This paper surveys questions about the nature of the Modern Synthesis as a historical event : was it rather theoretical than institutional? When and where did it actually happen? Who was involved? It argues that all answers to these questions are interrelated, and that systematic sets of answers define specific perspectives on the Modern Synthesis that are all complementary.

The article examines two cases of adoption of evolutionary ways of thinking by modern economists: Nelson and Winter’s (Evolutionary Theory of Economic Change, 1982), and evolutionary game theory (1990s and after). In both cases, the authors explicitly refer to natural selection in an economic context. I show that natural selection is taken in two different senses, which correspond to two general conceptions of the principle of natural selection, one of which contains reproduction and heredity as key elements, whereas the other (...) does not. (shrink)

L.A. Paul has recently defended the methodology of metaphysics on the grounds that it is continuous with the sciences. She claims that both scientists and metaphysicians use inference to the best explanation to choose between competing theories, and that the success of science vindicates the use of IBE in metaphysics. Specifically, the success of science shows that the theoretical virtues are truth-conducive. I challenge Paul’s claims on two grounds. First, I argue that, at least in biology, scientists adhere to the (...) vera causa ideal, which allows the theoretical virtues to play a much more limited role in scientific reasoning than Paul requires for metaphysical reasoning. The success of biology thus does not vindicate the methodology of metaphysics. Second, I argue that, at least in many cases, the successful reliance on the theoretical virtues in scientific contexts shows only that the theoretical virtues are truth-conducive within those local contexts, and not that they are truth-conducive generally. The upshots are that Paul’s defense of the methodology of metaphysics fails, and that any attempt to rescue her defense must pay more careful attention to what precisely is vindicated by successful science. (shrink)

This paper questions the form and prospects of “extended theories” which have been simultaneously and independently advocated both in the philosophy of mind and in the philosophy of biology. It focuses on Extend Mind Theory (EMT) and Developmental Systems Theory (DST). It shows first that the two theories vindicate a parallel extension of received views, the former concerning extending cognition beyond the brain, the latter concerned with extending evolution and development beyond the genes. It also shows that both arguments rely (...) on the demonstration of causal parities, which have been undermined by the classical received view. Then I question whether the argument that there is an illegitimate inference from parities or coupling to constitution claims, which has been objected by Adams and Aizawa in The bounds of cognition, (2008) to EMT, also holds against DST. To this aim, I consider two defenses against DST that are parallel to two defenses against EMT, one about intrinsic content, the other about the difference between what’s in principle possible and what happens in practice. I conclude by claiming that the weaknesses and strengths of both theories are different regarding these two kinds of objections. (shrink)

During the 1920s and 1930s, many biologists questioned the viability of Darwin’s theory as a mechanism of evolutionary change. In the early 1940s, and only after a number of alternatives were suggested, Darwinists succeeded to establish natural selection and gene mutation as the main evolutionary mechanisms. While that move, today known as the neo-Darwinian synthesis, is taken as signalling a triumph of evolutionary theory, certain critical problems in evolution—in particular the evolution of animal function—could not be addressed with this approach. (...) Here I demonstrate this through reconstruction of the evolutionary theory of Joseph Needham (1900–1995), who pioneered the biochemical study of evolution and development. In order to address such problems, Needham employed Herbert Spencer’s principles of emergence and Ernst Haeckel’s theory of recapitulation. While Needham did not reject Darwinian theory, Spencerian and Haeckelian frameworks happened to better fit his findings and their evolutionary relevance. He believed selectionist and genetic approaches to be important but far from sufficient for explaining how evolutionary transformations occur. (shrink)

Theoreticians that defend a form of realism regarding natural kinds minimally entertain the belief that the world features divisions into kinds and that the natural kind concept is a useful tool for philosophy of science. The objective of this paper is to challenge these assumptions. First, we challenge realism toward natural kinds by showing that the main arguments for their existence, which rely on the epistemic success of natural kinds, are unsatisfactory. Second, we show that, whether they exist or not, (...) natural kinds are expendable when it comes to describing and analyzing scientific explanations accurately. (shrink)

