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)

Clade selection is unpopular with philosophers who otherwise accept multilevel selection theory. Clades cannot reproduce, and reproduction is widely thought necessary for evolution by natural selection, especially of complex adaptations. Using microbial evolutionary processes as heuristics, I argue contrariwise, that (1) clade growth (proliferation of contained species) substitutes for clade reproduction in the evolution of complex adaptation, (2) clade-level properties favoring persistence – species richness, dispersal, divergence, and possibly intraclade cooperation – are not collapsible into species-level traits, (3) such properties (...) can be maintained by selection on clades, and (4) clade selection extends the explanatory power of the theory of evolution. (shrink)

There are several different ways in which chance affects evolutionary change. That all of these processes are called “random genetic drift” is in part a due to common elements across these different processes, but is also a product of historical borrowing of models and language across different levels of organization in the biological hierarchy. A history of the concept of drift will reveal the variety of contexts in which drift has played an explanatory role in biology, and will shed light (...) on some of the philosophical controversy surrounding whether drift is a cause of evolutionary change. (shrink)

The idea of ‘reversion’ or ‘atavism’ has a peculiar history. For many authors in the latenineteenth and early-twentieth centuries – including Darwin, Galton, Pearson, Weismann, and Spencer, among others – reversion was one of the central phenomena which a theory of heredity ought to explain. By only a few decades later, however, Fisher and others could look back upon reversion as a historical curiosity, a non-problem, or even an impediment to clear theorizing. I explore various reasons that reversion might have (...) appeared to be a central problem for this first group of figures, focusing on their commitment to a variety of conceptual features of evolutionary theory; discuss why reversion might have then ceased to be an interesting phenomenon; and, finally, close with some more general thoughts about the death of scientific problems. (shrink)

The Darwinian concept of natural selection was conceived within a set of Newtonian background assumptions about systems dynamics. Mendelian genetics at first did not sit well with the gradualist assumptions of the Darwinian theory. Eventually, however, Mendelism and Darwinism were fused by reformulating natural selection in statistical terms. This reflected a shift to a more probabilistic set of background assumptions based upon Boltzmannian systems dynamics. Recent developments in molecular genetics and paleontology have put pressure on Darwinism once again. Current work (...) on self-organizing systems may provide a stimulus not only for increased problem solving within the Darwinian tradition, especially with respect to origins of life, developmental genetics, phylogenetic pattern, and energy-flow ecology, but for deeper understanding of the very phenomenon of natural selection itself. Since self-organizational phenomena depend deeply on stochastic processes, self-organizational systems dynamics advance the probability revolution. In our view, natural selection is an emergent phenomenon of physical and chemical selection. These developments suggest that natural selection may be grounded in physical law more deeply than is allowed by advocates of the autonomy of biology, while still making it possible to deny, with autonomists, that evolutionary explanations can be modeled in terms of a deductive relationship between laws and cases. We explore the relationship between, chance, self-organization, and selection as sources of order in biological systems in order to make these points. (shrink)

Welch :263–279, 2017) has recently proposed two possible explanations for why the field of evolutionary biology is plagued by a steady stream of claims that it needs urgent reform. It is either seriously deficient and incapable of incorporating ideas that are new, relevant and plausible or it is not seriously deficient at all but is prone to attracting discontent and to the championing of ideas that are not very relevant, plausible and/or not really new. He argues for the second explanation. (...) This paper presents a twofold critique of his analysis: firstly, the main calls for reform do not concern the field of evolutionary biology in general but rather, or more specifically, the modern evolutionary synthesis. Secondly, and most importantly, these calls are not only inspired by the factors, enumerated by Welch, but are also, and even primarily, motivated by four problematic characteristics of the modern synthesis. This point is illustrated through a short analysis of the latest reform challenge to the modern synthesis, the so-called extended evolutionary synthesis. We conclude with the suggestion that the modern synthesis should be amended, rather than replaced. (shrink)

In this essay, I examine the usage of the term “just-so story.” I attempt to show that just-so storytelling can be seen as an epistemic concept that, in various ways, tackles the epistemological and methodological problems relating to evolutionary explanations qua historical/narrative explanations. I identify two main, yet mutually exclusive, strategies of employing the concept of a just-so story: a negative strategy and a positive strategy. Subsequently, I argue that these strategies do not satisfactorily capture the core of the “original” (...) meaning advanced by Stephen Jay Gould and Richard Lewontin at the end of the 1970s. I revisit the foundation(s) of their anti-adaptationist critique in order to reframe it as a critique of distinctive methodological manners and epistemic maxims related to historical inquiry. Last but not least, I suggest that contemporary evolutionary thinkers have two conceptually different options: they can either adhere to the “original” meaning of the term “just-so story” or accept that “just-so story” is a term equivalent to “implausible narrative explanation.”. (shrink)

