Abstract
Ecology in principle is tied to evolution, since communities and ecosystems result from evolution and ecological conditions determine fitness values (and ultimately evolution by natural selection). Yet the two disciplines of evolution and ecology were not unified in the twentieth-century. The architects of the Modern Synthesis, and especially Julian Huxley, constantly pushed for such integration, but the major ideas of the Synthesis—namely, the privileged role of selection and the key role of gene frequencies in evolution—did not directly or immediately translate into ecological science. In this paper I consider five stages through which the Synthesis was integrated into ecology and distinguish between various ways in which a possible integration was gained. I start with Elton’s animal ecology (1927), then consider successively Ford’s ecological genetics in the 1940s, the major textbook Principles of animal ecology edited by Allee et al. (1949), and the debates over the role of competition in population regulation in the 1950s, ending with Hutchinson’s niche concept (1959) and McArthur and Wilson’s Principles of Island Biogeography (1967) viewed as a formal transposition of Modern Synthesis explanatory schemes. I will emphasize the key role of founders of the Synthesis at each stage of this very nonlinear history.
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Notes
In the introduction to a special issue on ecology and evolutionary biology, James Collins, John Beatty, and Jane Maienschein consider “the changing role of evolutionary theory in the solution of ecological problems” (Collins et al. 1986, p. 169). The present paper attempts to provide a more systematic account of the acclimatizing of the Modern Synthesis, in particular within ecology, over the course of five decades.
Julian Huxley to Ernst Mayr, 3 September 1951. Papers of Ernst Mayr. HUGFP 14.15 Box 1. Harvard University Archives, Cambridge, MA.
This dual core of the Modern Synthesis is also how Smocovitis characterizes Dobzhanky’s network activity, which fostered the Synthesis at Columbia in the late 1930s and early 1940s: “in so doing they began to bind the heterogeneous practices of evolution into an evolutionary network grounded in genetics and selection theory“(Smocovitis 1992, p. 29).
Both Smocovitis (1992) and Cain (1993, 2009) concur in arguing that this eclipse relies mostly on the fact that, until the achievements of population genetics in the 1930s, evolution was addressed mostly in a natural history style and not through quantitative and experimental methods—which, in turn, were the hallmarks of genuine science. The rise of the Modern Synthesis should therefore be understood as a reaction against this eclipse, which set evolution among the scientific objects approachable through experiments and measures, especially because of its genetic material basis, as begun by Dobzhansky (1937).
Evolution was at the same time crucial and controversial for ecologists in the early twentieth century. As Ilerbaig indicates, this provided room for an emphasis on experimental physiology or natural history (2013) descriptions rather than evolutionary reasoning. “One common concern seemed destined to keep together this increasingly heterogeneous biological community: the centrality of the problem of evolution. However, the alignment of particular theoretical, methodological, and institutional positions fed the existing dismembering tendencies, making the study of evolution more a ground for dispute than an occasion for unification” (Ilerbaig 1999, p. 456).
On Warming’s Darwinism, see Coleman (1986, p. 192).
See van der Valk (2014) on Clements’s climax and its reception.
“The equations that formed the theoretical core of population ecology—were hardly ‘evolutionary’ in any standard sense of that term” (Collins et al. 1986, p. 174).
“Ecologists argued that present environmental conditions could be invoked to account for plant distribution and abundance, that is, in a manner more analogous to a physiological explanation” (Collins et al. 1986, p. 171).
“In contrast, the development of the science of ecology has been hindered in its organization and distorted in its growth by the separate development of plant ecology on the one hand and animal ecology on the other” (Clements and Shelford 1939, p. v).
On Shelford, see Benson (1992).
“A signal extension of ecological ideas is involved in the application of climax and succession, that is of development, to lake and ocean” (Clements and Shelford 1939, p. 4).
“Development is the basic process of ecology, as applicable to the habitat and community as to the individual and species (Clements 1904, 1905). It recognizes that life constitutes a dynamic system and that static studies are valuable only as they throw light on development or serve some practical purpose in this connection.” (Clements and Shelford 1939, p. 3).
“Plant ecology is physiology carried into the actual habitat, and in consequence its paramount theme is stimulus and response. It confines itself primarily and exhaustively with the cause-and-effect relation between the habitat on the one hand, and the organism and the community on the other. All further relations arise out of this, and all other approaches are incomplete unless they lead back to it. With the inclusion of animals in the biotic formation (biome), this relation naturally becomes more complex, but it is none the less valid” (Clements and Shelford 1939, p. 3).