The aim of this paper is to explain why, while Charles Darwin was well recognized as a scientific leader of his time, Claude Bernard never really regarded Darwinism as a scientific theory. The lukewarm reception of Darwin at the Académie des Sciences of Paris and his nomination to a chair only after 8 years contrasts with his prominence, and Bernard’s attitude towards Darwin’s theory of species evolution belongs to this French context. Yet we argue that Bernard rejects the scientific value (...) of Darwinian principles mainly for epistemological reasons. Like Darwin, Bernard was interested in hereditary processes, and planned to conduct experiments on these processes that could lead to species transformation. But the potential creation of new forms of life would not vindicate Darwinism since biologists can only explain the origin of morphotypes and morphological laws by the means of untestable analogies. Because it can be the object neither of experiments nor of any empirical observation, phylogeny remains out of science’s scope. Around 1878 Bernard foresaw a new general physiology based on the study of protoplasm, which he saw as the agent of all basic living phenomena. We will analyze why Bernard regarded Darwinism as part of metaphysics, yet still referred to Darwinians in his latter works in 1878. Basically, the absence of a scientific reception of Darwinism in Bernard's work should not obscure its philosophical reception, which highlights the main principles of Bernard’s epistemology. (shrink)

There is hardly anything more central to our universe than conservation. Many scientific fields and disciplines view the law of conservation as one of the most fundamental universal laws. The Darwinian model pivots the process of evolution on variability, reproduction, and natural selection. Conservation plays a marginal role in this model and is not really universal, as the model allows exceptions to conservation, i.e. non-conservation, to play an equally important role in evolution. This anomalous role of conservation in the Darwinian (...) model raises questions: What is the reason for this anomaly? Is conservation really universal, as we tend to believe or is it not, as the Darwinian model suggests? This contribution proposes a new model of evolution that focuses on levels of organization, rather than of species, organisms, or populations. It argues that conservation is central to evolution. Not only does this new model restores the universal status of conservation but it also makes possible to resolve some outstanding problems and controversies that continue to plague the Darwinian model. The article tries to advance the broad Darwinian project that seeks to explain the process of evolution as a product of the spontaneous processes in nature. (shrink)

In this commentary of Koonin’s target paper, we defend an extended view of CRISPR-Cas immunity by arguing that CRISPR-Cas includes, but cannot be reduced to, defence against nonself. CRISPR-Cas systems can target endogenous elements and tolerate exogenous elements. We conclude that the vocabulary of “defence” and “nonself” might be misleading when describing CRISPR-Cas systems.

This is the first of a two-part essay on the history of debates concerning the creativity of natural selection, from Darwin through the evolutionary synthesis and up to the present. Here I focus on the mid-late nineteenth century to the early twentieth, with special emphasis on early Darwinism and its critics, the self-styled “mutationists.” The second part focuses on the evolutionary synthesis and some of its critics, especially the “neutralists” and “neo-mutationists.” Like Stephen Gould, I consider the creativity of natural (...) selection to be a key component of what has traditionally counted as “Darwinism.” I argue that the creativity of natural selection is best understood in terms of selection initiating evolutionary change, and selection being responsible for the presence of the variation it acts upon, for example by directing the course of variation. I consider the respects in which both of these claims sound non-Darwinian, even though they have long been understood by supporters and critics alike to be virtually constitutive of Darwinism. (shrink)