Conceptions of adaptation have varied in the history of genetic Darwinism depending on whether what is taken to be focal is the process of adaptation, adapted states of populations, or discrete adaptations in individual organisms. I argue that Theodosius Dobzhansky’s view of adaptation as a dynamical process contrasts with so-called “adaptationist” views of natural selection figured as “design-without-a-designer” of relatively discrete, enumerable adaptations. Correlated with these respectively process and product oriented approaches to adaptive natural selection are divergent pictures of organisms (...) themselves as developmental wholes or as “bundles” of adaptations. While even process versions of genetical Darwinism are insufficiently sensitive to the fact much of the variation on which adaptive selection works consists of changes in the timing, rate, or location of ontogenetic events, I argue that articulations of the Modern Synthesis influenced by Dobzhansky are more easily reconciled with the recent shift to evolutionary developmentalism than are versions that make discrete adaptations central. (shrink)

Attempts to explain the origin of macroevolutionary innovations have been only partially successful. Here it is proposed that the patterns of major evolutionary transitions have to be understood first, before it is possible to further analyse the forces behind the process. The hypothesis is that major evolutionary innovations are characterized by an increase in organismal autonomy, in the sense of emancipation from the environment. After a brief overview of the literature on this subject, increasing autonomy is defined as the evolutionary (...) shift in the individual system–environment relationship, such that the direct influences of the environment are gradually reduced and a stabilization of self-referential, intrinsic functions within the system is generated. This is described as relative autonomy because numerous interconnections with the environment and dependencies upon it are retained. Features of increasing autonomy are spatial separations, an increase in homeostatic functions and in body size, internalizations and an increase in physiological and behavioral flexibility. It is described how these features are present in different combinations in the major evolutionary transitions of metazoans and, consequently, how they should be taken into consideration when evolutionary innovations are studied. The hypothesis contributes to a reconsideration of the relationship between organisms and their environment. (shrink)

David Lack of Oxford University and V. C. Wynne- Edwards of Aberdeen University were renowned ornithologists with contrasting views of the modern synthesis which deeply influenced their interpretation and explanation of bird behavior. In the 1950's and 60's Lack became the chief advocate of neo-Darwinism with respect to avian ecology, while Wynne- Edwards developed his theory of group selection. Lack 's position was consistent with the developing focus on individual level adaptation, which was a core concept of the modern synthesis. (...) Alternatively, Wynne- Edwards viewed the emphasis on populations as the most important development provided by the modern synthesis. In this paper, I present the development of these two positions and trace their roots in the literature of the synthesis. Through an analysis of Lack 's 1966 critique of Wynne- Edwards I conclude that Wynne- Edwards was, in many ways, justified in his pursuit of group level explanations. (shrink)

The 1940s and 1950s were marked by intense debates over the origin of drug resistance in microbes. Bacteriologists had traditionally invoked the notions of ‘training’ and ‘adaptation’ to account for the ability of microbes to acquire new traits. As the field of bacterial genetics emerged, however, its participants rejected ‘Lamarckian’ views of microbial heredity, and offered statistical evidence that drug resistance resulted from the selection of random resistant mutants. Antibiotic resistance became a key issue among those disputing physiological vs. genetic (...) explanations of variation in bacteria. Postwar developments connected with the Lysenko affair gave this debate a new political valence.Proponents of the neo-Darwinian synthesis weighed in with support for the genetic theory. However, certain features of drug resistance seemed inexplicable by mutation and selection, particularly the phenomenon of ‘multiple resistance’—the emergence of resistance in a single strain against several unrelated antibiotics. In the late 1950s, Tsutomu Watanabe and his collaborators solved this puzzle by determining that resistance could be conferred by cytoplasmic resistance factors rather than chromosomal mutation. These R factors could carry resistance to many antibiotics and seemed able to promote their own dissemination in bacterial populations. In the end, the vindication of the genetic view of drug resistance was accompanied by a recasting of the ‘gene’ to include extrachromosomal hereditary units carried on viruses and plasmids. (shrink)

This paper extends previous arguments against the assumption that the study of variation at the molecular level was instigated with a view to solving an internal conflict between the balance and classical schools of population genetics. It does so by focusing on the intersection of basic research in protein chemistry and the molecular approach to disease with the enactment of global health campaigns during the Cold War period. The paper connects advances in research on protein structure and function as reflected (...) in Christian Anfinsen’s The molecular basis of evolution, with a political reading of Emilé Zuckerkandl and Linus Pauling’s identification of molecular disease and evolution. Beyond atomic fallout, these advances constituted a rationale for the promotion of genetic surveys of human populations in the Third World, in connection with international health programs. Light is shed not only on the experimental roots of the molecular challenge but on the broader geopolitical context where the rising role of biomedicine and public health had an impact on evolutionary biology. (shrink)