See also Smith (1952).
On Shelford, see, for instance, Ilerbaig (1999, p. 457): “Physico-chemical reductionism and experimental manipulation, the hallmarks of physiology, were the bandwagon on which Shelford jumped in his attempt to make ecology more legitimate.”.
As Kingsland noted: “by providing ecology with a solid theoretical base, mathematics would raise the status of ecology to the level of the physical sciences” (Kingsland 1986, p. 243).
“In ecology the use of mathematics was parallel to, but mostly separate from, its use in population genetics” (Kingsland 1986, p. 237).
Kingsland (1986) indicates that Thompson’s skepticism was also caused by his defiance regarding the recently mathematized population genetics. “After 1930 Thompson feared that the mathematical arguments in population genetics put forth with such assurance by Fisher and Haldane were giving natural selection a new popularity, even though the lack of solid supporting evidence was almost as great as before’’ (p. 252).
“Since Elton’s treatment of communities is without question the best of the decade, we can do no better than examine the state of this phase of ecology as seen through his eyes” (Allee et al. 1949, p. 58).
This is sometimes considered as part of the foundation of the discipline, and volumes like Richardson (2011) are reflection upon its seminal impact and legacy.
Cain (2010) developed this interpretation of Huxley’s trajectory by focusing on his career and activity as director of the London Zoo.
Another social metaphor through which he addressed animal communities was human industry (see Hagen 1992, p. 56).
For instance, in a book Elton published 3 years later that was wholly devoted to evolution and animal ecology, he noted that “in most cases of irregular migration on a large scale the migrants perish, [thus] the instinct to migrate cannot therefore have been produced by natural selection, since it is the butterflies which do not migrate that survive, and those that migrate that perish. If migration is a biological advantage to the lemmings, how is the instinct to migrate perpetuated in the species, since all the animals that carry and exhibit this impulse march downhill into the lowlands, to be eaten by dogs?” (1930, p. 36). From cases like these, he infers that migration is an irreducible animal impulse driving evolution in addition to selection. Given that migration can lead to habitat selection, he concludes that there are two processes at play in evolution, so that we see “a process which may be called the selection of the environment by the animal as opposed to the natural selection of the animal by the environment. In evolution there are two variables—variations of the outer environment in place and time, and variations of the characters of species in place and time. From the interaction of these two variables, adaptation has been produced” (1930, p. 51). Notice how close this formulation is to the dual-process view of adaptive evolution vindicated today by niche construction theorists, even though, of course, Elton did not consider the same evidence and rather considered what we call habitat selection.
“The core of the synthetic theory is pretty much just the theory of population genetics” (Beatty 1986, p. 125).
A geographical comment: the fact that the Synthesis, at least in institutional terms, has been mostly elaborated in the UK and US explains why my story focuses on those two countries. Granted, there was development of the Synthesis in Russia, Germany, and many other countries (as Mayr and Provine acknowledged (1980) and many subsequent scholars confirmed), and major ecological work has been done in Russia, Denmark, and Germany, but asking about the Synthesis in ecology implies focusing on the US and UK.
Smocovitis (1994, p. 277) suggests that the name of the journal Evolution—a single word—and its two column style was inspired by the journal Ecology; Mayr and Emerson were regularly corresponding during this period.
As for Shelford, Ilerbaig shows how he decided for physiological viewpoints to animal-environment interactions against an evolutionary approach: “Shelford recognized two distinct points of view for biological investigation, namely evolution and physiology. In his opinion, the former had repeatedly failed to organize properly the facts of natural history. Thus, it was time to move along more physiological lines” (1999, p. 457). Regarding Clements, van der Valk notes: “Clements famously stated that ecology was ‘nothing but a rational field physiology.’ Even interactions among plants, like competition, could be explained by changes in the physical environment caused by the plants, like the reduction of available light or soil moisture” (2014, p. 3).
In detail, however, their conceptions did not emphasize the same aspects: “Allee saw cooperation as a principle underlying the evolution of sociality and also embodying the tenets of his Quaker philosophy, while for Emerson, cooperation was important because it contributed to greater homeostatic control; it was homeostasis that was the phenomenon of interest” (Mitman 1988, p. 187).