This is the second of a two-part essay on the history of debates concerning the creativity of natural selection, from Darwin through the evolutionary synthesis and up to the present. In the first part, I focussed on the mid-late nineteenth century to the early twentieth, with special emphasis on early Darwinism and its critics, the self-styled “mutationists.” The second part focuses on the evolutionary synthesis and some of its critics, especially the “neutralists” and “neo-mutationists.” Like Stephen Gould, I consider the (...) creativity of natural selection to be a key component of what has traditionally counted as “Darwinism.” I argue that the creativity of natural selection is best understood in terms of selection initiating evolutionary change, and selection directing evolutionary change, for example by creating the variation that it subsequently acts upon. I consider the respects in which both of these claims sound non-Darwinian, even though they have long been understood by supporters and critics alike to be virtually constitutive of Darwinism. (shrink)

During the 1920s and 1930s, many biologists questioned the viability of Darwin’s theory as a mechanism of evolutionary change. In the early 1940s, and only after a number of alternatives were suggested, Darwinists succeeded to establish natural selection and gene mutation as the main evolutionary mechanisms. While that move, today known as the neo-Darwinian synthesis, is taken as signalling a triumph of evolutionary theory, certain critical problems in evolution—in particular the evolution of animal function—could not be addressed with this approach. (...) Here I demonstrate this through reconstruction of the evolutionary theory of Joseph Needham (1900–1995), who pioneered the biochemical study of evolution and development. In order to address such problems, Needham employed Herbert Spencer’s principles of emergence and Ernst Haeckel’s theory of recapitulation. While Needham did not reject Darwinian theory, Spencerian and Haeckelian frameworks happened to better fit his findings and their evolutionary relevance. He believed selectionist and genetic approaches to be important but far from sufficient for explaining how evolutionary transformations occur. (shrink)

An important historical relation that has hardly been addressed is the influence of Prosper Lucas's Treatise on Natural Inheritance on the development of Charles Darwin's concepts related to inheritance. In this article we trace this historical connection. Darwin read Lucas's Treatise in 1856. His reading coincided with many changes concerning his prior ideas on the transmission and expression of characters. We consider that this reading led him to propose a group of principles regarding prepotency, hereditary diseases, morbid tendencies and atavism; (...) following Lucas, he called these principles: laws of inheritance. (shrink)

Besides mechanistic explanations of phenomena, which have been seriously investigated in the last decade, biology and ecology also include explanations that pinpoint specific mathematical properties as explanatory of the explanandum under focus. Among these structural explanations, one finds topological explanations, and recent science pervasively relies on them. This reliance is especially due to the necessity to model large sets of data with no practical possibility to track the proper activities of all the numerous entities. The paper first defines topological explanations (...) and then explains why topological explanations and mechanisms are different in principle. Then it shows that they are pervasive both in the study of networks—whose importance has been increasingly acknowledged at each level of the biological hierarchy—and in contexts where the notion of selective neutrality is crucial; this allows me to capture the difference between mechanisms and topological explanations in terms of practical modelling practices. The rest of the paper investigates how in practice mechanisms and topologies are combined. They may be articulated in theoretical structures and explanatory strategies, first through a relation of constraint, second in interlevel theories, or they may condition each other. Finally, I explore how a particular model can integrate mechanistic informations, by focusing on the recent practice of merging networks in ecology and its consequences upon multiscale modelling. (shrink)

I argue that the propensity interpretation of fitness, properly understood, not only solves the explanatory circularity problem and the mismatch problem, but can also withstand the Pandora’s box full of problems that have been thrown at it. Fitness is the propensity (i.e., probabilistic ability, based on heritable physical traits) for organisms or types of organisms to survive and reproduce in particular environments and in particular populations for a specified number of generations; if greater than one generation, “reproduction” includes descendants of (...) descendants. Fitness values can be described in terms of distributions of propensities to produce varying number of offspring and can be modeled for any number of generations using computer simulations, thus providing both predictive power and a means for comparing the fitness of different phenotypes. Fitness is a causal concept, most notably at the population level, where fitness differences are causally responsible for differences in reproductive success. Relative fitness is ultimately what matters for natural selection. (shrink)