The 1920-1960 period saw the creation of the conditions for a unification of disciplines in the area of evolutionary biology under a limited number of theoretical prescriptions: the evolutionary synthesis. Whereas the sociological dimension of this synthesis was fairly successful, it was surprisingly loose when it came to the interpretation of the evolutionary mechanisms per se, and completely lacking at the level of the foundational epistemological and metaphysical commitments. Key figures such as Huxley, Simpson, Dobzhansky, and Rensch only paid lip (...) service to the conceptual dimension of the evolutionary synthesis, as they eventually realized that a number of evolutionary phenomena could not be explained by its narrow theoretical corpus. Apparently, the evolutionary synthesis constituted a premature event in the development of evolutionary biology. Not only are the real achievements of the evolutionary synthesis in need of reevaluation, but this reassessment also has important implications for the historiography of Darwinism and the current debates about the darwinian movement. (shrink)

The Evolutionary Synthesis is often seen as a unification process in evolutionary biology, one which provided this research area with a solid common theoretical foundation. As such, neo-Darwinism is believed to constitute from this time onward a single, coherent, and unified movement offering research guidelines for investigations. While this may be true if evolutionary biology is solely understood as centred around evolutionary mechanisms, an entirely different picture emerges once other aspects of the founding neo-Darwinists’ views are taken into consideration, aspects (...) potentially relevant to the elaboration of an evolutionary worldview: the tree of life, the ontological distinctions of the main cosmic entities , the inherent properties of self-organizing matter, evolutionary ethics, and so on. Profound tensions and inconsistencies are immediately revealed in the neo-Darwinian movement once this broader perspective is adopted. This pluralism is such that it is possible to identify at least three distinct and quasi-incommensurable epistemological/metaphysical frameworks as providing a proper foundation for neo-Darwinism. The analysis of the views of Theodosius Dobzhansky, Bernhard Rensch, and Ernst Mayr will illustrate this untenable pluralism, one which requires us to conceive of the neo-Darwinian research agenda as being conducted in more than one research programme or research tradition at the same time. (shrink)

ABSTRACT. This paper contemplates Organicism and its relation with molecular and evolutionary biology. We explore whether twentieth-first century biology is returning to positions held at the beginning of the twentieth century and then abandoned. The guiding line is a history of theoretical biology in which we distinguish three periods: 1. The 20s-30s, and the Theoretical Biology Club (Needham, Woodger, and Waddington, among others); 2. An intermediate period in the 60s-70s, in which, in spite of the eclosion of the molecular and (...) evolutionary biologies, there is some recovery of organicist positions, and 3. The present post-genomic situation, which is demanding a systemic approach. (shrink)

This book is divided in two parts, the first of which shows how, beyond paleontology and systematics, macroevolutionary theories apply key insights from ecology and biogeography, developmental biology, biophysics, molecular phylogenetics, and even the sociocultural sciences to explain evolution in deep time. In the second part, the phenomenon of macroevolution is examined with the help of real life-history case studies on the evolution of eukaryotic sex, the formation of anatomical form and body-plans, extinction and speciation events of marine invertebrates, hominin (...) evolution and species conservation ethics. The book brings together leading experts, who explain pivotal concepts such as Punctuated Equilibria, Stasis, Developmental Constraints, Adaptive Radiations, Habitat Tracking, Turnovers, (Mass) Extinctions, Species Sorting, Major Transitions, Trends, and Hierarchies – key premises that allow macroevolutionary epistemic frameworks to transcend microevolutionary theories that focus on genetic variation, selection, migration and fitness. Along the way, the contributing authors review ongoing debates and current scientific challenges; detail new and fascinating scientific tools and techniques that allow us to cross the classic borders between disciplines; demonstrate how their theories make it possible to extend the Modern Synthesis; present guidelines on how the macroevolutionary field could be further developed; and provide a rich view of just how it was that life evolved across time and space. In short, this book is a must-read for active scholars and, because the technical aspects are fully explained, it is also accessible for non-specialists. Understanding evolution requires a solid grasp of above-population phenomena. Species are real biological individuals, and abiotic factors impact the future course of evolution. Beyond observation, when the explanation of macroevolution is the goal, we need both evidence and theory that enable us to explain and interpret how life evolves at the grand scale. (shrink)

Recent work on the heat-shock protein Hsp90 by Rutherford and Lindquist (1998) has been included among the pieces of evidence taken to show the essential role of developmental processes in evolution; Hsp90 acts as a buffer against phenotypic variation, allowing genotypic variation to build. When the buffering capacity of Hsp90 is altered (e.g., in nature, by mutation or environmental stress), the genetic variation is "revealed," manifesting itself as phenotypic variation. This phenomenon raises questions about the genetic variation before and after (...) what I will call a "revelation event": Is it neutral, nearly neutral, or non-neutral (i.e., strongly deleterious or strongly advantageous)? Moreover, what kinds of evolutionary processes do we take to be at work? Rutherford and Lindquist (1998) focus on the implications of non-neutral variation and selection. Later work by Queitsch, Sangster, and Lindquist (2002) and Sangster, Lindquist, and Queitsch (2004) raises the possibility that Hsp90 buffering may play the role that was played by drift in Sewall Wright's shifting balance model, permitting transition from one adaptive peak to another. However, Ohta (2002) suggests that much of this variation may be nearly neutral, which in turn, would imply a strong role for drift as well as selection. The primary goal of this paper is to illuminate the alternative scenarios and the processes operating in each. At the end, I raise the possibility of a synthesis between evo-devo and nearly neutral evolution. (shrink)

