long enough, the opponents can catch the next swing of the pendulum back in their direction. seems to The pendulum may swing faster in other fields. "Selection" thema in the behavioral and social sciences after World
at a 1961 conference on social Speaking change and evolu was a revival of interest in this tionary theory, he noted that there currently in after decades scientists which social topic, rejected evolutionary theory because of its association with reactionary political views such as social to any scien newly discovered tific evidence for evolutionary theories but simply to the tendency of social scientists to reject the status quo, whatever itmay be; bored with the anti evolution stance of the previous generation, they now accepted evolution The ism.203Other as a model for the modification revival was not credited
Darwinism.
psychologists used natural selection error not mentioned learning (and, by Campbell, behavior by "operant conditioning"). was According associated to Hamilton with Cravens (1988), extreme hereditarianism
for trial and of
the de Vries mutation
theory in the nature versus nurture
controversy of the early twentieth century; its replacement by natural selec a shift tion in the 1930s and 1940s was by, and supported, accompanied toward the recognition of environmental influences in the behavioral and social sciences. Fisher s tions are seldom perfect. used the selection analogy to describe the history of sci Philosophers ence and technology. Stephen Toulmin argued that evolutionary biology provides the best analogy for the development of scientific ideas: assessing the merit of a theory depends on "seeing in how many ways a novel scien tific idea may ... be better adapted than its predecessors or rivals" (Toulmin advocacy of eugenics reminds us that such correla
1961,17; see alsoToulmin 1966).
until 1972, described his method of "conjectures and refutations" as a kind of natural selection. A hypothesis that survives tests and criticism better than its competitors As forDarwin's
delivered in 1961 but not published per 1972). In fact, Popper, in a lecture
as a nonfalsifiable the Karl Popper, despite his criticism of Darwinism for the growth of knowledge ory, considered it a plausible mechanism (Pop
tific teaching"(Popper 1972,261).
"may, temporarily, be accepted
as
part of current scien
theory of biological evolution, Popper made
an interest
203. Campbell (1960, 1965, 1970); see the essaysbyDavid Hull and others in Heyes and Hull (2001) and Popper (1972).
118
Context
and Conclusions
in existence and pur ofdesign by explaining, purely physical terms, the a connection with the fact, well known to in world'."This the suggests pose theoretical physicists since the seventeenth century, that a large number of can be described in two ways that are natural phenomena mathematically different: variational but prin quite teleological equivalent philosophically causal equations. As the great Swiss mathematician ciples and mechanistic Euler wrote in 1744, "All effects of the universe
ing statement (probably not original but worth repeating): "Darwins theory to to reduce of natural selection showed that it is in teleology principle possible causation
Leonhard minima Two
method ofmaxima and by the equallyhappily fromfinal causes [teleology]

and from the effective causes themselves" later, Princeton hundred (Truesdell 1960,200-1). Edwin G. Conklin invoked
can be explained
years biologist the authority of Einstein, Bohr, and Planck to support his view that both and natural selection and purpose are factors of evolution: "Mechanism teleology The are complementary views of nature, neither excluding the other"
(Conklin 1944,132).
earliest and principle in one medium
to point B in another inwhich its speed is different, itwill seems follow the path that makes the total time as short as possible. This to to achieve a future is goal. Yet it imply that the path of the ray designed correct answer (Snell's law of refraction), the gives previously derived by Descartes from a mechanistic from a mechanistic make particle model and later derived by Huygens wave model. Descartes had to assume that light moves
was Pierre de Fermat s simplest variational principle in optics, which asserted that if a ray of light travels from point A
could faster in a denser medium, which is hard to believe, whereas Huygens the more reasonable opposite assumption. So, Huygens succeeded to in a in reducing Fermat's teleological explanation physical causation; one to is advocate of mathematical could that Darwin sense, say Paley (the as is to Fermat. Of course no sensible physicist intelligent design) Huygens an believes that the of ray light is constructed by intelligent designer today no reasonable to get from A to B as fast as and biologist today possible, believes that organisms have been constructed by an intelligent designer to have the s, shows how properties they do. Darwin's theory, like Huygens the appearance of design can be simulated by a causal mechanistic process
(cf.Sober 1996).
Reasons for Accepting
Natural the
Selection Hypothesis
of metaphysics; let us now turn to table 5 (p. 135), where I have Enough tabulated the most popular reasons for accepting the natural selection hy
119
CHOOSING
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in is, the empirical evidence most often mentioned pothesis?that Anglo American books and technical reviews on evolution by biologists (now in in the cluding "Founders and Leaders") period 1951-1970. Notice one wants thatwhile Kettlewell s
and afterward, came too late to have much effect in chang same is true for views of evolutionists (Brouwer 1958, 1960). The the ing Cain and Sheppard s work on snails, the connection between sickle cell the late 1950s and malaria, and the connection between blood groups and diseases. In stead, we have two rather different kinds of evidence playing an important role in addition to protective coloration: on one hand, the development
before Kettlewell s research. The recent controversy about the swork is therefore to the scientific of Kettlewell validity largely irrelevant more status of natural selection in the 1960s.205 rigorous Similarly, the new, in research on mimicry by Jane Van Zandt Brower and others, published
to understand most why biologists had accepted the primacy of natural selection by 1960, one has to startwith the more traditional illustra tions: various kinds of protective coloration, including industrial melanism, as observed
itdid not play a very significant role in thesebooks until after1955.204 So if
peppered moth
is the most popular
overall,
of insecticide-resistant
and oth 1950s);206 and on the other hand, the experiments of Dobzhansky ers on inversions in and Adalia cyclic variations of chromosome Drosophila acts much more show Both that natural selection punctata. quickly than had previously been believed?a few years in the first case, a few weeks or months in the second. The data for textbooks, popular corresponding articles, and books are shown in table 4 (p. 134). The most striking differ
pests (going back to the scale insects in the cit rus groves of California in the beginning of the twentieth century and the similar development of resistance toDDT and antibiotics in the 1940s and
ence is the greater importance given to protective coloration and wingless insects, and the much smaller weight for chromosome inversions, snails, sickle cell/malaria, and blood groups. the 1970s,
itwas not enough to show that natural selection was than other evolutionary processes; the goal posts had been important itself. W. H. Dowdeswell moved, thanks in part to the success of the NSH ac was so far very little that there complained "precise evidence for the tion of selective agents" (Dowdeswell 1971, 369). Fifteen years later, John By more Endler compiled a list of more than one hundred empirical demonstra
204. See only one
chapter 1955)
19; of the seven references specifically mentioned
to industrial melanism work.
in the period
1951-1955,
205.Hooper (2002) andRudge (2000b,2003,2005). 206. Simon (2003 [and andCreager (2007). works cited therein])
(Ford
Kettlewell's
120
Context
and
Conclusions
tions of natural the 1960s
selection
in the wild;
or later and thus had answer s
of theNSH
(Endler 1986 126-53). Even themore recentones did not,

the important quantitative were questions about
in most of them were published little influence on the earlier acceptance
he argued, generally selection. Some of Dobzhansky
to test genetic designed drift, and contrary to his expectations, they showed that the effect of drift was to that of natural selection; as a prominent leader negligible compared to the natural selection hy of the evolutionary synthesis, his conversion early experiments pothesis undoubtedly influenced many other biologists. The drift in combination with experiments
in
the 1950s byKerr and Wright and byDobzhansky and Pavlovsky showed
that the effects of genetic natural selection, pre
But there was no designed laboratory experiments with small populations. credible evidence that genetic drift by itself played a significant role in evo
could be detectedby specially dicted by theFisher-Haldane-Wright theory,
nature. lutionary processes in The fact that old evidence, going back to the period before 1930, was so in support of the new version of natural selection theory frequently invoked leads us to ask: did the new evidence really play a major role in persuad two answers to that ing evolutionists to accept natural selection? I suggest labors of Fisher, Hal First, it seems clear to me that the mathematical dane, andWright were essential to the success of the new natural selection
question.
One could support thisview by quotingDobzhansky, Huxley, Simpson,