Biologists and philosophers have been extremely pessimistic about the possibility of demonstrating random drift in nature, particularly when it comes to distinguishing random drift from natural selection. However, examination of a historical case-Maxime Lamotte's study of natural populations of the land snail, Cepaea nemoralis in the 1950s - shows that while some pessimism is warranted, it has been overstated. Indeed, by describing a unique signature for drift and showing that this signature obtained in the populations under study, Lamotte was able (...) to make a good case for a significant role for drift. It may be difficult to disentangle the causes of drift and selection acting in a population, but it is not impossible. (shrink)

This paper attempts to reconcile the conceptual conflict between typological and population thinking to provide a philosophical foundation for extended evolutionary synthesis. Typological thinking has been considered a pre-Darwinian, essentialist dogma incompatible with population thinking, which is the core notion of Darwinism. More recent philosophical and historical studies suggest that a non-essentialist form of typology has some advantages in the study of evolutionary biology. However, even if we adopt such an epistemological interpretation of typological thinking, there still remains an epistemological (...) and methodological conflict between these two styles of thinking. How can we relate typological thinking with population thinking in pursuit of more integrated or interconnected research into evolutionary biology? I propose that homology thinking, which is another style of thinking that recognizes homologous characters, provides a common basis for typological representations of character states and for character dynamics in an evolving population. Good examples of this bridging role are found in teratology and breeding, where variation and novelty are recognized in developmental and morphological traits, gene expression patterns, and so on. Essentialism-free, dynamic views of homology have great potential to reconcile typological and population thinking and to set the stage for the EES. (shrink)

In June 1909, scientists and dignitaries from 167 different countries gathered in Cambridge to celebrate the hundredth anniversary of Charles Darwin’s birth and the fiftieth anniversary of the publication of Origin of Species. The event was one of the most magnificent commemorations in the annals of science. Delegates gathered within the cloisters of Cambridge University not only to honor the “hero” of evolution but also to reassess the underpinnings of Darwinism at a critical juncture. With the mechanism of natural selection (...) increasingly under attack, evolutionary theory was in disarray. Against this backdrop, biologists weighed the impact of several new developments—the rediscovery of Mendel’s laws of heredity, de Vriesian mutation theory, and the linkage of sex‐cell division to the mechanism of heredity. The 1909 Darwin celebration thus represents a significant watershed in the history of modern biology that allows historians to assess the status of evolution prior to the advent of the chromosome theory of genetics. (shrink)

In the wake of recent attempts at alternate history (Bowler 2013), this paper suggests several avenues for a pluralistic approach to Charles Darwin and his role in the history of evolutionary theory. We examine in what sense Darwin could be described as a major driver of theoretical change in the history of biology. First, this paper examines how Darwin influenced the future of biological science: not merely by stating the fact of evolution or by bringing evidence for it; but by (...) discovering natural selection, and giving it pre-eminence over any other mechanism for evolution; and also by proposing a masterful and quite unique synthesis of many scientific fields. Contrasting Darwin’s views with those of A.R. Wallace, I conclude that “natural selection” is clearly an original contribution, that it had no forerunners or co-discoverers, and could barely have appeared after Darwin conceived of it. This specificity of Darwin’s contribution is an invitation to be strongly presentist (Loison 2016) and to adopt only weak counter-factuals. In contrast, there are possible ways to use strong counter-factuals as attempts to “pluralize” the history of biological theory: i.e. imagine new possible avenues for the development of evolutionary biology. The idea that evolution was a theory “in the air” suggests that evolutionary theory could have developed in a world without Darwin, especially if we accept to delete not only “Darwin” but “England”. France and Germany are examined as possible countries where evolutionary ideas would have thrived even with no contribution from the English scientists. Finally, the paper suggests another counter-factual hypothesis: deleting not Darwin and his Origin but the Darwin Industry itself. This may allow us to read the Origin of Species with fresh eyes and to discover Darwin’s life-long interest in variation and its laws, as many of his early readers did. (shrink)