Darwinism is defined here as an evolving research tradition based upon the concepts of natural selection acting upon heritable variation articulated via background assumptions about systems dynamics. Darwin’s theory of evolution was developed within a context of the background assumptions of Newtonian systems dynamics. The Modern Evolutionary Synthesis, or neo-Darwinism, successfully joined Darwinian selection and Mendelian genetics by developing population genetics informed by background assumptions of Boltzmannian systems dynamics. Currently the Darwinian Research Tradition is changing as it incorporates new information (...) and ideas from molecular biology, paleontology, developmental biology, and systems ecology. This putative expanded and extended synthesis is most perspicuously deployed using background assumptions from complex systems dynamics. Such attempts seek to not only broaden the range of phenomena encompassed by the Darwinian Research Tradition, such as neutral molecular evolution, punctuated equilibrium, as well as developmental biology, and systems ecology more generally, but to also address issues of the emergence of evolutionary novelties as well as of life itself. (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)

Although sciences are often conceptualized in terms of theory confirmation and hypothesis testing, an equally important dimension of scientific reasoning is the structure of problems that guide inquiry. This problem structure is evident in several concepts central to evolutionary developmental biology (Evo-devo)—constraints, modularity, evolvability, and novelty. Because problems play an important role in biological practice, they should be included in biological pedagogy, especially when treating the issue of scientific controversy. A key feature of resolving controversy is synthesizing methodologies from different (...) biological disciplines to generate empirically adequate explanations. Concentrating on problem structure illuminates this interdisciplinarity in a way that is often ignored when science is taught only from the perspective of theory or hypothesis. These philosophical considerations can assist life science educators in their continuing quest to teach biology to the next generation. -/- . (shrink)

Contingency-theorists have gestured to a series of phenomena such as random mutations or rare Armageddon-like events as that which accounts for evolutionary contingency. These phenomena constitute a class, which may be aptly called the ‘sources of contingency’. In this paper, I offer a probabilistic conception of what it is to be a source of contingency and then examine two major candidates: chance variation and genetic drift, both of which have historically been taken to be ‘chancy’ in a number of different (...) senses. However,contrathe gesturing of contingency-theorists, chance variation and genetic drift are not always strong sources of contingency, as they can be non-chancy in at least one sense that opposes evolutionary contingency. The probabilistic conception offered herein allows for sources of contingency to appropriately vary in strength. To this end, I import Shannon’sinformation entropyas a statistical measure for systematically assessing the strength of a source of contingency, which is part and parcel of identifying sources of contingency. In brief, the higher the entropy, the greater the strength. This is also empirically significant because molecular, mutational, and replicative studies often contain sufficient frequency or probability data to allow for entropies to be calculated. In this way, contingency-theorists can evaluate the strength of a source of contingency in real-world cases. Moreover, the probabilistic conception also makes conceptual room for the converse of sources of contingency: ‘sources of directionality’, which ought to be recognised, as they can interact with genuine sources of contingency in undermining evolutionary contingency. (shrink)

This essay interprets Schelling's positive philosophy as an effort to conceive nature as a gift. Schelling ruminated throughout his career on the paradoxical relation between humanity and nature that is expressed in the contemporary term “Anthropocene,” but this essay argues that Schelling's most productive response to this paradox can be found in his reflections on how to receive the gift of nature. After laying out the project of positive philosophy, the essay first explores Schelling's effort to conceive nature as a (...) gift in On the World-Soul, and then considers how this project founders in the First Projection of a System of Nature Philosophy. After the resources of the natural sciences showed themselves to be inadequate to the task of conceiving nature as a gift, Schelling returned to mythology in his late philosophy, and the concluding section of this essay traces how an often overlooked Epicurean strain in Schelling's late philosophy can help us see what is entailed in conceiving nature as a gift. (shrink)