My emphasis. Granted, Elton (1930) had a similar systemic view, as when he says: “animal community forms a highly intricate system of interlocking parts, and the actions of any one species affect not only its next neighbor in the chain of food and other relationships, but through this neighbor, and its neighbors, all other species. Thus a wave of disturbance set up in one part of the system may produce unexpected reverberations in other parts of ramifying branches” (p. 16), but this is rather an insight than a developed theoretical articulation as presented in Allee et al (1949).
As Hagen (1992) established, this idea of the metabolism of a community—later, an ecosystem—has been foundational for “ecosystem ecology,” developed by Odum on the tracks of Lindemann and Hutchinson. However, this ecosystem ecology relaxed the ties to evolution, while here it is presented as a concept proper to an evolutionary framework.
Elton already anticipated this view (even though he had a heterodox conception of the scope of selection) when he said that “the whole of an animal community can act as a biological unit, operated upon by natural selection so as to bring about the best compromise in the way of optimum population for all” (1930, p. 75). Also, that “adaptation is produced by the selection of whole populations rather than the selection of individuals, and whole it raises one huge difficulty by reopening the species problem, it does away with another huge difficulty, of seeing how in practice natural selection could ever be effective in picking out a single individual and succeed in leading it, as it were, through the perils of life in a fluctuating animal population” (p. 30). This “difficulty” shows once again that Elton was quite far removed from the Synthesis understanding of natural selection, as the population geneticists initiated it. Unlike his 1927 treatise, this 1930 book was not influential.
“The protocooperation and cooperation of organisms of the same and of different species have great survival values. This is evidently an important corrective of the naive view that evolution is promoted exclusively by the struggle for existence. The ‘struggle’ involves cooperative as well as disoperative elements, and the former are adaptively more efficient” (Dobzhansky 1950, p. 279). “Disoperation” was coined by Clements and Shelford (1939) to “indicate organismic operations that have immediately harmful effects” (Mitman 1988, p. 181).
Such a Clementsian-Wrightian view of community ecology would be fatally affected by the demise of group selection in the 1960s, when Williams (1966) issued his devastating critique of Wynne-Edwards (1962), who in answer to Lack (1954) had made explicit the thesis of a selection on groups to explain the self-restricted consumption of organisms. On Lack and Wynne-Edwards, see the contrasting views of Kimler (1986) and Borello (2003). Interestingly, Borello traces Wynne-Edwards’s interest in population ecology to Elton’s influence, his mentor at Oxford (532). Kimler argues that Williams’s critique of group selectionism was a key advance in forming “evolutionary ecology,” though this is mostly valid for behavioral ecology; the integration of Synthesis views in ecology overall—addressed here—is a much more complicated story.
“When the number of scales became exceptionally high, I argued, this constitutes an increase in the food supply of the enemies, which consequently increase in numbers and so collectively search the trees more intensively. Carrying on this type of argument, I concluded that the scale insects and their enemies would strongly tend to reach a balance at which the number of scale insects is just sufficient to support the right sized population of enemies and to destroy the surplus numbers of scale insects produced” (Egerton 2014, p. 157).
Ironically, in the early 1930s Andrewartha had reviewed the manuscript of Nicholson’s intended book on natural control and rejected it, so Nicholson finally had published some of the materials as the 1933 paper on the “balance of animal populations.”
Such a point was dramatically underscored in the short paper Sewall Wright published in Science to honor the accession of ecologist Thomas Park to the presidency of the AAAS in 1960: “Humankind question of the very persistence of mankind, or at least of civilized man, in the explosive situation brought about by the world-wide decrease in mortality rates and the lack of compensating decreases in birth rates. The alternatives are violent reduction or even extinction of the human species, perhaps by way of the hydrogen bomb: expansion to a violently fluctuating upper limit, controlled by the availability of necessities for bare subsistence: or attainment of ecologic equilibrium with the resources of the world at such a level that progress in civilization remains possible. Park brings a keen awareness of the population problem to the thinking of organized science” (Wright 1961, p. 502).
As Palladino (1991) indicates, the status of ecology in the 1960 s was deeply affected by the rising public concern about problems that were usually dealt with theoretically by ecologists: “The rapid expansion of theoretical population ecology and systems ecology during the 1950 s and 1960 s [was] indicative of a general desire to project an image of scientific respectability for the field as a whole. However, this depressed state of affairs began to change quite rapidly in the early 1960s in the wake of growing public concern over the degradation of the natural environment—radioactive fallout from the testing of atomic weapons and Rachel Carson’s writings had done much to arouse public interest” (p. 233).