Chance has been a focus of attention ever since the beginning of population genetics, but neutrality has not, as natural selection once appeared to be the only worthwhile issue. Neutral change became a major source of interest during the neutralist–selectionist debate, 1970–1980. It retained interest beyond this period for two reasons that contributed to its becoming foundational for evolutionary reasoning. On the one hand, neutral evolution was the first mathematical prediction to emerge from Mendelian inheritance: until then evolution by natural (...) selection was considered the alternative to the fixity of species; now it appears to be the alternative to continuous change. Second, neutral change generated a set of clear predictions on standing variation. These could be used as a reference for detecting more elusive alternative mechanisms of evolution including natural selection. In the wake of the transition from Mendelism to genomics, the combination of coalescent theory, DNA sequence variation, and numerical analysis made it possible to integrate contingent aspects of the history of species into a new null model, thus opening a new dimension in the concept of population that the Modern Synthesis formerly considered as a mere gene pool. (shrink)

Prior to August Weismann’s 1889 germ-plasm theory, social reformers believed that humans could inherit the effects of a salubrious environment and, by passing environmentally-induced modifications to their offspring, achieve continuous progress. Weismann’s theory disrupted this logic and caused many to fear that they had little control over human development. As numerous historians have observed, this contributed to the birth of the eugenics movement. However, through an examination of the work of social scientists Lester Frank Ward, Richard T. Ely, Amos Griswold (...) Warner, James Mark Baldwin, Simon Nelson Patten, Alfred Kroeber, Walter Robinson Smith, and Luther Lee Bernard, I demonstrate that Weismann’s ideas also prompted scholars to create of theories of human progress in which the social environment had a central role and biological heredity had a diminished one. Furthermore, in creating a new theory of social progress based on a concept called “social heredity,” the thinkers surveyed in this article separated biological and social thought and asserted the independence of the American social sciences. I argue that this represents an important moment in the maturation of the human sciences, and I suggest that the germ-plasm theory of heredity deserves a larger place in histories of the development of the American social scientific disciplines. (shrink)

In the advertising discourse of human genetic database projects, of genetic ancestry tracing companies, and in popular books on anthropological genetics, what I refer to as the anthropological gene and genome appear as documents of human history, by far surpassing the written record and oral history in scope and accuracy as archives of our past. How did macromolecules become "documents of human evolutionary history"? Historically, molecular anthropology, a term introduced by Emile Zuckerkandl in 1962 to characterize the study of primate (...) phylogeny and human evolution on the molecular level, asserted its claim to the privilege of interpretation regarding hominoid, hominid, and human phylogeny and evolution vis-à-vis other historical sciences such as evolutionary biology, physical anthropology, and paleoanthropology. This process will be discussed on the basis of three key conferences on primate classification and evolution that brought together exponents of the respective fields and that were held in approximately ten-years intervals between the early 1960s and the 1980s. I show how the anthropological gene and genome gained their status as the most fundamental, clean, and direct records of historical information, and how the prioritizing of these epistemic objects was part of a complex involving the objectivity of numbers, logic, and mathematics, the objectivity of machines and instruments, and the objectivity seen to reside in the epistemic objects themselves. (shrink)

Work throughout the history and philosophy of biology frequently employs ‘chance’, ‘unpredictability’, ‘probability’, and many similar terms. One common way of understanding how these concepts were introduced in evolution focuses on two central issues: the first use of statistical methods in evolution (Galton), and the first use of the concept of “objective chance” in evolution (Wright). I argue that while this approach has merit, it fails to fully capture interesting philosophical reflections on the role of chance expounded by two of (...) Galton's students, Karl Pearson and W.F.R. Weldon. Considering a question more familiar from contemporary philosophy of biology—the relationship between our statistical theories of evolution and the processes in the world those theories describe—is, I claim, a more fruitful way to approach both these two historical actors and the broader development of chance in evolution. (shrink)