Historians and philosophers of science have furnished a wide array of theoretical-historiographical terms to emphasize the discontinuities among different systems of knowledge. Some of the most famous include Thomas Kuhn’s “paradigm”, Michel Foucault’s “episteme”, and the notion of “styles of reasoning” more recently developed by Ian Hacking and Arnold Davidson. This paper takes up this theoretical-historiographical thread by assessing the values and limitations of the notion of “style” for the historical and philosophical study of science. Specifically, reflecting on various methodological (...) and theoretical concerns prompted by sexuality, translation, and East Asian studies, this paper argues that the heretofore ways in which historians and philosophers of science have used the notion of “style” are severely restricted in terms of its mere applicability to the intellectual history of Western science. The particular example of the translation of “homosexuality” into Chinese during the May Fourth era reveals that the notion of “style” has the potential of carrying a much more dynamic conceptual weight, as when used in “styles of argumentation”. The paper also engages briefly with the historiography of scientific “national styles” and ends with some concluding remarks on the limitations of “social histories from below” and the under appreciated importance of “epistemological histories of possibilities”. (shrink)

Despite the amount of work that has been produced on the subject over the years, the ‘transformation of cladistics’ is still a misunderstood episode in the history of comparative biology. Here, I analyze two outstanding, highly contrasting historiographic accounts on the matter, under the perspective of an influential dichotomy in the philosophy of science: the opposition between Scientific Realism and Empiricism. Placing special emphasis on the notion of ‘causal grounding’ of morphological characters in modern developmental biology’s theories, I arrive at (...) the conclusion that a ‘new transformation of cladistics’ is philosophically plausible. This ‘reformed’ understanding of ‘pattern cladistics’ entails retaining the interpretation of cladograms as ‘schemes of synapomorphies’, but in association to construing cladogram nodes as ‘developmental-genetic taxic homologies’, instead of ‘standard Darwinian ancestors’. The reinterpretation of pattern cladistics presented here additionally proposes to take Bas Van Fraassen’s ‘constructive empiricism’ as a philosophical stance that could properly support such analysis of developmental-genetic data for systematic purposes. The latter suggestion is justified through a reappraisal of previous ideas developed by prominent pattern cladists , which concerned a scientifically efficient ‘observable/non-observable distinction’ linked to the conceptual pair ‘ontogeny and phylogeny’. Finally, I argue that a robust articulation of Antirealist alternatives in systematics may provide a rational basis for its disciplinary separation from evolutionary biology, as well as for a critical reconsideration of the proper role of certain Scientific Realist positions, currently popular in comparative biology. (shrink)

In the received view of the history of the Modern Evolutionary Synthesis, paleontology was given a prominent role in evolutionary biology thanks to the significant influence of paleontologist George Gaylord Simpson on both the institutional and conceptual development of the Synthesis. Simpson's 1944 Tempo and Mode in Evolution is considered a classic of Synthesis-era biology, and Simpson often remarked on the influence of other major Synthesis figures – such as Ernst Mayr and Theodosius Dobzhansky – on his developing thought. Why, (...) then, did paleontologists of the 1970s and 1980s – Stephen Jay Gould, Niles Eldredge, David M. Raup, Steven Stanley, and others – so frequently complain that paleontology remained marginalized within evolutionary biology? This essay considers three linked questions: first, were paleontologists genuinely welcomed into the Synthetic project during its initial stages? Second, was the initial promise of the role for paleontology realized during the decades between 1950 and 1980, when the Synthesis supposedly "hardened" to an "orthodoxy"? And third, did the period of organized dissent and opposition to this orthodoxy by paleontologists during the 1970s and 1980s bring about a long-delayed completion to the Modern Synthesis, or rather does it highlight the wider failure of any such unified Darwinian evolutionary consensus? (shrink)

The Human Genome Project (HGP) has been criticised from an evolutionary perspective for three reasons: completely ignoring genetic variation; improperly treating either all or some genetic variation as deviation from a norm; and mistakenly seeking to define species in terms of essential properties possessed by all and only member organisms. The first claim is unfounded; the second and third claims are more on target. Nevertheless, it is a mistake to use the typological-population distinction to oppose molecular genetics and evolutionary genetics (...) in order to characterise HGP mapping and sequencing aims, especially the production of a DNA reference sequence, as 'anti-evolutionary' and 'pre-Darwinian.' These aims are consistent with certain strands in twentieth-century evolutionary thought: Muller's classical theory, Kimura's neutral and 'effectively neutral' theories, and, to a lesser extent, Dobzhansky's balance theory of the genetic structure of natural populations. In practice, population-based approaches to human genetic variation are similarly vulnerable to charges of 'typological' and 'essentialist' thinking in their treatment of genetic variation as deviation. This means that the Human Genome Diversity Initiative will not provide a population-based panacea for an overly typological HGP, as its proponents contend. Substituting related, and less rhetorically-charged, conceptual, empirical, and metaphysical distinctions for the typological-population distinction furnishes a better way to assess the concept of the normal genome from an evolutionary perspective. (shrink)