In the same way, in his 1954 book, Lack worried about theory construction in population research: “The only theoretical concepts so far put forward are highly simplified, based on a priori arguments, and expressed in mathematical terms in a few abstract and difficult papers. These concepts have been distrusted by naturalists, as is both understandable and partly, but only partly, justified. In the early stages of a science a way has to be steered between two opposite dangers, on the one hand of theoretical ideas so simplified that they have no value in application, on the other hand of facts so disorganized that no coherent theme is apparent. The first danger is greatest when the ideas have been expressed mathematically, as has happened in population research” (1954, p. 3).
On Thompson’s views, see Kingsland (1995, pp. 166ff).
On Cole’s career as an ecologist, see Blomquist (2007).
This follows a smaller identical experiment in Scotland a few years earlier (Ramsden and Adams 2009).
Ramsden and Adams (2009) described the pathologies of the rats and how Calhoun’s notion of “behavioral sink,” used to label the shift in behavior due to overcrowding, resonated with contemporary social concerns about megacities in popular culture. They also note Calhoun’s lesser-known later work, devoted to the identification and understanding of the few individuals that manifested enhanced abilities to cope with overcrowding.
Calhoun and Wellington are among the few names cited by pioneer behavioral ecologist Krebs in his paper about Denis Chitty and the evolutionary viewpoint within ecology in the 1950s (Krebs 1995).
Gleason’s individualistic conception of communities is often claimed to have dismissed Clements’s superorganism. But on this precise issue, Gleason’s view of communities neglected individual differences within a species, to the extent that Janis Antonovics saw this concept as typically typological, and therefore not compatible with the Synthesis. “It seems that much ecological thinking is still a generation behind that of the systematist, in that it remains locked into a typological view of the species. The current ‘individualistic’ view of communities, first championed by Gleason, has embedded in it a view of a species that is remarkably typological: the ‘individualistic species’ is almost synonymous in usage with ‘typological species’” (Antonovics 1976, p. 239).
The Institute was near the Bureau of Animal Populations led by Elton, but the two groups apparently did not get along very well (Gay 2013).
“Lack originally considered the differences among the Galapagos finches to be a matter of random drift. Thus, he neatly reflects the changing attitude toward the importance of natural selection during the evolutionary synthesis” (Collins et al. 1986, p. 176).
With “both Darwin’s Finches and ‘the Significance of Clutch-Size’ in 1947, Lack inaugurated the application of the Modern Synthesis (the successful integration of Darwinian natural selection with Mendelian genetics that was forged in the 1930s and early 1940s) to the field of ecology” (Anderson 2013, p. vii).
“The reproductive rate of each species is a result of natural selection, and is not, as often supposed, adjusted to the mortality rate of the species; and the critical mortality factors are density-dependent, hence climate per se cannot be the primary factor controlling numbers” (Lack 1954, p. 8).
Once again, notice the shifting sense of mechanism versus pattern difference: now the whole density-dependence is the pattern and no longer the explanation.
Krebs and Davies (1995).
Chitty, a British-born ecologist, received his PhD at Oxford with Elton and then worked with him for 26 years at the Bureau of Animal Population before going to the University of British Columbia in 1960.
Chitty (1952, 1957, 1960).
“Voles probably exemplify a general law that all species are capable of limiting their own population densities without either destroying the food resources to which they are adapted, or depending upon enemies or climatic accidents to prevent them from doing so. If this is true, self-regulatory mechanisms have presumably been evolved through natural selection, and arguments in support of this view can certainly be advanced” (Chitty 1960, p. 111).
Although dated 1957, the volume did not appear until 1958. In 1959 another landmark work was the publication of an issue of Ibis on population regulation that included a paper by Wynne-Edwards and a paper on Lack, presenting opposed versions of the explanation of natural control based on natural selection (see Borelllo 2003). Here I focus on the Cold Spring Harbor Symposium, which mobilized many of the protagonists I consider here and presented a larger set of issues.
“Some of the factors influencing animal numbers, such as size, the length of the breeding season, or migration, are products of evolution, and their causes can be considered under two distinct heads. For example, a bird may be said to breed in spring because the longer days stimulate the growth of its sex organs, and also because it is only in spring that there is enough food for it to raise young. In this example, daylength is a proximate factor helping to bring the bird into breeding condition at a suitable season; but the suitability of the season depends on the food supply, which has been an ultimate factor in the evolution of the breeding season of the species. Ultimate factors are concerned with survival value, proximate factors with adaptations in physiology and behaviour. … An effective adaptation is often thus ‘anticipatory,’ but the anticipation is not, of course, conscious, nor a result of the immediate situation; it is a long-term product of evolution” (Lack 1954, p. 5).
