Change is the fundamental idea of evolution. Explaining the extraordinary biological change we see written in the history of genomes and fossil beds is the primary occupation of the evolutionary biologist. Yet it is a surprising fact that for the majority of evolutionary research, we have rarely studied how evolution typically unfolds in nature, in changing ecological environments, over space and time. While ecology played a major role in the eventual acceptance of the population genetic viewpoint of evolution in the (...) synthetic era (circa 1918-1956), it held a lesser role in the development of evolutionary theory until the 1980s, when we began to systematically study the evolutionary dynamics of natural populations in space and time. As a result, early evolutionary theory was initially constructed in an abstract vacuum that was unrepresentative of evolution in nature. The subtle synthesis between ecology with evolutionary biology (eco-evo synthesis) over the past 40 years has progressed our knowledge of natural selection dynamics as they are found in nature, thus revealing how natural selection varies in strength, direction, form, and, more surprisingly, level of biological organization. Natural selection can no longer be reduced to lower levels of biological organization (i.e., individuals, selfish genes) over shorter timescales but should be expanded to include adaptation at higher levels and over longer timescales. Long-term and/or emergent evolutionary phenomena, such as multilevel selection or evolvability, have thus become tenable concepts within an evolutionary biology that embraces ecology and spatiotemporal change. Evolutionary biology is currently suspended at an intermediate stage of scientific progress that calls for the organization of all the recent knowledge revealed by the eco-evo synthesis into a coherent and unified theoretical framework. This is where philosophers of biology can be of particular use, acting as a bridge between the subdisciplines of biology and inventing new theoretical strategies to organize and accommodate the recent knowledge. Philosophers have recommended transitioning away from outdated philosophies that were originally derived from physics within the philosophical zeitgeist of logical positivism (i.e., monism, reductionism, and monocausation) and toward a distinct philosophy of biology that can capture the natural complexity of multifaceted biological systems within diverse ecosystems—one that embraces the emerging philosophies of pluralism, emergence, and multicausality. Therefore, I see recent advances in ecology, evolutionary biology, and the philosophy of biology as laying the groundwork for another major biological synthesis, what I refer to as the Second Synthesis because, in many respects, it is analogous to the aims and outcomes of the first major biological synthesis (but is notably distinct from the inorganic and contrived progressive movement known as the extended evolutionary synthesis). With the general development of a distinctive philosophy of science, biology has rightfully emerged as an autonomous science. Thus, while the first synthesis legitimized biology, the Second Synthesis autonomized biology and afforded biology its own philosophy, allowing biology to finally realize its full scientific potential. (shrink)

August Weismann is famous for having argued against the inheritance of acquired characters. However, an analysis of his work indicates that Weismann always held that changes in external conditions, acting during development, were the necessary causes of variation in the hereditary material. For much of his career he held that acquired germ-plasm variation was inherited. An irony, which is in tension with much of the standard twentieth-century history of biology, thus exists – Weismann was not a Weismannian. I distinguish three (...) claims regarding the germ-plasm: (1) its continuity, (2) its morphological sequestration, and (3) its variational sequestration. With respect to changes in Weismann’s views on the cause of variation, I divide his career into four stages. For each stage I analyze his beliefs on the relative importance of changes in external conditions and sexual reproduction as causes of variation in the hereditary material. Weismann believed, and Weismannism denies, that variation, heredity, and development were deeply intertwined processes. This article is part of a larger project comparing commitments regarding variation during the latter half of the nineteenth century. (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)

In earlier work I have argued that the most substantial threat to scientific realism arises from the problem of unconceived alternatives: the repeated failure of past scientists and scientific communities to conceive of alternatives to extant scientific theories, even when such alternatives were both (1) well confirmed by the evidence available at the time and (2) sufficiently scientifically serious as to be later embraced by actual scientific communities. In this paper I explore Charles Darwin's development and defense of his 'pangenesis' (...) theory of inheritance and conclude that this particular historical example offers impressive support for the challenge posed to realism by this problem of unconceived alternatives. (shrink)

Ecological Psychology and Enactivism both challenge representationist cognitive science, but the two approaches have only begun to engage in dialogue. Further conceptual clarification is required in which differences are as important as common ground. This paper enters the dialogue by focusing on important differences. After a brief account of the parallel histories of Ecological Psychology and Enactivism, we cover incompatibility between them regarding their theories of sensation and perception. First, we show how and why in ecological theory perception is, crucially, (...) not based on sensation. We elucidate this idea by examining the biological roots of work in the two fields, concentrating on Gibson and Varela and Maturana. We expound an ecological critique of any sensation-based approach to perception by detailing two topics: classic retinal image theories and perception in single-celled organisms. The second main point emphasizes the importance of the idea of organism-environment mutuality and its difference from structural coupling of sensations and motor behavior. We point out how ecological - phenomenological methods of inquiry grow out of mutualism and compare Gibson’s idea of visual kinesthesis to Merleau-Ponty’s idea of the lived body. Third, we conclude that Ecological Psychology and varieties of Enactivism are laying down different paths to pursue related goals. Thus, convergence of Ecological Psychology and Enactivism is not possible given their conflicting assumptions, but cross-fertilization is possible and desirable. (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)