One of the central aims of science is explanation: scientists seek to uncover why things happen the way they do. This chapter addresses what kinds of explanations are formulated in biology, how explanatory aims influence other features of the field of biology, and the implications of all of this for biology education. Philosophical treatments of scientific explanation have been both complicated and enriched by attention to explanatory strategies in biology. Most basically, whereas traditional philosophy of science based explanation on derivation (...) from scientific laws, there are many biological explanations in which laws play little or no role. Instead, the field of biology is a natural place to turn for support for the idea that causal information is explanatory. Biology has also been used to motivate mechanistic accounts of explanation, as well as criticisms of that approach. Ultimately, the most pressing issue about explanation in biology may be how to account for the wide range of explanatory styles encountered in the field. This issue is crucial, for the aims of biological explanation influence a variety of other features of the field of biology. Explanatory aims account for the continued neglect of some central causal factors, a neglect that would otherwise be mysterious. This is linked to the persistent use of models like evolutionary game theory and population genetic models, models that are simplified to the point of unreality. These explanatory aims also offer a way to interpret many biologists’ total commitment to one or another methodological approach, and the intense disagreements that result. In my view, such debates are better understood as arising not from different theoretical commitments, but commitments to different explanatory projects. Biology education would thus be enriched by attending to approaches to biological explanation, as well as the unexpected ways that these explanatory aims influence other features of biology. I suggest five lessons for teaching about explanation in biology that follow from the considerations of this chapter. (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)

A number of areas of biology raise questions about what is of value in the natural environment and how we ought to behave towards it: conservation biology, environmental science, and ecology, to name a few. Based on my experience teaching students from these and similar majors, I argue that the field of environmental ethics has much to teach these students. They come to me with pent-up questions and a feeling that more is needed to fully engage in their subjects, and (...) I believe some exposure to environmental ethics can help focus their interests and goals. I identify three primary areas in which environmental ethics can con- tribute to their education. The first is an examination of who (or what) should be considered to be part of our moral community (i.e., the community to whom we owe direct duties). Is it humans only? Or does it include all sentient life? Or all life? Or ecosystems considered holistically? Often, readings implicitly assume one or more of these answers; the goal is to make the student more sensitive to these implicit claims and to get them to think about the different reasons that support them. The second area, related to the first, is the application of the different answers concerning the extent of the ethical community to real environmental issues and problems. Students need to be aware of how the different answers concerning the moral community can imply conflicting answers for how we should act in certain cases and to think about ways to move toward conflict resolution. The third area in which environmental ethics can contribute is a more conceptual one, focusing on central concepts such as biodiversity, sustainability, species, and ecosystems. Exploring and evaluating various meanings of these terms will make students more reflective and thoughtful citizens and biologists, sensitive to the implications that different conceptual choices make. (shrink)

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)

This article scrutinizes in detail much of the extant historiography on the controversy between biometricians and Mendelians, considering in particular how this controversy is related to the evolutionary synthesis. While the historiographic critique concentrates on William Provine’s standard account, it also extends to the proposal by Donald MacKenzie and Barry Barnes. What Provine and these sociologists of scientific knowledge have in common is a set of unquestioned assumptions about the nature of Darwinism, about William Bateson’s anti-Darwinism, and about the very (...) idea of an evolutionary “synthesis.” While these assumptions make for a compelling history of the synthesis, they engender an endemically asymmetrical perspective and bias historiography toward the mere confirmation of antecedent expectations, which renders the years from 1859 to 1929 an age of ignorance and misunderstanding. In contrast, a return to what was probably the original meaning of evolutionary “synthesis” allows for a symmetrical account. It yields an appreciation of the positive contributions made both by Mendelians and biometricians to the gradual development of Darwinism. The article concludes with a synopsis of this alternative account. (shrink)

La crítica kantiana legó una doble herencia a la biología decimonónica: su noción de ciencia basada en el mecanicismo newtoniano configuró epistemológicamente la teoría de la evolución darwinista, mientras que su comprensión de los organismos se tradujo en una morfología teleológica. En este artículo planteamos dos cuestiones en torno la relación entre las ideas de Kant y Darwin: 1) si Kant habría considerado a Darwin el Newton de la biología, a lo que, con matices, respondemos afirmativamente; 2) si la física (...) newtoniana es hoy suficiente para naturalizar lo orgánico. Nuestra respuesta negativa se fundamenta en las ciencias de la autoorganización y la evo-devo, que permiten plantear la naturalización de la concepción kantiana de los organismos. (shrink)