and many others.207 But perhaps he is notorious for his attempts in the 1950s to minimize the best authority is Ernst Mayr, because (along with those of C. H. Waddington) relative the contributions of the mathematicians,
hypothesis
and
indeed
to the success of the evolutionary
synthesis
itself.
and (unfairly)Wright.208 Mayr was "beanbag genetics" of Fisher, Haldane, the senior author of a zoology textbook inwhich one finds, near the begin ning, the flat statement: Fisher's (1930) demonstration that even a very small selective advantage of a new gene or gene combination would cause in due time a genetic transformation of populations was a tremendously important contribu tion. (Mayr et al. 1953,12)
to those of the field and laboratory biologists, and forhis ridicule of the
207. Dobzhansky (1955,14); Huxley (1942,123); Provine (1978); Sheppard (1954, 201-2); Watson (1951,192). Simpson (1949b,266); and 208.Mayr (1959) andProvine (1986,480-4).
121
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At
the end of our period, Mayr The modern mathematical
summed up the situation as follows: is a
attitude toward natural selection has two roots. One
root is the overwhelming mass responsible for these advantages.The other ofmaterial gathered by naturalists on the effect of the environment. This evidence was given a largely Lamarckian interpretation in the days when small mutations were claimed to be saltational and cataclysmic. When was realized that all variation had mutations were discovered, and when it a source of a powerful ultimately mutational origin, this evidence became documentation for the selectionist viewpoint. (Mayr 1970,108) of of protective coloration, mimicry, and development So, the phenomena resistance to pesticides and antibiotics became evidence for natural selec
and analysis (R. A. Fisher, J.B. S. Haldane, Sewall Wright, even minute ad selective others), demonstrating conclusively that very vantages eventually lead to an accumulation in the population of the genes
tion, at least in part because of the theoretical work of Fisher, Haldane, and the development Fifteen years later, industrial melanism andWright. were still considered the best empirical evidence for natural of resistance selection
a leader of the in this controversy, wrote pro-evolutionists Eugenie C. Scott, that "the evidence for the operation of natural selection is so overwhelm Earth as
course (Moore 1984,515-6). to present in an introductory over on the an This evidence has also had ongoing controversy impact in American the teaching of evolution and creationism public schools.
ing thatboth IDCs
to account for these introduction). When phenomena, whose pressed to medicine and agriculture is obvious, creationists like Henry importance Morris make an arbitrary distinction between microevolution (which unde 2004, niably does occur) and macroevolution second reason is mentioned The
Creationists] pesticide resistance
now
Design Creationists] andYECs [Intelligent
accept that it is responsible for such phenomena in insects or antibiotic resistance in bacteria" (Scott
[Young
(which, they assert, does not).209 inMayr's 1970 summation quoted mutations Muller's pro discovery of radiation-induced previously: H. J. vided an independent confirmation of Fisher s geometrical argument that can contribute to evolution. Large mutations only very small mutations or are occur to but may sterility.Those biologists who likely produce death for other reasons were now forced and orthogenesis rejected Lamarckism to and were leftwith the reject also the de Vries version of neo-Darwinism
209. Morris for Creation and many articles inActs & Facts, the monthly newsletter of the Institute See Numbers (2004) for the acceptance of microevolution by creationists.
(1974)
Research.
122
Context
and Conclusions
Fisher-Haldane-Wright
alternative.
version
of natural
selection
as the only plausible
s research on D. an Third, Dobzhansky important pseudoobscura forged link between laboratory genetics and field observations, allowing evolution to become an in which experimental discipline hypotheses about natural selection and drift could be tested. The selection Lakatos success of the natural to helps explain the is effect." Imre be called the "Lakatos what might hypothesis, was one of the first of science to point out that sci philosophers fourth answer, which
entists do not simply weigh the evidence on both sides of an issue; they also compare the track records of competing theories (or what Lakatos called "research programmes"?series of theories). If theory A is found to
forA, they are likely to switch their allegiance to A, even ifmost A will eventually evidence the of stillfavors B, because they anticipate that win out, and this can become a self-fulfilling prophecy. In fact, during the 1950s and 1960s, several supporters of natural selection (theory A) claimed evidence that this was the situation: since many characters previously thought
are more "degenerating." If scientists "progressive," while theory B becomes recent research has transformed evidence for B into that all of the persuaded
becomes more B, it thereby ously thoughtto be betterexplainedby theory
successfully predict
or
give
a better explanation
of a phenomenon
previ
to
be nonadaptive and therefore determinedby genetic drift (theory B) had

been shown by more detailed research to be adaptive, while there were no should be that examples of the opposite; therefore, the default assumption all characters are adaptive (or genetically linked to other characters that are in the absence of definite proof that they are nonadaptive. To the adaptive) extent that evolutionists accepted this claim, the natural selection hypoth esis was considered
was consid progressive, while the genetic drift theory a ered degenerating?even though there might still be large number of cases of no selective basis had characters for which apparently nonadaptive yet been found.210 Similarly, H. Muller J. we argued thatwhen see phenom
210. Beatty (1987b, 57); Cain (1951a; 1951b; 1964: 37); Darlington (1963, 61-62); Dob zhansky (1962,280; 1970, 262); Dunn (1959, 103); Ford (1964, 35; 1971, 39-41); Lack (1961, preface); Mayr (1970,120-4); Sheppard (1958,123; 1967,110); and Stebbins (1959a, 50; 1966,
Another version of this effect is described by John R. G.Turner: "Repeated too strong to allow signifi [by the Oxford School] of rather strong selective effects, cant drift, or left an onus of proof producing clear adaptive differences between local populations, on the supporters of Wright's theory" (1987, 342). 74-76; 1970,184-5). observations are still to define drift and whether Philosophers debating about how as a separate force or factor in evolution: natural from selection tinguished it can be clearly dis Sober (1984); Mill
stein(2002,2005); Walsh et al. (2002); Sober (1984a); Stephens (2004); Brandon (2005); Pfeifer (2005); andPlutynski(2007b).
123
CHOOSING
SELECTION
makes it thepreferred defaulthypothesis (Muller 1949,440).

Of course the Lakatos evidence the side whose ismore
ena that look we should not orthogenetic, postulate "mysterious processes" we have not like orthogenesis because just yet explained them by natural selection; the success of natural selection in explaining other phenomena effect can help either side; itgives an advantage to recent. For in the late nineteenth example,
century debate about the atomic structure ofmatter, the atomists had shown around 1870 that evidence from several different phenomena led to similar ber. Despite estimates for the size of an atom and forAvogadro this, the anti-atomist thermodynamics s or Loschmidt's num seemed
to be program more progressive at the end of the nineteenth century, at least by Lakatosian standards (Clark 1976). Then, thanks to the development of new theories von Smoluchowski, of Brownian movement by Albert Einstein andMarion was new to estimates Perrin size able obtain of the of the atom and of Jean s number, which on the Avogadro agreed fairlywell with the old estimates; basis of this agreement and other experiments, he was able to establish once and for all the atomic nature of matter.211 The new evidence was not really much better than the old; itwas just newer. The alternatives to the natural selection hypothesis included, tion to genetic drift, Lamarckism and orthogenesis. Throughout in addi
the pe in riod 1930-1970, Lamarckism was almost universally dismissed (except as unproven and France and in German implausible. The paleontology) was taken was the advocate who only seriously by biologists psychologist William peated whose McDougall, and finally discredited were re on the training of rats experiments a at the in a two decades-long study by group et al. 1954). had been popular (Agar Orthogenesis
Melbourne University of in the 1930s but was strongly attacked by leaders of the evolutionary syn or vitalist thesis as a vestige of unscientific metaphysical thinking; Simpson
not all, that their observations could be persuaded some, but paleontologists better explained by natural selection.212 are essential for spe A fourth alternative, the view that macromutations a to attract small minority of biologists. The only major ciation, continued
win
as "the Bald was theory sometimes discussed "organic selection," also known or it I have not discussed it in the text because effect," "canalization," "genetic assimilation." on this in to is it be textbooks. The literature but rather appears rarely appeared theory confusing, an that seem at first glance to be La attempt to explain by natural selection certain adaptations 212. Another (Waddington characteristic can its own evolution by the organism that guide in the 1950s on the of the "veinless" 1961). Waddingtons experiments development to were in subjected high temperatures highly regarded (Huxley 1963; Drosophila 1962; Moody 1962), but his attempt to explain the results by genetic assimilation was in nature or to involve some behavior
211. Brush (1968,1976) andNye (1972).
marckian
Merrell
124
Context
and Conclusions
s of this view was Richard Goldschmidt, and like genetic Wright a as a macromutations rather than he drift, supplement replace regarded was ment for natural selection. Like drift and orthogenesis, macromutation an idea to the holistic German 1930s. As Gold of the biology congenial advocate schmidt himself the 1940s recognized, "All theories of evolution
tend to reflect the
trendsof the time" (Goldschmidt 1940, 397), and the trendof scientific
favored the natural selection hypothesis. But Gold so his to be to be too important a theory had biologist ignored, given the courtesy of serious consideration?followed by definitive rejec were not in tion.Macromutations evolution because: (1) they are important and 1950s schmidt was that speciation worked that way; (3) there was no reason to make a because every qualitative distinction between micro- and macromutation, evidence step in between had been observed and seemed to follow the same rules as
was no solid empirical always lethal (Fisher 1930a;Muller 1938); (2) there
of thedouble helixmodel of molecular biology followingthe establishment