This paper considers recent heated debates led by Jerry A. Coyne andMichael J. Wade on issues stemming from the 1929–1962 R.A. Fisher-Sewall Wrightcontroversy in population genetics. William B. Provine once remarked that theFisher-Wright controversy is central, fundamental, and very influential.Indeed,it is also persistent. The argumentative structure of therecent (1997–2000) debates is analyzed with the aim of eliminating a logicalconflict in them, viz., that the two sides in the debates havedifferent aims and that, as such, they are talking past each other. (...) Given aphilosophical analysis of the argumentative structure of the debates,suggestions supportive of Wade's work on the debate are made that areaimed, modestly, at putting the persistent Fisher-Wright controversy on thecourse to resolution. (shrink)

: This paper explores the calibration of laboratory models in population genetics as an experimental strategy for justifying experimental results and claims based upon them following Franklin (1986, 1990) and Rudge (1996, 1998). The analysis provided undermines Coyne et al.'s (1997) critique of Wade and Goodnight's (1991) experimental study of Wright's (1931, 1932) Shifting Balance Theory. The essay concludes by further demonstrating how this analysis bears on Diamond's (1986) claims regarding the weakness of laboratory experiments as evidence, and further how (...) the calibration strategy fits within Lloyd's (1987, 1988) account of the confirmation of ecological and evolutionary models. (shrink)

Julian Huxley’s eclipse of Darwinism narrative has cast a long shadow over the historiography of evolutionary theory around the turn of the nineteenth century. It has done so by limiting who could be thought of as Darwinian. Peter Bowler used the eclipse to draw attention to previously understudied alternatives to Darwinism, but maintained the same flaw. In his research on the Non-Darwinian Revolution, he extended this problematic element even further back in time. This paper explores how late nineteenth-century neo-Darwinian conceptualizations (...) of Darwinism were later utilized by several advocates and detractors of the Modern Synthesis. John Beatty has shown how this continuity hinges at least partly on the perceived importance of the creativity of natural selection. The paper provides a more thorough look at Darwin’s two conflicting accounts of variation, ascribed to struggles in explaining quantitative versus qualitative characters. In doing so, it suggests that other forms of Darwinism persisted, in both the non-Darwinian revolution and eclipse periods, because of tension between contingency and creativity in Darwin’s own work. This tension is traced out from Darwin’s conceptions of variation into the work of Alfred Russel Wallace, Hugo de Vries, and Thomas Henry Huxley. Based on this, the eclipse narrative is criticized for not considering the meaning of Darwinism in different geographical locations. Britain and the United States showed few signs of an eclipse. Rather, the rise of German debates about Haeckel’s vision of Darwinism have been mistaken for a universal decline in support. (shrink)

Causalism is the thesis that natural selection can cause evolution. A standard argument for causalism involves showing that a hypothetical intervention on some population-level property that is identified with natural selection will result in evolution. In a pair of articles, one of which recently appeared in the pages of this journal, Jun Otsuka has put forward a quite different argument for causalism. Otsuka attempts to show that natural selection can cause evolution by considering a hypothetical intervention on an individual-level property. (...) Specifically, Otsuka identifies natural selection with the causal relationship between a trait and fitness, claims an intervention on the strength of this relationship can cause evolution, then concludes that natural selection can cause evolution. Below I describe why Otsuka’s argument for causalism is unconvincing. Central to my criticism is that Otsuka’s argument works only if one adopts an indefensible account of natural selection, according to which natural selection can occur in the absence of trait or fitness variation. I go on to explain why any attempt to demonstrate the truth of causalism via a hypothetical intervention on an individual-level property would appear to require one to adopt an account of natural selection that is inadequate for the same reason. This in turn means the plausibility of causalism does indeed depend on the plausibility of the claim that population-level properties, which supervene on the properties of the individuals in the population, can be causally efficacious. (shrink)