Huxley coined the phrase, the “evolutionary synthesis” to refer to the acceptance by a vast majority of biologists in the mid-20th Century of a “synthetic” view of evolution. According to this view, natural selection acting on minor hereditary variation was the primary cause of both adaptive change within populations and major changes, such as speciation and the evolution of higher taxa, such as families and genera. This was, roughly, a synthesis of Mendelian genetics and Darwinian evolutionary theory; it was a (...) demonstration that prior barriers to understanding between various subdisciplines in the life sciences could be removed. The relevance of different domains in biology to one another was established under a common research program. The evolutionary synthesis may be broken down into two periods, the “early” synthesis from 1918 through 1932, and what is more often called the “modern synthesis” from 1936-1947. The authors most commonly associated with the early synthesis are J.B.S. Haldane, R.A. Fisher, and S. Wright. These three figures authored a number of important synthetic advances; first, they demonstrated the compatibility of a Mendelian, particulate theory of inheritance with the results of Biometry, a study of the correlations of measures of traits between relatives. Second, they developed the theoretical framework for evolutionary biology, classical population genetics. This is a family of mathematical models representing evolution as change in genotype frequencies, from one generation to the next, as a product of selection, mutation, migration, and drift, or chance. Third, there was a broader synthesis of population genetics with cytology, genetics, and biochemistry, as well as both empirical and mathematical demonstrations to the effect that very small selective forces acting over a relatively long time were able to generate substantial evolutionary change, a novel and surprising result to many skeptics of Darwinian gradualist views. The later “modern” synthesis is most often identified with the work of Mayr, Dobzhansky and Simpson. There was a major institutional change in biology at this stage, insofar as different subdisciplines formerly housed in different departments, and with different methodologies were united under the same institutional umbrella of “evolutionary biology.” Mayr played an important role as a community architect, in founding the Society for the Study of Evolution, and the journal Evolution, which drew together work in systematics, biogeography, paleontology, and theoretical population genetics. (shrink)

Barely a decade after the discovery of the chromosomal basis of inheritance, and the articulation of the genetical theory of population change, the gene came to be widely regarded as the fundamental unit of biological organization. This is hardly surprising. The gene concept is a powerful one; it plays a unifying role in our understanding of evolution. Darwin told us that evolution by natural selection occurs in a population when organisms survive, die and reproduce differentially on account of their heritable (...) form. This is a very schematic theory. It requires an account of the process of inheritance and also an account of the generation of phenotype. The gene concept plays a prominent role in explaining, and uniting, these phenomena. Genes are the units of inheritance; they are passed from parents to offspring in reproduction. Moreover, they are seen as units of phenotypic control. Evolutionary biologists often speak of the genome as a program for the production of an organism. Genes also became the elements of which populations are composed. Our best theory of population dynamics—inherited from Fisher, Haldane, and Wright—is a theory of changes in the relative frequencies of gene types. Genes are not just the principal causes of evolutionary change, they are also the units over which evolutionary change is defined and measured. So, at least, the orthodox reading of the Modern Synthesis theory of evolution would have us believe... (shrink)

Erwin Schrödinger’s 1944 publication What is Life? is a classic of twentieth century science writing. In his book, Schrödinger discussed the chromosome fibre as the seat of heredity and variation thanks to a hypothetical aperiodic structure – a suggestion that famously spurred on a generation of scientists in their pursuit of the gene as a physico-chemical entity. While historical attention has been given to physicists who were inspired by the book, little has been written about its biologist readers. This paper (...) examines the case of the English evolutionary botanist and cytologist Irène Manton, who took an interest in What is Life? for its relevance to her own research in chromosome structure as a clue to plant phylogeny. Drawing on recently discovered correspondence between Manton and Schrödinger, the paper reconstructs Manton ‘s path to the book and her response to it by way of throwing new light on a pivotal moment in the history of the debate on chromosome structure. (shrink)