DNA was produce As previously noted, at least three predictions based on natural selection were confirmed between 1930 and 1970: Darwins prediction that flying on a organisms windy island would tion that chromosome Drosophila evolve a wingless inversions would incompatible new species. with the existence of macromutations that could
be micromutations; (4) there was no need to postulate macromutations, cause micromutations could account for all evolutionary processes, as soon as one in recognized the tremendous power of selection; and (5) research
(or if you prefer, Dobzhansky s opposite result); and Ford prediction associated with
s form;Mayr predic to be prove adaptive s refutation of his own expectation of the that certain blood groups would be school, evolution
of a prediction is "satisfying" referred not to natural selection but to s chromosome Dobzhansky prediction that certain previously unobserved as are intermediate would be discovered because needed arrangements they mation
did they mention the confirmations of evolutionary theory's predictions about the effects of genetic drift by Kerr andWright, by Dobzhansky and Pavlovsky, and by Lamotte. Bruce Wallace seems to have been the statement that the confir only exception, but his
disease. Aside from Mayr and the Ford ists gave these successful predictions no extra credit for novelty and did not even mention evo them in the debate with philosophers about whether lutionary theory is falsifiable. Nor
controversial. The andWilliams
thirdeditionof his textbook(1970). See also Schmalhausen (1949); Richards (1987); B. Hall Weber andDepew (2003). (2001); and
use of the was (1963), (1953b), Mayr concept strongly criticized by Simpson was to withdraw his support in the (1966); Moody persuaded by these criticisms
125
CHOOSING
SELECTION
stages in the evolution from one known arrangement to another. Moreover, this statement about the value of a prediction was made in a popular work, not one intended for other biologists. If one wanted to make the case that evolutionists were converted natural
to the
selection hypothesis by following the scientific method, the best on to outcome have based would be the of the second argument prediction. a in make he did did, effect, compare itwith ob Dobzhansky prediction;
servations, and then he tested the conclusion by controlling the variables in a selective laboratory experiment, which indeed showed that the proposed did the result.213 agent (seasonal temperature changes) produce predicted Dobzhansky himself stated that the result of this research led
Moreover,
him to adopt the natural selectionhypothesis (which he had previously

in this connection by several and the research was mentioned doubted), authors of books on evolution. On the other hand, this was originally a to the natural test of the "contraprediction"?a prediction that, contrary
events itwas correlated with in gene changes frequencies. This sequence of seems consistent with s idea that the result should ideally Popper predicted be one thatwould nolbt expected in the absence of the hypothesis. Yet only a on evolution even mentioned this test (table minority of authors of books 2); and as far as I know, none of them asserted that the result was persuasive because itwas a successful novel prediction. Moreover, when philosophers and a few biologists criticized evolutionary theory for not being testable, no evolutionist seems to have brought forward this example in response.214 I can conclude only that while biologists do make testable predictions and sometimes confirm them, they do not (despite the rhetoric of textbook on scientific method) in their introductory really believe chapters
selection hypothesis, genetic drift does have a significant effect on the evo lution of gene frequencies. When the contraprediction was refuted by field a he then invented observations, possible selective agent and showed that
writers
was not a crucial test and that 213. In retrospect one could argue that strictly speaking, this the laboratory experiment was not really a 1995), but it replication of the field results (Lewontin did seem to have an impact on the views of himself and others. Dobzhansky 214. In a recent issues in biology, Marjorie Grene and "episodic history" of philosophical it is not, David Depew the question, "Is evolutionary If (2004, 249) mention theory predictive? how because can it be state that explanatory?" They Popper called natural selection the new Darwinism; he misunderstood the chance "Overstressing small step [leading to an adaptation] is the result of a purely nonfalsifiable, element, accidental partly he thought " mutation
that each to mention
with
any of the confirmed predictions of evolutionary theory, and the reader may be left not a law-based the impression that the view ofMayr science and Lewontin?that biology is and does not need to be predictive?is and 2004,265-6; (Grene Mayr generally accepted Depew
(Popper 1972,269-70 [as quoted byGrene andDepew 2004, 262]). But Grene andDepew fail
1985,1988; Lewontin 1991).
126
Context
and Conclusions
is the most important criterion for judging a new theory. the other hand, they expect that an established theory should be able to make predictions and that if one of those predictions is falsified, one need that predictiveness On necessarily abandon ments to accommodate physical scientists. The were more not the theory itself but perhaps make minor adjust the data. In this respect, they are very much like difference is that, at least during this period, their than quantitative.
predictions
qualitative
127
ACKNOWLEDGMENTS
Will This book is greatly indebted to the pathbreakingpublications of

Provine
of the linger high schoolbiology circa 1948was incredibly boring,a result of the Scopes Trial (Grabiner andMiller 1974), so I avoided ing effects
taking any biology in
and to his extensive critique of an earlier draft; I regret that I have as as he has. I got a late start: not been able tomaster the subject thoroughly
s can to the truth of Dobzhansky testify college.2151 (1973) maxim, "Nothing in biology makes sense except in the light of evo lution." Lindley Darden also provided a very helpful and detailed critique, to her stan not to quite been able bring my exposition up although I have dards of accuracy and clarity. I thank Francisco Sandra Ayala,
John Beatty, Matt Chew, James Crow, George Richard Richard Lewontin, David Herbert, Garratty, Highton, O'Brochta, Anya Plutynski, David Rudge, Ezra Shahn, V. Betty Smocovi and Nick Zimmermann tis,Carol Sokolski, Roger Thomas, Polly Winsor, information in for answering my questions and providing much valuable correspondence. tiques. My Two anonymous referees provided extremely useful cri
a supported in part by fellowship from the John Simon Foundation and by the Institute for Physical Sci Guggenheim Memorial ence and at the Technology University ofMaryland. research was
were not as successful in texts. A banning evolution from college biology a of loan from textbook obtained (Mavor 1948) copy Liberty Baptist Col popular by interlibrary a sticker in has the title that reads: page lege Lynchburg, Virginia, preceding 215. Creationists to the reader: Use of this volume as a text for reference in is Liberty Baptist College from the standpoint of morals, philosophy, theology, cannot contents the It is necessary to use books whose college of Liberty Baptist College life iswell known. on the fundamentals of the faith
not an endorsement
of its contents
or scientific hypotheses. wholly endorse. The position and the separated Christian Mavor, Vries' refute. a
at Union selection (de College, professor of biology accepted evolution by natural not and four of evidence for which the sticker does it, version) gave attempt to pieces
129
TABLES
TABLE
Founders
and leaders of the evolutionary synthesis
Theodosius Dobzhansky (1900-1975):MNAS, FMRS; zoology;Columbia

University
Ronald AylmerFisher (1890-1962): FRS, FANAS; genetics,statistics,

eugenics; University College Edmund Briscoe Ford London, Cambridge University (1901-1988): Oxford anatomy; University Sanderson Haldane London, FRS; genetics, zoology, comparative
John Burdon
University College
(1892-1964): FRS; genetics and biometry; Indian Statistical Institute (Calcutta)
JulianSorell Huxley (1887-1975): FRS; King sCollege London, Zoological

Society of London
Ernst Harvard University, Mayr (1904-2005):MNAS, FMRS; ornithology;

American Museum ofNatural History (New York)
George Gaylord Simpson (1902-1984):MNAS, FMRS; paleontology;

Harvard University, American Museum of Natural History (New York)
G. Ledyard StebbinsJr. (1906-2000):MNAS;

of California
genetics, University botany;
Sewall (Green) Wright (1889-1988):MNAS, FMRS; Universityof

Chicago, U.S. Department ofAgriculture
Note: Limited to thosewith significantEnglish-language publications, 1930-1950. FANAS = Foreign Associate of theNational Academy of Sciences (United States); FMRS = Foreign Member of theRoyal Society of London; FRS = Fellow of the =Member of theNational Royal Society of London; MNAS Academy of Sciences (United States), elected before 1964.
131
Tables
TABLE
Factors in evolution1.Textbooks,
2
popular articles, and books
19411945 mutations NS, small mutations NS, small RejectsNS 11 2
19461950 17 3 9 7
19511955 14
19561960 16
19611965 27 13
1966 1970 33 26
Total 118 60
(drift is insignificant in evolution or not mentioned) is also significant in some cases)
(but drift, including founder principle,
11110
0 5 10 110 1
etc.)
16 15 NS, includinglarge mutations (deVries, Goldschmidt) NS + Lamarckian

effects (including Baldwin
3 1
2 56
4 6
1 1 15 41 12 41
effect, canalization,
NS + orthogenesis
Number ofpublications 29
28
29
42
60
229
Note: Limited to publications inEnglish by authorswho are either on the faculty of colleges or universities orwho are professional staffatmuseums or other research institutions; not including founders and leaders, listed in table 1.NS = natural selection.
132
Tables
TABLE
Factors
in evolution: Books and technical review articles on evolution and related topics 194119461951 195619611966
1945 mutations NS, small mutations NS, small RejectsNS NS, includinglarge
mutations
1950 6 11 58
1955 11 8
1960 1217
1965
1970 12
Total
12 3 5 13 12 3 3 33 0
70 42
(drift is insignificant in evolution or not mentioned)
(but drift, including founder principle,
is also significant in some cases)
2 2 2 1 0 0
14 11 11 12 137
(de Vries, Goldschmidt)
NS + Lamarckian NS + orthogenesis
effects (including Baldwin
effect, canalization,
etc.
3 33 22 22
2 24
1 23
0 20
Number of publications 26
Note: Limited to publications inEnglish by authorswho are either on the faculty of colleges or universities orwho are professional staffatmuseums or other research institutions;not including founders and leaders, listed in table 1.NS = natural selection.
133
Tables
TABLE
Evidencefor in biology textbooks, popular 19411946-
4
mentioned frequently articles, and books 195619611966
natural selection: Most
1951-
1945
Pepperedmoth
(Kettlewell) Protective
1950
1955
1960
1965
13
1970
29
Total 52
1117
2 15 6 peppered moth, snails)
and other types of industrial melanism 31
coloration
(not including industrial melanism, Resistance to insecticides 0
23 2
9 6 10 9 5
20 14 13 13 13
Mimicry Darwin's finches(Lack)