The analogy between artificial selection of domestic varieties and natural selection in nature was a vital element of Darwin's argument in his Origin of Species. Ever since, the image of breeders creating new varieties by artificial selection has served as a convincing illustration of how the theory works. In this paper I argue that we need to reconsider our understanding of Darwin's analogy. Contrary to what is often assumed, nineteenth-century animal breeding practices constituted a highly controversial field that was fraught (...) with difficulties. It was only with considerable effort that Darwin forged his analogy, and he only succeeded by downplaying the importance of two other breeding techniques - crossing of varieties and inbreeding - that many breeders deemed essential to obtain new varieties. Part of the explanation for Darwin's gloss on breeding practices, I shall argue, was that the methods of his main informants, the breeders of fancy pigeons, were not representative of what went on in the breeding world at large. Darwin seems to have been eager to take the pigeon fanciers at their word, however, as it was only their methods that provided him with the perfect analogy with natural selection. Thus while his studies of domestic varieties were important for the development of the concept of natural selection, the reverse was also true: Darwin's comprehension of breeding practices was moulded by his understanding of the working of natural selection in nature. Historical studies of domestic breeding practices in the eighteenth and nineteenth century confirm that, besides selection, the techniques of inbreeding and crossing were much more important than Darwin's interpretation allowed for. And they still are today. This calls for a reconsideration of the pedagogic use of Darwin's analogy too. (shrink)

The paper treats several ontological questions about certain nineteenth-century and contemporary medical and scientific conceptualizations of hereditary relation. In particular, it considers the account of mid-nineteenth century psychiatric thought given by Foucault in Psychiatric Power: Lectures at the Collège de France, 1973–1974 and Abnormal: Lectures at the Collège de France, 1974–1975 . There, Foucault argues that a fantastical conceptual prop, the ‘metabody,’ as he terms it, was implicitly supposed by that period’s psychiatric medicine as a putative ground for psychiatric pathology. (...) After presenting the heart of Foucault’s thought on the ‘metabody,’ the paper investigates the possibility that a contemporary version of a ‘metabody’ may operate today as a conceptual analog of the nineteenth-century psychiatric theory and practice that Foucault began to expose in the texts examined here. It speculates that we might identify a contemporary genetic version of a ‘metabody’ in a particular current conception of the gene as replicator, an item marked by an ambiguous temporal ontology. (shrink)

No single author presented Darwin with a more difficult question about his priority in discovering natural selection than the British comparative anatomist and paleontologist Richard Owen. Owen was arguably the most influential biologist in Great Britain in Darwin’s time. Darwin wanted his approbation for what he believed to be his own theory of natural selection. Unfortunately for Darwin, when Owen first commented in publication about Darwin’s theory of descent he was openly hostile. Darwin was taken off-guard. In private meetings and (...) correspondence prior to 1860 Owen had been nothing but polite and friendly, even helping Darwin in cataloguing and analyzing Darwin’s zoological specimens from the Beagle voyage. Every early indication predicted a life-long friendship and collaboration. But that was not to be. Owen followed his slashing review with a mounting campaign in the 1860s to denounce and discredit both Darwin and his small but ascendant circle of friends and supporters. But that was not enough for Owen. Starting in 1866, perhaps by now realizing Darwin had landed the big fish, Owen launched a new campaign, to claim the discovery of “Darwin’s theory” for himself. Darwin naturally fought back, mainly in the “Historical Sketch” that he prefaced to Origin starting in 1861. But when we peel back the layers of personal animus and escalating vituperation we discover in fact their quarrel was generated more by mutual misunderstanding than scientific disagreement. The battle ended only when Darwin finally penetrated to the crux of the matter and put an end to the rivalry in 1872, in the final version of the Sketch. (shrink)