The Modern Synthesis has been receiving bad press for some time now. Back in 1983, in an article entitled “The Hardening of the Modern Synthesis” Stephen Jay Gould criticized the way the Modern Synthesis had developed since its inception in the 1930s and early 1940s (Gould 1983). Back then, those who would later become known as ‘architects’ of the synthesis were united in their call for explaining evolution at all levels in terms of causation at one level: genetics. What drove (...) changes in gene frequency remained an open question. It could be mainly selection, or drift, or some (other) form of constraint. But in the two decades that followed, the synthesis underwent a major change. By the late 1940s the synthesis had ‘hardened’ around adaptationism, according to Gould. Influential contributors like Dobzhansky, Simpson and Wright had increasingly expressed adaptationist views in the later editions of their landmark books. Not because evidence had piled up, showing that selection was in fact pervasive. Instead, Gould argued, adaptationist tendencies had been preserved by some kind of cultural inertia, and were now being revived. “Certain ‘national styles’ persisted from the eighteenth century, through Darwin’s era, and into our own time. Views on adaptation provide a good example” (Gould 1983). Gould did not just argue that some form of adaptationism had resurfaced. He became well-known for his efforts to intervene on this status quo by attempting to make evolutionary biology more ‘pluralistic’. In collaborative work with Richard Lewontin (Gould and Lewontin 1979), Elisabeth Vrba (Gould and Vrba 1982; Vrba and Gould 1986) and Niles Eldredge (Eldredge and Gould 1972; Gould and Eldredge 1977) he criticized the synthesis for its adaptationism and its lack of appreciation for hierarchical perspectives. Gould exerted his influence in a different way as well. Together with Eldredge, he had facsimiles reprinted of the first editions of two books that had shaped synthesis, but with their own critical introductions (Eldredge 1982; Gould 1982). Dobzhansky’s Genetics and the Origin of Species and Mayr’s Systematics and the Origin of Species appeared as reprints in the ‘Columbia Classics in Evolution’ series, sending an unambiguous message to the readers: these are foundational works, but they have been superseded. In the summer of 2008, some 25 years after Gould made his point about the hardening of the Modern Synthesis, a group of sixteen biologists and philosophers gathered at the Konrad Lorenz Institute for Evolution and Cognition Research (KLI) near Vienna, Austria, to discuss cutting-edge research that reaches beyond the synthesis framework. Before it even started, this workshop on the ‘Extended Synthesis’ had already attracted a fair share of attention in the blogosphere and had resulted in a news feature in Science (Pennisi 2008). After the meeting, Nature weighed in on the matter (Whitfield 2008). The results of over 3 days of presentations and extensive discussion have now been published as Evolution—The Extended Synthesis. 1 The publication of this collection of sixteen essays is accompanied by the republication of Julian Huxley’s Evolution: The Modern Synthesis; the book that introduced the term ‘Modern Synthesis’. Both books are introduced by the organizers of the KLI workshop, Massimo Pigliucci and Gerd Müller. Like Gould and Eldredge before them, Pigliucci and Müller did not reissue one of the canons of the Modern Synthesis without giving the readers some ‘guidance’. Starting with the cover, the editors proclaim boldly that this is ‘the definitive edition’ of Huxley’s book. In a new foreword, they sketch the context in which the book was written and assess some of its features. They voice some mild criticism of alleged ‘adaptationism’. But their tone is different from that of Gould and Eldredge. Pigliucci and Müller praise Huxley for his pluralistic outlook, which has again become essential in the forging of an Extended Synthesis. That makes Huxley’s book more than just an interesting but obsolete classic. Instead, it can teach valuable lessons about how to ‘soften up’ a synthesis that has become hardened over time. (shrink)

The dichotomy between ‘typological thinking’ and ‘population thinking’ features in a range of debates in contemporary and historical biology. The origins of this dichotomy are often traced to Ernst Mayr, who is said to have coined it in the 1950s as a rhetorical device that could be used to shield the Modern Synthesis from attacks by the opponents of population biology. In this two-part essay, I argue that the origins of the typology/population dichotomy are considerably more complicated and more interesting (...) than is commonly thought. In the first part, I argued that Mayr's dichotomy was based on two distinct type/population contrasts that had been articulated much earlier by George Gaylord Simpson and Theodosius Dobzhansky. Their distinctions made eminent sense in their own, isolated contexts. In this second part, I will show how Mayr conflated these type/population distinctions and blended in some of his own, unrelated concerns with ‘types’ of a rather different sort. Although Mayr told his early critics that he was merely making “a temporary oversimplification,” he ended up burdening the history and philosophy of biology with a troubled dichotomy. (shrink)

The dichotomy between ‘typological thinking’ and ‘population thinking’ features in a range of debates in contemporary and historical biology. The origins of this dichotomy are often traced to Ernst Mayr, who is said to have coined it in the 1950s as a rhetorical device that could be used to shield the Modern Synthesis from attacks by the opponents of population biology. In this two-part essay I argue that the origins of the typology/population dichotomy are considerably more complicated and more interesting (...) than is commonly thought. In this first part, I will argue that Mayr's dichotomy was based on two distinct type/population contrasts that had been articulated much earlier by George Gaylord Simpson and Theodosius Dobzhansky. Their distinctions made eminent sense in their own, isolated contexts. In the second part, I will show how Mayr conflated these type/population distinctions and blended in some of his own, unrelated concerns with ‘types’ of a rather different sort. Although Mayr told his early critics that he was merely making “a temporary oversimplification,” he ended up burdening the history and philosophy of biology with a troubled dichotomy. (shrink)

The term “genetic load” first emerged in a paper written in 1950 by the geneticist H. Muller. It is a mathematical model based on biological, social, political and ethical arguments describing the dramatic accumulation of disadvantageous mutations in human populations that will occur in modern societies if eugenic measures are not taken. The model describes how the combined actions of medical and social progress will supposedly impede natural selection and make genes of inferior quality likely to spread across populations – (...) a process which in fine loads their progress. Genetic load is based on optimal fitness and emerges from a “typological view” of evolution. This model of evolution had previously, however, been invalidated by Robert Wright and Theodosius Dobzhansky who, as early as 1946, showed that polymorphism was the rule in natural populations. The blooming and persistence of the concept of genetic load, after its theoretical basis had already expired, are a historical puzzle. This persistence reveals the intricacy of science and policy-making in eugenic matters. The Canguilhemian concept of ‘scientific ideology’ is used along with the concept of ‘immutable mobile’ and compared with the concept of ‘co-production’, to provide complementary perspectives on this complex phenomenon. (shrink)