Resistance Chromosome to antibiotics inversions
0 0
0 0
0 00 0
00 2 3 2
12
5 3 2
and other seasonal insects Wingless on ocean islands Number
cycles inDrosophila, Adalia punctata, etc.
3
30
3
43
10
25 32
14
of publications
42
60
232
Note: Authors limited as in tables 2 and 3, but now including founders and leaders. Publications that rejected natural selection are not included in number of publications at bottom.
Frequency = mentioned ten or more times.
134
Tables
TABLE
Evidence for natural selection: Most frequently mentioned in books and technical reviews on evolution
19411945 moth Peppered

(Kettlewell)
19461950 3
0
19511955
19561960
19611965
14
1966 1970 13 13
Total 51 32 33 25
2 712
and other types of industrial melanism
Blood groups 0
and disease polymorphism
Chromosome inversions Snails (Cain and Sheppard) Sickle cell andmalaria Mimicry
1 0 0 0
3 0 0 2 3 2
2 115
11 3 3 3 7 7
4
etc.
7 8 4 8 5 5 10 4 4 2 4
and other seasonal cycles inDrosophila, Adaliapunctata,
6 8 5 3 3
snails)
25 22 23 22 13 172
Resistance to insecticides 1 coloration Protective

Resistance to antibiotics
3
0
(not including industrial melanism,
peppered moth,
Number ofpublications 29
24
28
38
26
27
Note: Limited as in tables 2 and 3, but now including founders and leaders. Publications that rejected natural selection are not included in number of publications at bottom.
Frequency = mentioned ten or more times.
135
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INDEX
Abrahamson, accommodation,
S., 31n68 4n5
Adalia punctata, 120

Adams, M.B.,8nl5,25
Avogadro s number, 124 Ayala, F.J.,Hn21,35,35n79,38n87, 108nl91,129 Babcock, E. B., 31n67,34n77

bacteria, 74-75
adaptation,vii, 1,25n53,47
animals, 44-45
polymorphism, 63
Agar,W.E.,124 age of universe, Aird, 105
21,25, 32,57, nonadaptive differences, 116 plants, 43
Baker, J.J.W., 109nl92 Balbiani, ?-G., 37n84 Baldwin effect, 6,124n212

Barash, D. P., 34n77 Barker, A. D.,110nl95
Barnes, E.W.
of Ford's prediction, Barnett,
I., confirmation
(Bishop of Birmingham), 30-31,31n66

S. A., 92nl65
69, 84,104 Alaudidae, 53nll0 Allchin, D., 4n5

Allee,W.C.,34n77,103
Barricelli,N., 112
Bateson,W., Bauer, H., 95 6nl2 151
Wright's theoryovershadowed by
evidence for selection,
Allen,EJ.,6,6nl3 Allen, G. E., 5n9,30n65,31n65,31n69, 109nl92,115 Allen's rule,44 Allison, A.C.,70, 73nl42 Alverdes, 37n84 American Indians, blood groups, 72 Amish communities, 80nl52 Anderson, E., 8, 60nll5
anemia,
bean bag genetics, 17,17n32 Beatty,J., 10,Hn21,70,70nl33,72nl40, 123n210,129

reason for selectionism, Beaver, W., 102 Dobzhansky 66-67 s change to
Beck,W.S.,58, 92,109nl92 behavioristic psychology and natural

selection, 118
H., 30n63, 87nl57 Bennett, J.

Benson, Bentall, S.B.,53nllO H. H., confirmation of Ford's
groupA, 69
antibiotics, resistance 25 to, 74-75,120 antireductionism,
pernicious,
correlated
with
blood
prediction, 69 RJ.,106nl85 Berry, Beurlin, 58

Beurton, size of atom, P., In2
Ariew, A., 2n3, 9nl6, Hn21

Arkansas, atomic equal structure time law, 113 of matter,
124
big bang theory,107 bildung (cultivation),114 billion, 15n28
171
Index
Biological Sciences Curriculum Study, 97nl72 biology and biologists, 1,4n5 based on DNA,115 philosophy of, 3,23-24 predictiveness,110,110nl96
resistance to mathematics, 91-92 4 9nl6, 31,51
Carlson, E. A., 109nl92

Carpenter, H. Carson, Carter, 77 Cavalli-Sforza, Ceccarelli, Cepaea Cepaea L., L. L., 37 72,72nnl40-141 G. G. D. H., L.,27 S., loses debate with A. J. Cain, 33
unification of, 1
Bis ton betularia, Blackmun, H. A.,
hortensis, 76 nemoraiis, 76-78
blood groups, 22,54,55,68-73

Blossom, P. M.,53nll0 Bonner,J.T.,109nl92 Bossert,W.,112
chemistrybased on molecular [quantum] orbitals, 115

S. S., 9
Chetverikov,
Chew, M.,
Chironomus, chromosomes,
129
37n84 25,33
botany, 1,44, 60-61 Bowler, P.J., lnl, 6nl2,12,30n65, 114
31n66,
R, 10nl7,14n26,16-17,18n35, Box, J. 31n66,69nl30

brain-computer Braithwaite, analogy, R. B., 105 115
Chiricahua (CH), 64 chromosome theoryof heredity,3 cyclic (seasonal) changes in gene giant chromosomes in salivaryglands of flies,36,37n84,63
inversions, 62-67,72, 36 82 speciation, arrangements, 65
Brandon, R., 24n48,123n210

Breder,C.M.Jr.,97
Bridges,C, 3 broccoli/radish,35n80
Br?mer, Brousseau, BrouwerJ. Brownian Brues,A. Brush, R., 8nl5 G. E. Jr., 120 124 84nl54
cinquefoil,43
citricola scale, 42
V. Z.,
Clark, P., 124 Clark, R.W., onHaldane, 28n59 on Huxley, 47nl00 Clarke, B.C.,68nl27 Clarke, C. A., 70nl34,73nl42, Clausen, J.,43,43n95,61nll8
Cockrum, Coloration, E. L., 90nl63 and warning, 45, protective
movement, M.,72 S. G.,
77-78, 84
2n4,3,3n5,4,17n31,17n33,
26n55,37n84,40n90,107,115, 115nl99,116,124 BuckwalterJ. A., 69nl32 Buican, D., 8nl5,40n91

Burian, R. M.. on macromutations, Dobzhansky 66nl23 and
103,120,122 Conklin, E. G., 119

Cooper, Cooper,J.E.,75nl46 R. A., 103 Copenhagen mechanics, interpretation 115 of quantum
85 butterfly, Cain, A J., 17,70-72,76-77,120 wins debate with G. S. Carter, 77
cosmogony, Cott, H.
105
B., 45 118
Cain J., 2n3,57nll3,60nll9,62nl20, 123n210 Campbell, D.T.,118 canalization, 6,124n212

cancer of stomach correlated with
A.,24n50 Coyne, J.
Cravens, H., Creager,A.,74-75 vs. evolution, creationism blood
5,12,28-29,
groupA, 69
103,113,122,129 Crick, F.,34n77,117
172
Index
Crombie, A. C,
species are
differences between bird

adaptive, 97
Crow, J.R, 9nl6, lln21,14n25-26, 19n37,21,21n41,22,23,25n51, 30n63,31n68,129 Curtis,H., 75nl45 Dalbiez, R., 5n9 dandelion, 43n95 Darden, L., 2n3,37,37n86,129 Darlington, C. D., 17,46,123n210
Darwin, C. R.
37n84,38n87,106nl85,114,121, 123,123n210,129 blood groups, 70-71

chromosome inversions, 62-67, 70, 83,
120,125-126
Demerec experiment 75 on bacterial resistance, evidence
entomology,35, 83
for natural selection and NSH,
39-45,43n95, 64-65
Filipchenko and macromutations,
66nl23
is toPaley asHuygens is toFermat, 119 theoryof evolution, 1-4, 6nl3,11,35 Madeira Islands, wingless beetles of 39-41,39n89,125
G. H., 17
founderprinciple, 71 genetic drift,36, 70-71, 84-85,104,
120
Geneticsand theOrigin ofSpecies,36-39