Recent discoveries of geographical patterns in microbial distribution are undermining microbiology’s exclusively ecological explanations of biogeography and their fundamental assumption that ‘everything is everywhere: but the environment selects’. This statement was generally promulgated by Dutch microbiologist Martinus Wilhelm Beijerinck early in the twentieth century and specifically articulated in 1934 by his compatriot, Lourens G. M. Baas Becking. The persistence of this precept throughout twentieth-century microbiology raises a number of issues in relation to its formulation and widespread acceptance. This paper will (...) trace the conceptual history of Beijerinck’s claim that ‘everything is everywhere’ in relation to a more general account of its theoretical, experimental and institutional context. His principle also needs to be situated in relationship to plant and animal biogeography, which, this paper will argue, forms a continuum of thought with microbial biogeography. Finally, a brief overview of the contemporary microbiological research challenging ‘everything is everywhere’ reveals that philosophical issues from Beijerinck’s era of microbiology still provoke intense discussion in twenty-first century investigations of microbial biogeography. (shrink)

The cell is not only the structural, physiological, and developmental unit of life, but also the reproductive one. So far, however, this aspect of the cell has received little attention from historians and philosophers of biology. I will argue that cell theory had far-reaching consequences for how biologists conceptualized the reproductive relationships between germs and adult organisms. Cell theory, as formulated by Theodor Schwann in 1839, implied that this relationship was a specific and lawful one, that is, that germs of (...) a certain kind, all else being equal, would produce adult organisms of the same kind, and vice versa. Questions of preformation and epigenesis took on a new meaning under this presupposition. The question then became one of whether cells could be considered as autonomous agents producing adult organisms of a given species, or whether they were the product of external, organizing forces and thus only a stage in the development of the whole organism. This question became an important issue for nineteenth-century biology. As I will demonstrate, it was the view of cells as autonomous agents which helped both Charles Darwin and Gregor Mendel to think of inheritance as a lawful process. (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)

In this paper, I analyze the disruptive impact of Darwinian selectionism for the century-long tradition in which the environment had a direct causative role in shaping an organism’s traits. In the case of humans, the surrounding environment often determined not only the physical, but also the mental and moral features of individuals and whole populations. With its apparatus of indirect effects, random variations, and a much less harmonious view of nature and adaptation, Darwinian selectionism severed the deep imbrication of organism (...) and milieu posited by these traditional environmentalist models. This move had radical implications well beyond strictly biological debates. In my essay, I discuss the problematization of the moral idiom of environmentalism by William James and August Weismann who adopted a selectionist view of the development of mental faculties. These debates show the complex moral discourse associated with the environmentalist-selectionist dilemma. They also well illustrate how the moral reverberations of selectionism went well beyond the stereotyped associations with biological fatalism or passivity of the organism. Rereading them today may be helpful as a genealogical guide to the complex ethical quandaries unfolding in the current postgenomic scenario in which a revival of new environmentalist themes is taking place. (shrink)

It is well known that William James’s thinking was influenced by evolutionary theory and by Darwin’s theory of natural selection in particular. It is easy to misunderstand James’s evolutionary thinking, however, if one is tempted to read contemporary evolutionary views back into James. In this article I try to avoid such anachronism by carefully distinguishing James’s evolutionary views from some of their nearest conceptual neighbors. I focus in particular on James’s social evolutionism, especially as he expounds it in his 1880 (...) essay “Great Men, Great Thoughts, and the Environment.”1 By distinguishing James’s social evolutionism from sociobiology, from social Darwinism, and from the theory of memes, I underscore .. (shrink)

The article addresses the question whether culture evolves in a Lamarckian manner. I highlight three central aspects of a Lamarckian concept of evolution: the inheritance of acquired characteristics, the transformational pattern of evolution, and the concept of directed changes. A clear exposition of these aspects shows that a system can be a Darwinian variational system instead of a Lamarckian transformational one, even if it is based on inheritance of acquired characteristics and/or on Lamarckian directed changes. On this basis, I apply (...) the three aspects to culture. Taking for granted that culture is a variational system, based on selection processes, I discuss in detail the senses in which cultural inheritance can be said to be Lamarckian and in which sense problem solving, a major factor in cultural change, leads to directed variation. (shrink)