Futuyama later made the same point: “In many of these fields [of functional ecology] the major questions were and are functional rather than historical in nature; evolution and history need not be invoked if we wish to know what immediate factors govern the course of succession, the rate of phosphorus turnover, or the distribution of a species, given its physiology” (1986, p. 306).
r- and K- selection are two modes of selection, distinguished by the ratio of offspring number and parental investment. r selection favors a strategy of leaving many offspring and not caring about them (e.g., laying thousands of fish eggs), while K selection favors the strategy of leaving a few mature offspring requiring substantial parental care (e.g., primates’ reproductive life). The environmental conditions, especially the frequency of predators, decide which selective regime dominates.
Lerner regularly corresponded with Dobzhansky (sometimes in Russian, their common native language). Dobzhansky relied on Lerner’s mathematical ability to support him, for example, in the controversy about balanced versus classical accounts of polymorphisms.
Actually, Park and colleagues continued to work on the experiment, integrating the genetic viewpoint and controlling strains; Leslie et al. (1968) still concludes that some cases are not explainable by competitive exclusion.
Mayr to Lerner, 2 January 1962, I. Michael Lerner Papers, American Philosophical Society, Philadelphia, PA.
Together with Eric Charnov, Gordon Orians in the 1970s authored a textbook on behavioral ecology of foraging that, although not published, was nonetheless circulated and was very influential.
Gay, referring to the important historical role of this piece, states: “Judging from the flurry of work following Hutchinson’s ‘concluding remarks’ it appears that he gave ecology a considerable heuristic boost” (2013, p. 121).
A hyperspace is a mathematical abstract space with a high number of dimensions (much higher than 3).
“The only conclusion that one can draw at present from the observations is that although animal communities appear qualitatively to be constructed as if competition were regulating their structure even in the best studied cases there are nearly always difficulties and unexplored possibilities. These difficulties suggest that if competition is determinative it either acts intermittently, or it is a more subtle process than has been supposed” (Hutchinson 1957, p. 419).
“Competing species seeking the same plant, prey or host, can coexist if their numbers are controlled by genetic feedback. For example, let assume that two aphid populations feed on sap from the same plant species. The two aphid species can coexist because the more abundant aphid species will eventually be controlled through the processes of genetic feedback” (p. 1436).
Kingsland speaks of a “shift away from descriptive or story-telling narratives to a more analytical, hypothesis-testing style, complete with attempts to mathematize the biological world” (1997, p. 417).
Hagen (1992, 96ff.).
A hypothetico-deductive method, drawing testable predictions from simple mathematical models, and identical to the method in population and quantitative genetics, is something MacArthur made pervasive in ecology through his influence. Fretwell notes that “prior to MacArthur’s 1957 paper on relative abundance, it had been little used in the study of natural history (about 5% of papers in biology from 1950 to 1956 tested predictions, compared to almost 50% nowadays)” (Fretwell 1975, p. 3).
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Acknowledgements
The author is grateful to the group of historians and philosophers committed since 2014 to the projet ‘Revisiting the Modern Synthesis’, especially Jean Gayon, Dick Burian, Edna Suarez and the audiences at the workshop organised by the group and the HSS conference in Chicago 2014. He warmly thanks Antoine Dussault, Sébastien Dutreuil, Jean Baptiste Grodwohl, John Huss, and Sophie Rousseau Mermans for their insightful comments and criticisms. He is greatly indebted to Chris Donohue for a thorough language checking, and a careful reading. He is extremely thankful to two anonymous reviewers whose criticisms helped to significantly improve the manuscript, as well as the JHB editors, whose remarks and copyediting crucially helped the paper. Je also thanks Jean-Baptiste Grodwohl for providing materials from the archives. This work has been supported by the GDR CNRS 3770 Sapienv, and the LIA CNRS Paris-Montréal ECIEB. The author and his paper are especially indebted to Jean Gayon.
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Huneman, P. How the Modern Synthesis Came to Ecology. J Hist Biol 52, 635–686 (2019). https://doi.org/10.1007/s10739-019-09570-9
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DOI: https://doi.org/10.1007/s10739-019-09570-9