Genetics of Natural
Darwin,
Darwin, L. 30n65 Darwinism, 2,28n59 France, 40n91,41n92 centennial of Origin, 103 social, 118 Daston, L., 114nl99 de Beer,G., 92nl65 deer mouse, 53nll0
Delbruck,M.,74 Delsol,M.,ln2 Demerec, Demigretta M., 75 sacra, 53 reception, 4-5,5n7,35n78
64nl21
hardening influence of view on biologists 8nl5 et al.
Populations on
series, 81-84
selection, in several
countries, on L'H?ritier
41n93 Mayr,51,52,
Mexican D.
experiment,
41,
71
Pseudoobscura, 67nl25 crossed, barrier
micro/macroevolution 65
personal data, 131 predictions, 104,106,106nl87, 109nl92

45 and warning coloration, protective reason for views on selection changing
Depew, D. J., 10nl7, lln21,27n58,90, 117n202,125n212,126n214

R., 119
Descartes,
119,122 design, intelligent, refutedbyHaldane, 28-29

de Vries, H.
and drift,66,66-67nl25 Wright, 22,23,36,51, 65

Dodson, dominance, E. O., 89 evolution
of, 16-17
mutations, 5, 6nl2, 36 theoryof large with blood diabetes mellitus correlated 60 A, group
Dice, L. R.,44,53nll0
Dorippe jap?nica, 49-50

Dove,W.F.,9nl5,14n26
Dietrich, M. R, 7,34n77
Divall,C.,114
Dowdeswell, W. H., 70nl34, 85, 92, 109nl92,120 Drabble, M., 93 drift, genetic, 1,2 Dronamraju, K. R., 13n24,46n98, 52 Drosophila, 6,22,35,37, 83
D. azteca, 63 44 D.funebris, D. melanogaster,
Diver, C, 52, 76 DNA double helixmodel, 34n77,117,
125
Dobzhansky,T., lnl, 6-7, 7nl4, 8,12, 21n42,30n65,35-38,35nn78-80,
40-41,44
173
Index
Drosophila {continued) D. persimi/isy 36

D.
82, 84 testof prediction,40-41 Dubinin, N. R, 66, 88 21n42,25 genetic drift, 66, 94 strongselection inD. funebris,
Dubos,R.J.,74 Dunkers, Dunn,E. 79-80 R.,31n70
pseudoobscura,
35, 36,54,
62-63,
Huxley, 47 hardening, 7,57-59, 81-90,115 metaphysical foundations, 114 similar to unification of physics by
energy conservation, 116
Simpson, 56 Stebbins, 60
evolutionism in behavioral/social sciences,
118 Ewens,W. J.,27nS7,52nl07

experimentalists experiments, vs. naturalists, 31n69
Dunn, L. C, 2n3,37n85,123n210 with blood duodenal ulceration correlated groupO, 69

ecotypes, 43
controlgroup, 41n92
3
crucial,
Eddington,A. S., 114 Eden, M., 112 Edwards, A. W. R, 9nl6,72nl41 eggs,price of,22 Einstein,A., 3,5,25,124 Eldredge, N., 51nl04
elements,
explanation,vii, 3,4n5, 69nl31,105,107 extinction refutesintelligent design 28 theory,

Falconer. D. S., 80nl52
and falsification, 4,4n5,105 falsifiability M., 20,35 Faraday,

Fermat,P.de,119
Elliott,A. M.
emergence, Emerson, empiricism, A.
origin
in stars, 107
80nl52, 92,102
E., 2 36n82
Filipchenko, I., 66nl23 finalism,116
28,114
Fisher,R. A., 9-18,12n23,13n24,112 blood group prediction based on theory, 68-70

determinism and indeterminism,
and idealism, 116 EndlerJ., 2,2n3, Hn21,120-121

Engels, E.-M., 52 8nl5
Epling,C,
essentialism,
36,54, 62
13n24,116 eugenics, 13n24,17

evolution of dominance, 16-17, 31
eugenics, 13n24,28n59 Euler, L., explanation by teleology

equivalent evolution, factors to effective in, 132-133 causes, 119
experiments,17n33,18,18n34 eyesight,14n26 on genetic drift, 3, 7,23,24,24n50,

mathematics, 9nl6,14,20,51
87
evolutionarybiologists, 2,3,31n65 fashions in theory, 89-90 random drift, 67 replies to Popper and Scriven, 107-109
survey of views survey of views on evidence for natural
mutations and evolution, 14,15n27,125 personal data, 131 on psychological opposition to
Darwinism, 13n24 reception of Gen?tica/ Theory,30, 30n63 scarlet tiger moth, 78, 79
sexual vs. asexual reproduction, 13-14
selection, 120,135
on factors in evolution,
97-99,133 evolutionary synthesis,1,34,36-37,120 founders and leaders, 131 Haldane, 27
speciation, 17,17n31 theoryof natural selection, lnl, 5, 5n8,

7,10-17,10nnl7-18, Hnnl9-21,
15n28,17n32,21,27,30,31,31n67, 32,68,85,104,121-123
174
Index
Fitch, WM.,35n79 fitness,11, lln21,12,22,

five-petalled rose, 43 Flew, A. G.N.,110
68
genetics, 1, 35,37,46 Sixth InternationalCongress, 30n63 bean bag, 17

systematics/taxonomy, 44-45 variation, geographical and 52
Forber,P., 2n3,24n48, 84nl54 Ford, E. B., 17,18,48, 68-70,123n210 balanced polymorphism, 16, 68 blood groups prediction,68-73,104,125 Fisher, 9nl6,30n63,51, 68 genetic drift,87-88, 87nl57 Huxley, 46
industrial melanism, 42
geology based on plate tectonics, 115 Gerson, E. M., 1,13n24, 95nl70, 114nl99 Ghiselin,M., 108-109 Gigerenzer, G., 70nl33, 72nl40 Glass, B., 79-80, 80nl52,102
Glick,T.F.,5n7
natural selection,68, 69, 83, 85 peppered moth, 91, 92 personal data, 131 scarlet tiger moth, 78
P., 114,115
Gloger's rule,44
Glymour, B., 2n3 Goethe,J.W.Von,114
Wright, 23,87
Forman,
founderprinciple, 51-53,58, 71, 84, 86

Fox, H.M.,44
Goldschmidt, R. B., 6nl2,31n67,33-34, 34n77,36,54,58, 89,95,125 hopefulmonsters, 33

gonorrhea, 75
Franklin, A., 18n36 French biology,40n91

Freud, S., 5 D. J., 116 Futuyma,
Goodnight, C.J., 24-25,25n51

Goudge,T.A.,110
fuzzy set,36
patruelis, 92 95
Gould, S.J., 6nl2, 7,32,33,34n77,37n85, 57-58,67 Dobzhansky, 81-83 Goldschmidt, 58 57,58nll4, hardening of synthesis, 81-83
early selectionist view, 86
Gambusia Gardner,
71,
E.J., R. R., G. R,
G., 68nl29, 72,129 Garratty,

Gates, Gause, 32 88
Ruse critique,58nll4 Simpson, 57, 83 snail debate a victoryfor adaptation, 78

textbooks, 99 Grabiner,J.V.,97nl72,129
GayonJ.,l,40n92 Geddes,P.,40
genes, 5,25
arrangements in chromosomes, 16
gramicidin, 74
Grant, Grant, recessive, 27 36 Peter, 97 97 Rosemary,
36-37,54
dominant frequencies, speciation, and
Great Snail Debate 76-78
v. selection), (drift
genetic assimilation, 124n212 genetic code, 117 genetic drift,1,2,11,20-26,48,51, 79-86,94,102,115,120,123n210 and extinction,72,110
holistic antireductionism, 115 116
Greene, J.,46n96,52nl04,52nl06 Gregg, T. G., 60nll7, 73nl42

Gregory, W.K.,37n85 126 Grene, M.,
Grimoult, C,
Guterman, L.,
8nl5,40nn91-92
52nl05
<->nondrift),85 pendulum (drift

genetic homeostasis,
Hagedoorn, A. C,
Hagedoorn, A. L.,
10,21n42
10,21n42
175
Index
10 Hagedoorn effect, Hagen, J.B., 31n69,46n96,52nl04, 93nl68 HaldaneJ. B. S., 27-29,28n59,50nl03, 112 vii, 5,5n8, 7, 9, evolutionary theory, 85, 17n32,27-29,31,31n67,32,33, 104,120-122 on Fisher, 9nl6,28n59 genetic drift,87
on L'H?ritier mathematician, et al. experiment, 28,51 41
Hill,W.G.,19n37 historicism, 104

Hitchcock, C.,4n5
Hodge, M. J. S., 10nl7,23n47 holism, 25,28,34,114,117,125 holisticmaterialism, 115

Holman, R.M.,31n68
Holmes, S.J.,6nl2, 60nll7 Holton, G.,115nl99 Hossfeld, U.,8nl5

Hoyle, R, prediction to make needed of nuclear elements reaction in stars,
personal data, 131 prediction,vii, 103 J.M. Smith, 51 on Wright, 28n59, 87

Haldane,J. S., confused with
107
Hull, D. L., 4n6,106nl85,110nl96 Huxley, J. S., In2,2n3,13n24,15-16,

J. B. S.,
28n59
Hall, B., 125n212

Hall,N.S.,10nl7,18n35
16n29,33,34,34n77,46-50, 46nn96-98,50nl03,121 blood groups, 88

evolutionary synthesis, 47
Hapgood, F, 52nl04 Hardin, G., 68nl29,102,102nl84 Hardy-Weinberg law,9nl6

Har?an, Harman, H. O. V., 61nll8 S., 17 Hardy,A.,47nl00
Fisher, 47,48 Ford, 48 47,48-49, 88 genetic drift, Haldane, 48

Heikecrab, 49-50
Harrington, A., 26n55,115 W. Heslop, 6,48 Harrison, J.