According to a classical narrative, early geneticists, failing to see how Mendelism provides the missing pieces of Darwin’s theory, rejected gradual changes and advocated an implausible yet briefly popular view of evolution-by-mutation; after decades of delay (in which synthesis was prevented by personal conflicts, disciplinary rivalries, and anti-Darwinian animus), Darwinism emerged on a new Mendelian basis. Based on the works of four influential early geneticists – Bateson, de Vries, Morgan and Punnett –, and drawing on recent scholarship, we offer an (...) alternative that turns the classical view on its head. For early geneticists, embracing discrete inheritance and the mutation theory (for the origin of hereditary variation) did not entail rejection of selection, but rejection of Darwin’s non-Mendelian views of heredity and variation, his doctrine of natura non facit saltum, and his conception of “natural selection” as a creative force that shapes features out of masses of infinitesimal differences. We find no evidence of a delay in synthesizing mutation, rules of discrete inheritance, and selection in a Mendelian-Mutationist Synthesis. Instead, before 1918, early geneticists had conceptualized allelic selection, the Hardy–Weinberg equilibrium, the evolution of a quantitative trait under selection, the probability of fixation of a new mutation, and other key innovations. Contemporary evolutionary thinking seems closer to their more ecumenical view than to the restrictive mid-twentieth-century consensus known as the Modern Synthesis. (shrink)

There are ongoing debates in philosophy of biology about what falls within natural selection's explanatory scope. These include debates about whether selection can explain individual-level traits, the extent to which selection can explain distributions of trait frequencies, and whether selection can explain the origin of novel traits. Here I'll survey these debates, suggest which views seem most plausible, and describe some useful conceptual frameworks for thinking about the issues involved.

Textbook descriptions of the foundations of Genetics give the impression that besides Mendel’s no other research on heredity took place during the nineteenth century. However, the publication of the Origin of Species in 1859, and the criticism that it received, placed the study of heredity at the centre of biological thought. Consequently, Herbert Spencer, Charles Darwin himself, Francis Galton, William Keith Brooks, Carl von Nägeli, August Weismann, and Hugo de Vries attempted to develop theories of heredity under an evolutionary perspective, (...) and they were all influenced by each other in various ways. Nonetheless, only Nägeli became aware of Mendel’s experimental work; it has also been questioned whether Mendel even had the intention to develop a theory of heredity. In this article, a short presentation of these theories is made, based on the original writings. The major aim of this article is to suggest that Mendel was definitely not the only one studying heredity before 1900, if he even did this, as may be inferred by textbooks. Although his work had a major impact after 1900, it had no impact during the latter half of the nineteenth century when an active community of students of heredity emerged. Thus, textbooks should not only present the work of Mendel, but also provide a wider view of the actual history and a depiction of science as a social process. (shrink)

Recently, much philosophical discussion has centered on the best way to characterize the concepts of random drift and natural selection, and, in particular, whether selection and drift can be conceptually distinguished (Beatty, 1984; Brandon, 2005; Hodge, 1983, 1987; Millstein, 2002, 2005; Pfeifer, 2005; Shanahan, 1992; Stephens, 2004). These authors all contend, to a greater or lesser degree, that their concepts make sense of biological practice. So it should be instructive to see how the concepts of drift and selection were distinguished (...) by the disputants in a high-profile debate; debates such as these often force biologists to take a more philosophical turn, discussing the concepts at issue in greater detail than usual. Moreover, it is important to consider a debate where the disputants are actually trying to apply the models of population genetics to natural populations; only then can their proper interpretations become fully apparent. (Indeed, I contend that some of the philosophical confusion has arisen because authors have considered only the models themselves, and not the phenomena that the models are attempting to represent). A prime candidate for just such a case study is what Provine (1986) has termed “The Great Snail Debate,” that is, the debates over the highly polymorphic land snails Cepaea nemoralis and C. hortensis in the 1950s and early 1960s. These studies represent one of the best, if not the best, of the early attempts to demonstrate drift in natural populations. (shrink)