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, the nature of science (NOS) has become recognized as an important element within the K-12 science curriculum. Despite differences in the ultimate lists of recommended aspects, a consensus is emerging on what specific NOS elements should be the focus of science instruction and inform textbook writers and curriculum developers. In this article, we suggest a contextualized, explicit approach addressing one core NOS aspect: the human aspects of science that include the domains of creativity, social influences and subjectivity. To illustrate (...) these ideas, we have focused on Charles Darwin, a scientist whose life, work and thought processes were particularly well recorded at the time and analyzed by scholars in the succeeding years. Historical facts are discussed and linked to core NOS ideas. Creativity is illustrated through the analogies between the struggle for existence in human societies and in nature, between artificial and natural selection, and between the division of labor in human societies and in nature. Social influences are represented by Darwin’s aversion of criticism of various kinds and by his response to the methodological requirements of the science of that time. Finally, subjectivity is discussed through Darwin’s development of a unique but incorrect source for the origin of variations within species. (shrink)

This paper explores Charles Darwin’s hypothesis of pangenesis through a popular and professional reception history. First published in The Variation of Animals and Plants under Domestication (1868), pangenesis stated that inheritance can be explained by sub-cellular “gemmules” which aggregated in the sexual organs during intercourse. Pangenesis thereby accounted for the seemingly arbitrary absence and presence of traits in offspring while also clarifying some botanical and invertebrates’ limb regeneration abilities. I argue that critics largely interpreted Variation as an extension of On (...) the Origin of Species by Means of Natural Selection (1859), while pangenesis was an extension of natural selection. Contrary to claims that pangenesis was divorced from natural selection by its reliance on the inheritance of acquired characters, pangenesis’s mid nineteenth-century reception suggests that Darwin’s hypothesis responded directly to selection’s critics. Using Variation’s several editions, periodical reviews, and personal correspondence I assess pangenesis popularly, professionally, and biographically to better understand Variation’s impact on 1860s and 70s British evolutionism and inheritance. -/- . (shrink)

This inaugural handbook documents the distinctive research field that utilizes history and philosophy in investigation of theoretical, curricular and pedagogical issues in the teaching of science and mathematics. It is contributed to by 130 researchers from 30 countries; it provides a logically structured, fully referenced guide to the ways in which science and mathematics education is, informed by the history and philosophy of these disciplines, as well as by the philosophy of education more generally. The first handbook to cover the (...) field, it lays down a much-needed marker of progress to date and provides a platform for informed and coherent future analysis and research of the subject. -/- The publication comes at a time of heightened worldwide concern over the standard of science and mathematics education, attended by fierce debate over how best to reform curricula and enliven student engagement in the subjects There is a growing recognition among educators and policy makers that the learning of science must dovetail with learning about science; this handbook is uniquely positioned as a locus for the discussion. -/- The handbook features sections on pedagogical, theoretical, national, and biographical research, setting the literature of each tradition in its historical context. Each chapter engages in an assessment of the strengths and weakness of the research addressed, and suggests potentially fruitful avenues of future research. A key element of the handbook’s broader analytical framework is its identification and examination of unnoticed philosophical assumptions in science and mathematics research. It reminds readers at a crucial juncture that there has been a long and rich tradition of historical and philosophical engagements with science and mathematics teaching, and that lessons can be learnt from these engagements for the resolution of current theoretical, curricular and pedagogical questions that face teachers and administrators. (shrink)

Lucas Matthews and I substantially revised my SEP entry on Heritability. This version includes discussion of the missing heritability problem and other issues that arise from the use of Genome Wide Association Studies by Behavioral Geneticists.

Recent discussion of mechanism has suggested new approaches to several issues in the philosophy of science, including theory structure, causal explanation, and reductionism. Here, I apply what I take to be the fruits of the 'new mechanical philosophy' to an analysis of a contemporary debate in evolutionary biology about the role of natural selection in speciation. Traditional accounts of that debate focus on the geographic context of genetic divergence--namely, whether divergence in the absence of geographic isolation is possible (or significant). (...) Those accounts are at best incomplete, I argue, because they ignore the mechanisms producing divergence and miss what is at stake in the biological debate. I argue that the biological debate instead concerns the scope of particular speciation mechanisms which assign different roles to natural selection at various stages of divergence. The upshot is a new interpretation of the crux of that debate--namely, whether divergence with gene flow is possible (or significant) and whether the isolating mechanisms producing it are adaptive. (shrink)