Hartert,53nll0. Hartmann,M.,38,94 Harwood,J.,8nl5,114 Hazen,L.N.,22n44
Russian evolutionary biology in 1945,88 Waddington experiments,124n212

weakness science, Huxley, T.H., in mathematics, 47nl00 12-13 physical
idealism, 114 Mayr, 51-53,53nll0 moths, light and dark, 92 personal data, 131
Heike crab,49-50 Heikertinger, R, 45 Heitz, E., 36 Hempel, C. G., 105

Henderson, heredity, Heredity, An M. G., 5n7 3 theory of, Journal of chromosome International 17
Huygens, C., 119 hybridization,60nll5 hypothetico-deductivemethod, 2 not onlymethod allowed in science, 108 idealism and empiricism, 116
indeterminism, 25 hydrocyanic gas as agent of selection, 42
Genetics,
W., 6 Heslop Harrison, J. Hess, C, 79 19 heterozygosity, hierarchical thinking,81

Hiesey,W.M.,43
industrial melanism, 42, 91-92,120,122 110 reversible,

inheritance
Highton, R., 129 Hill, B., 36n82
blending, 11
particulate, insecticides, 10 resistance to, 74,120
176
Index
isolation,geographical, 1,2,21, 36, 37, 47,52

Ives,RT.,31n70
Lamarckian
effects, 2,5-6,
6nl3,28,
40n91,48,53nll0,57nll3,74-75, 95,115,122,124,132-133
Lamotte,M., 104
Jahn,E. R, 79 H., 10,10nl7,114,117 Jeans,J.

Jepsen, G. L., 6nl2
evidence fordrift,77, 86,125 Lancefield, D. E., 35

Landauer,W.,109nl94
W. L., 6nl2, 32n71, 60 Johannsen, pure line theory influencedby culture or fashion, 60nll6, 89 K., 52nl06 Johnson,
Johnson, P. E., 104
H., 109nl93 Langenheim, J. L. 57nll3 R, Laporte,

Larson, E. J., 97 Laubichler,M.D.,8nl5
T., In2, 8nl5,13n24 Junker,

"just so" stories, 50,103 6nl2
Laudan, L., 109nl94 Lawrence, C, 26n55

trial and error, analogous learning, 118 natural selection, Lederberg, E. M., 109 75 to
Kane,T.
C.,
Kaplan, J.,2n3 Kaplan,M.,lllnl97

Karpechenko, G. D., creation
Lederberg, J.L., 75
Lehman, of new H.,
species, 35n80
D. D.,43
Keck,
Leigh, E. G., Jr.,Iln21,106nl88 M., 22n44, 60nll7,115 Lerner, I. Lessard, S., Iln22 Levine, L., 8nl5,35n79,67nl25
Lewis, R.W.,106nl88 R. C., Iln21,17n33, Lewontin,
Kerr,J.G.,6nl3
W. E., 22-23,23n45,104,121,125 Kerr, Kettlewell,H. B. D., 91-92,120 W. C., 5n9,45,68nl27 Kimler,

Kimura,M.,9nl6,24
kinetic theoryof gases, 10,10nl7,14,24, 117 Kingsland, S. E., 31n69 A. C, 52 Kinsey, Kipling, R., 103 Kitchin,R D., 69nl30 Kleinman, K., 60nll5 Kohler,R. E.,31n69,40n90 Kolchinsky,E. L, 8nl5
Koprowski,H.,lllnl97 N. L., S., 3 36n82
23n46,36n83,38n86,73nl42, 108nl91,109nnl92-193,114nl99, 126nn213-214,129 Dobzhansky sGenetics ofNatural Populations, 64nl21 Liberty Baptist College, 129 lightbending, 3 Lip ton,P., 4n5 Lloyd, E. A., 106nl87,110nl96 Lloyd Morgan C, 114
Losee, Luria, J., 4n5 S., 74 Lindsey,A.W.,5n9 Cartesian-analytic method, 117
Krementsov,
Krimbas, C. B., 8nl5,37n85

Kuhn,T.
Lurgin, P. R, 72 Lustig,A., 6nll

Lysenko,T.D.,74,88 40-41 MacBride,E.W.,5,6nl3,32
LH?ritier, P., 31,40nn91-92

test of Darwinian
population cage/box,41n92, 82 Lack, D.,5n9,22,123n210 changed toNSH because of own

research, 95-96
prediction,
Maienschein, J., 114nl99

macroevolution and microevolution, no
Lakatos effect, 123-124
36,53, 65, 83, qualitative difference,
125
177
Index
malaria/sickle cell polymorphism, 73

Malecot,G.,22n43 Malthus,T.,9nl6
McCauley,WJ.,90nl63
McDouga?,W.,124 McKerrowJ. C.,5n9
Mani?la jurtina, 85
Manser, Martini, E., M. 109 L.,61nll8 109nl93 monism, 28,
McKusick,V.A.,80nl52 McOuat,G.,28n59,29n62
Maryland, equal time law, 113

Mason, H. L., materialism, materialistic
114,115 holistic, 115 Mather, K., 15,30n65,34n77, 86nl56

Matthen, MavorJ.W., Maxwell,J.C.,20 M. 2n3 101-2,129
mechanistic biology, 34,114,117 complementary to teleological biology, 119 Medawar, P., 109nl93, lllnl97 A., confirmationof Ford's Mehigan, J. prediction, 69 melanism, industrial, 42,91-92,122
G. results, 18 experimental mendelism
Mendel,
theoryof heredity,1,10,14n25,16,49
Mayr, E., In2,5n9, 6nl2, 8, 8nl5,13n24, 21n42,31n69,32,33,34,52nl04, 112,114,123n210 bean bag genetics, 17n32,52nl07,120
chromosome inversions, 62
faddish,95
Mercury, perihelion advance, 3
Merrell, DJ.,92,124n212 Mettler, L. E., 60nll7,73nl42

microbes, 74
Dobzhansky, 52nl07,54
"father of modern
biology," 52
evolutionary
Fisher, 52nl07,121-122 Ford, 52nl07,55 founderprinciple, 51-53,55, 84, 86, 86nl56 genetic drift rejected,54, 88-89 Goldschmidt, 34n77,54 Haldane, 52nl07,122 historian, 52
mathematics, 52
midcenturymodernism, 115 Miller, A. L, 14 Miller, P. D.,97nl72,129 Mills, S. K., Iln21 Millstein, R. L., 2n3,23n47,24n48, 25n53,77nl48, 86,123n210 mimicry, 18,49,68,103,120,122 Moberg, C. L., 74nl43,75nl46 models in scientifictheories,17,17n33,
112
predictions cant be made from theory

of natural selection, 107-108,
natural selection,53nll0,94 organic selection criticized,125n212 personal data, 131 predictionbased on Epling data, 54,125
Moody, P.A., 13n24,92,93,124n212, 125n212 Moon, origin, 17 Moore, J., 17,31n66

Moore, Morgan, Morgan, R., 24n49 114 3,5n9,6nl2,12,30n65, C. L., T. H.,
126n214 Rensch's priorityforfounderprinciple, 53nl09

sexual selection, 13n24
35,35n78,46 on evolution, 30, 30n65

Morris, Morris, Morrison, C.W.,109nl93 H., M., melanic 122 10nl7 <-? nonmelanic
species definition,36n82
turn toward selectionism, 54
mosquito fish,95
moths,
Wright, 52nl07,53-54,122
McAtee,W.L.,32n71,45
forms,
6,
McCarthy, J.R., 13n24
27,42,68nl27,91-92,110,120
Mourant, A. E.,69nl31
178
Index
Muller, H J., 14n25,16n29,30n65,31n70, 116,125 21n42,25 genetic drift, mutations produced byX-rays, 6, 7nl4, 31,31n68,36,37n84, 67,122
mutations, 2, 6nl3,15,31,36,37,48,115,
Neefs,Y. 40-41 NeelJ.V.,72
, test of Darwinian
prediction,
neo-Darwinism, neorealism, neutral 117
1,2,111-112
neoromanticism,
25,34 evolution, 88 71, 85
122,142-133 idealistphilosophy, 116 large,5, 6,14,33,34n77,36,48,58, 132-133

in production of resistance to
(non-Darwinian) name, 5n9
Newman,
inappropriate H. H.,
Noland, G. B., 102

nondisjunction, 3
fumigation,42 Nagylaki,T.,22n43
Nanay,B.,2n3,61nll7 natural selection, 1,2,4-7
Norton, B., 13n24,17

Norton, novel H.T.J.,15 fact, 4n5
Numbers, R L., 5,122n209 Nye,MJ.,124n211

0'Brochta,D.A.,41n92,129 Oenothera, 60
as a cause,
analogywith history of science and technology,118

2,2n3,16
as a fact, 2
confirmedpredictions, 39-41, 54, 68-69,111 denying it is "like flyingin the face of the wind," 41n92
dissatisfaction with
Olby, R., 6nl2, lln21,17,17n31 Oppenheim, R, 105 organic selection, 124n212 ornithology,51
Orr,H.A.,36n81 orthogenesis, 6,6nl2,28,48,56,59,95,
109nl93
evidence from inversions, as a force, 2n3 fundamental 65
the concept, chromosome
115,122,124,132-133
seasonal Osborn,H.R,5n9 Otte,M., 11 Painter, T. 114
theorem,
S., 36 with 57nll3
and human (artificial)selection,42 hypothesis (NSH), 2,7,35,48,53,65, 82,120 and kinetic theoryof gases, 10nl7 as only a negative factor, 5,31, 61nll7 13 force, peaceful
prediction as a process, reasons for from Darwin's 2 accepting, 119,135 Origin, 39-41
paleontology, 1, 56, 64,124

collaboration
Gould critique of Simpson for devaluing it,56-57

Lamarckism Panaxia of Germans, 78 57nll3 dominula,
systematics,
Papilio dardanus, 68
Park,T.,19n37
as thema
sculptor analogy, 60
in behavioral/social sciences, and
118
naturalists <-? vs. experimentalists ~
Parker, G., 5,5nl0 T., 4n5, 7nl4, 34n77 Patterson, J. 84-85,104,121,125 O., Pavlovsky, Pearl, R, 103 ^nondrift), pendulum (drift (holistic ^mechanistic),
(evolution <-? antievolution),
85; 118;
118
geneticists, 31n69,37
(nature nurture) mutations (large natural selection), 118
penicillin, 74-75 peppered moth, 91-93,120
Nazis, 12,13n24
179
Index
PeromyscuSy
53nllO
PerrinJ., 124 Pfeifer,J.,123n210 philosophy of biology, 3,23-24,123n210 philosophy of science, 3,10,103-106, 109-111
must be based on historical research,
logical, 108 novel, 2n4,3,4n5,47,64,72,125 about past, 108 temporal, 108 testable,3,25,103 Price, G.R., 11,Hn21 Prout,T., 38n87,106nl86,106nl89 Provine, W., In2,6nl2,32,36n83,37n85, 46n96,52nl04,72,121nn207-208,
111 phylogeny,38,106 physics, 3,4n5

optics, variational
129
elementaryparticles, 115 119

principle of Fermat,
Dobzhansky's selectionism, evolutionary constriction, Genetics
turn toward 65 "synthesis" 1,47,115 Populations series, is really a
statistical,10nl7
theoretical, 9-10,17,20
of Natural
Pigliucci,ML, 2n3 Pilpel,A.,18n36 Plough, H.H.,31n70 A.,2n3, Hnn21-22,17n33, Plutynski, 23n47,24n48,123n210,129 polymorphism, 52nl07
adaptive, 63-64
64nl22 Great Snail Debate, 76-78 Great Tiger Moth Debate, 78nl51
on Mayr's 95 turn toward selectionism, 54 Sumner,
balanced, 16,64, 83,120 polyploidy, 1,2,33,36,60nnll5-116 Popper, K., 2,2n4,3, 3n5,40,126, 126n214
conjectures natural on Darwinism and refutations 118 analogous to selection,
on Wright, 7,19n37,20,20n38,21, 21n42,22,23nn46-47,24n50, 81-82, 86, 66nl23,75,75nl48, 115-116

pseudoscience, 2
psychoanalysis,5,5n7
Punnett, R.C.,5n9,15
and neodarwinism,
quantum
mechanics,
4n5,25,28,105,115,
13n24,104?l05,109-110,112nl97, 113 teleology reducible to causation, 119 population, 10, 37 cage, 40,41n92,64 genetics, 10-25,106,117
natural, Potentilla Poulton, 18,35,37 43
117n202
Quayle,HJ.,42 Quetelet, A.,9nl6

randomness, 25, 85
52 thinking,
g/andu/osa, E. B., 30nn63-64
metaphysical aspect, 115 outcome maybe predictable, 116-117

Raphanobrassica, 35n80
Powell, J. R., 30n65,36n81, 85 prediction,vii, 2,2n4, 3,4,18,105-111, 125-127 absolute, 108 in advance, 2n4,4n5,27, 111 blood groups, 69-71
chromosome inversions, 63-64
92 Ray, C., Jr., realism, 115,117,117n202

reception studies, 4-5,5n7
reductionism,34,114,115,117 refraction, explained by teleological or mechanistic principle, 119

Reif,W.-E.,8nl5
Reinig,W.R, 52 Reisman, K., 2n3,24n48, 84nl54
180
Index
5,5n7 relativity theory, 3 general theory,

religious influence on science, 40n91
M., Sch?tzenberger,
science can it be demarcated
112
from
Rensch, B., 52,53nll0,57nll3,116

reproduction asexual, sexual, resistance resistance 13-14 12-14 to antibiotics, to insecticides, 74-75,122 42, 74,122
pseudoscience?, 2,105,110 philosophy of, 3,4 scientific method, 4,40,103,126 5,129 Trial, Scopes Scott, E. C, 122 M, 25n54,105-107 Scriven, SewallWright effect, 21,24,47,48
sexual selection, 13n24,103 86
rhetoric,37, 71-72 Rhodes, RH.T.,13n24 Richards, O. W., 25,31n67,32 Richards, R. J.,6nll, 125n212 Richardson, R. C, 6nl2
Roberts, J.A. R, confirmation of Ford's
Reydon,T.A.C.,36n82
Shahn, E., 129

Shanahan,T., In2,
predictiion,69, 70
Robins, W.W.,31n68 Robson,G.C.,25,31n67,32
rodents in lava flows,53nll0

Roman, H., D. 56 43 7nl4
Shapere,D., 110 Sheppard, P.M., 17,34n77, 70, 70nl34, 72, 76-78,79, 92,109nl92,120, 121n207,123n210 Cold SpringHarbor paper suppressed, 78 Shull,A. E, vii, 21n42,31-32,32n71, 103 sickle cell/malariapolymorphism, 73,120 signs, theoryof, 109nl93
C.W.,74 G. G., 6nll, 49,56-59,
romanticbiology, 114,115
Romashov, Romer, A., D., 21n42,25
Simon, Simpson,
rose, 5-petalled, 92 Ross, H. H.,
Rudge, D. W., 64,91nl64, 93nl68,129 Rudwick,MJ. S.,109 Ruse, M., Iln21,20n38,22n44,24n50, 52nl04,58nll4, 68nl27, 69nl30, 84nl54,106nl88,110
Sacks, M. S.,79
109nnl92-193,114,120 56,58,58nll4 genetic drift, Gould's critique,57

horse an example of natural selection, 56
Mayr,56nll2 moths, light and dark, 92

macromutations and idealism, 116
Sagan,C, 50nl03 salivary glands, 36, 37n84

saltationism, Salzano, F M., 6, 37 72
Sarkar, S., 5n8,27,27nn56-57, M, 13n24 Savage, J.

scale insects, 42 view of theories, 10
38n87
predictionsmay be made about past, and tested,108 2n3, 7,17,20,20n38, Skipper,R. A., Jr., 86 23n47,24,68nl27,78nl51,
Slatis,H.M., 72
organic selection rejected, 125n212 orthogenesis rejected,56,124 personal data, 131
scarlet tiger moth, 78

semantic
Sloan, P. R., 52nl06 Slobodkin L. B., 109nl92 Smith,J.M.,109nl92 Haldane,28n59,51 Smocovitis,V. B., In2, 33,43n95, 50nl03, 60nll5,129
Schindewolf,O., 58 Schmalhausen, 1.1., 6nll, 125n212 Schmitt, S., 34n77 Schr?dinger,E., 117
181
Index
Smocovitis, reason
V. B. for
{continued) change 124 to
tapeworm refutesintelligent design, 28-29 Taraxacum officina/e, 43n95

teaching undergraduate taxonomy, 51
NSH,
Smoluchowski,
67
Dobzhansky's M. von,
courses
snails,72,76-78,120
Snyder,L. H.,22n44
Sober,E., 2n3,4n5,4n6, Hn21,110nl96, 119,123n210

social construction, 17,129 14 space, multidimensional,
failure,34n77 Teilhard de Chardin, P., 112nl97 59; reducible or equivalent to teleology,

causality, Teissier, G., 82 119
sign of
N. P., 104 Spassky, speciation, 1,17,35-36,49,52,125 36 hybrid sterility, species,definition,36n82 fuzzy set,36
species differences are
testofDarwinian prediction,40-41
A., 5n9 on evidence for natural 12
Tennyson, textbooks,
survey of views selection, 134 survey of views 21,
on natural
selection
and
non-adaptive,
other factors,99-102,132 thema,115nl99 Thomas, R., 129 Thompson, P., 52 Thomson, J. A., 40
N. W., 33,46,52 Timofeeff-Ressovsky, 66 seasonal cyclic changes, N., 91-92 Tinbergen,
25,32
Spencer, H., 114 Srb,A.M.,92 Stadler,L.J.,7nl4 75 Staphylococcus aureus, statistical methods, 18,22 Stebbins, G. L., Jr., 2n3,34n77,42,60-61, 62-63
123n210
chromosome inversions,
de Vries mutation theory, 60nll6 89-90 fashions in evolutionary theory, 89 drift, genetic 60nll6 Johannsenpure line theory,
mutations, 89
Tiniakov, G. G., 61,66,94,95nl69 Toulmin, S., 110-111

Turesson, G., 43
Turner, J.R. G., 23n47, 68nl27,71,72, 77nl49,91nl64,123n210 Ulam, S.,112

universe, age of, 105-106
personal data, 131

stem rust of wheat, 42
C, Stephens,
Stone, L., 72
123n210
Van Helden, 47
variation or variability, 2, 7,11,18, 35-37,
Stone,W.S.,4n5,34n77 Stratton,F.J.M.,10nl7 struggle for existence, A. H., 12
geographical, 44-45
human, 67
42,52,74
Struthers, D.,69nl31 Sturtevant, 64, 65nl22
variational principles (teleology) in physics

equivalent Veuille,M.,40n92 Villee,C.A.,4n5,25n54 to causal equations, 119
stylesof thought,114
Sukatschew,W.,43n95
Sumner,F. B., 34n77,45,53nll0,95

reluctant conversion 95 to selection by own experiments,
vitalism, 116
Vries, H. de
Supreme Court, 4 Sylv?n,61nll8

systematics, 51
mutations, 5,6,15, 36, 37, theoryof large 48,60,101; 122,124-125,129

and hereditarianism, 118
Vucinich, A.,35n78
182
Index
Waddington, C. H., 6nll, 50nl03,109, 109nl92,112 on genetic drift,87nl58 onT.H. Huxley, 13n24
on L'H?ritier et al. experiment, 41
Winsor, M. P., 28n59,29n62,36n82, 52nl06,129 Wright, S., 19n37,20n39,23n45,51, 115-116 blood groups, 22 Dobzhansky, 36,66nl23 5, 9,17n32,19-26, evolutionary theory, 20nn38-39,25,27,31,31n67,32,49, 70,85-86,104,112,121-122 genetic drift,3, 7,20-26,21n42,24n49, 85-86,104,125 genetics of natural populations, 62, 65, 66nl23
hardening of view on selection, 81-82
Wade, M.J., 24-25,25n51

Wallace, A. R., 6nl3
Wallace, B., 92,109nl92,125 Wallace, C, 13n24

Waters, C. K., 47
Watson, D. M. S., 32,34n77,56,121n207 Watson, J. D.,34n77,117

Weatherwax, R, 33
Weber, B. H., 10nl7, lln21,27n58, 117n202,125n212 Weismann, A R L., lnl, 5

Weisskopf,V.F.,112 Weisz,G.,26n55 Weisz,RB.,13n24
Mayr,52,52nl07 path analysis,22 personal data, 131

chooses selection not drift in Great
Wells, G.R,2n3
Wells, H. G.,2n3
White, M. J. D., 34n77,37n84

Whitehead,A.N.,114 Wigan, Willermet, L. G.,15 C.M.,36n82
Snail Debate, 77 balance theory, 7,20,22,24 shifting 21n42 effect, Wright-Dubinin

Yoon, C. K., 52nl05
Williams, G. C, 125n212 Williams, M. B., 106nl85,110nl96

Wilson, wingless E. O., insects 37 as evidence 39-41,110 for natural
Zachos, R,110nl96
Zahar, E., 4n5 N., 129 Zimmermann,
zoology, 1
selection,
183
CHOOSING
This
SELECTION
book describes the establishment of the hypothesis thatCharles

Darwins "natural selection,"reformulated by R.A. Fisher, J.B.S. Hal
Mendelian genetics, is the primary or dane, and S. Wright in the Hght of exclusive mechanism for biological evolution. During the 1930s, alterna
tives such as Lamarchism, macromutations, and
were dis in favor of natural selection acting on smallmutations, but there agreements about the role of random genetic drift in evolution. By the 1950s, research byT, Dobzhansky, E.B. Ford, and others persuaded leading was evolutionists that natural selectionwas so powerful that drift generally a was conclusion This most; however, unimportant. accepted by significant and articles of textbooks in the mentioned drift minority popular biology late 1960s.
orthogenesis
were
rejected
G. Brush was employed as a theoreticalphysicist at theLawrence Stephen Liverrnore Laboratory (1959-1965), a staff member at Harvard Project Physics (1965-1968, developing a historically oriented physics course for high schools), and Lecturer in Physics and History of Science at Harvard University (1966-1968), From 1968 through2006 he served on the facultyof theUniversity of Maryland, University Park,with a joint appointment in the Department ofHistory and the Institute forPhysical Science & Technology. retired in 2006 and holds the title Distinguished University Professor of
of Science, Emeritus. the History
He

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Choosing Selection: The Revival of Natural Selection in Anglo-American Evolutionary Biology, 1930-1970 Author(s): Stephen G.

The Revival ofNatural Selection in Biology, 1930-1970 Anglo-American Evolutionary

AMERICANPHILOSOPHICALSOCIETY | PHILADELPHIA | 2009

The <rR?vival D^atural Selectionin of

PHILOSOPHICAL "Philadelphia 2009

Useful Knowledge Held at Philadelphia forPromoting Volume 99, Part 3

American Philosophical Society for its Copyright? 2009 by the

and cover design

978-1-60618-993-1 issn: 0065-9746

Brush, Stephen G. : the revival of natural selection of the American

evolutionary biology,1930-1970 / StephenG. Brush,

forpromotingusefulknowledge, issn 0065-9746 ;v. 99, pt. 3)

selection?Research?Great selection?Research?United Britain.

Britain?History?20th States?History?20th States. 4. Biologists?United

century. century. I. Title.

from Collected Papers

Snails and Tiger Moths 79

FACT THAT NATURAE

planation of adaptation acceptance of Darwinian tion is very widespread, abundant

generallybelieved by historiansof biology to have been established in the

Why did biologists accept themodern version ofDarwins theory?I

Junker(2002) on the terminology.

the reception by evolutionists

and other biologists

of thought"(Depew and Weber 1985b,240)? If so, ing empiricistcurrents

pseudoscience?4 That question

and tested. Conversely, Einstein's

inphysics?the discovery Albert ofgravitationallight bending, by predicted

of the "predictivism" "accommodation") Sober (2004)

may be as good and Scerri andWorrall

6.Hull (1974); Sober (1984a, 1984b,2006); and recentissuesofStudiesin HistoryandPhilos

Selection in the 1930s

I startthe storyat thebeginning of the 1930s, with thepublication of the

But Lamarckism, though supportedinEngland byE.W. MacBride

revived, then and

search of the online

of the Isis Cumulative

and Darwin reception 8. For

seeBowler (1979; 1983,144,162,178-81); Holmes (1948a); Jepsen(1949); On orthogenesis,

not discuss period; I do of drift after 1965. This

what Stephen Jay Gould

of the synthesis.) [1983] called the "hardening"

directionsbeginning in the late 1960s (Dietrich 1994). lengedfrom several

14. Muller (1927); Pattersonand Muller (1930); andDobzhansky (1981).According toRo

cept for historical 1970. This American

relevant to the period before

as well study is limited in space community. Of biological

as in time: it covers only theAnglo course this includes community

15. For France,

Fisher: A New Language forEvolutionary Research

pecially, in the language cannot

biology, look up individual words

the background knowledge and styleof thought of thosewho use the

to keep the particles astir.

18.Fisher and Stock (1915, 60) andFisher (1922,321).

Fisher: A New Language for Evolutionary Research

genetic drift, discussed

next) must be insignificant

except in small popula

Tripos examination) would and molecular velocities. See

(Bowler 1978,55). by struggle"

Fisher (1930b). 23. Fisher (1930a,47); cf.