We trace the history of the Modern Evolutionary Synthesis, and of genetic Darwinism generally, with a view to showing why, even in its current versions, it can no longer serve as a general framework for evolutionary theory. The main reason is empirical. Genetical Darwinism cannot accommodate the role of development (and of genes in development) in many evolutionary processes. We go on to discuss two conceptual issues: whether natural selection can be the “creative factor” in a new, more general framework (...) for evolutionary theorizing; and whether in such a framework organisms must be conceived as self-organizing systems embedded in self-organizing ecological systems. (shrink)

The theory of evolution, which provides the conceptual framework for all modern research in organismal biology and informs research in molecular bi- ology, has gone through several stages of expansion and refinement. Darwin and Wallace (1858) of course proposed the original idea, centering on the twin concepts of natural selection and common descent. Shortly thereafter, Wallace and August Weismann worked toward the complete elimination of any Lamarckian vestiges from the theory, leaning in particular on Weismann’s (1893) concept of the separation (...) of soma and germ, resulting in what is some- times referred to as “neo-Darwinism”. (shrink)

This article addresses the emergence of human personality in evolution. The mechanisms of natural and sexual selection developed by Darwin are not sufficient to explain the sense of self. Therefore we attempt to trace the evolutionary process back to a form of selection termed “emotional selection.” This involves reconstructing selection out of subjective qualities and showing how emotions enable human forms of life that are relevant for the cultural level of cooperation that marks our species. We see a paradigm shift (...) in the concept of emotional selection that binds emotion and evolution closer together, thus closing the explanatory gap between classical ethology and modern evolutionary psychology. (shrink)

Laland and colleagues have sought to challenge the proximate–ultimate distinction claiming that it imposes a unidirectional model of causation, is limited in its capacity to account for complex biological phenomena, and hinders progress in biology. In this article the core of their argument is critically analyzed. It is claimed that contrary to their claims Laland et al. rely upon the proximate–ultimate distinction to make their points and that their alternative conception of reciprocal causation refers to phenomena that were already accounted (...) for by standard theory. (shrink)

The idea of genetic assimilation, that environmentally induced phenotypes may become genetically fixed and no longer require the original environmental stimulus, has had varied success through time in evolutionary biology research. Proposed by Waddington in the 1940s, it became an area of active empirical research mostly thanks to the efforts of its inventor and his collaborators. It was then attacked as of minor importance during the ‘‘hardening’’ of the neo-Darwinian synthesis and was relegated to a secondary role for decades. Recently, (...) several papers have appeared, mostly independently of each other, to explore the likelihood of genetic assimilation as a biological phenomenon and its potential importance to our understanding of evolution. In this article we briefly trace the history of the concept and then discuss theoretical models that have newly employed genetic assimilation in a variety of contexts. We propose a typical scenario of evolution of genetic assimilation via an intermediate stage of phenotypic plasticity and present potential examples of the same. We also discuss a conceptual map of current and future lines of research aimed at exploring the actual relevance of genetic assimilation for evolutionary biology. (shrink)

There are two motivations commonly ascribed to historical actors for taking up statistics: to reduce complicated data to a mean value (e.g., Quetelet), and to take account of diversity (e.g., Galton). Different motivations will, it is assumed, lead to different methodological decisions in the practice of the statistical sciences. Karl Pearson and W. F. R. Weldon are generally seen as following directly in Galton’s footsteps. I argue for two related theses in light of this standard interpretation, based on a reading (...) of several sources in which Weldon, independently of Pearson, reflects on his own motivations. First, while Pearson does approach statistics from this "Galtonian" perspective, he is, consistent with his positivist philosophy of science, utilizing statistics to simplify the highly variable data of biology. Weldon, on the other hand, is brought to statistics by a rich empiricism and a desire to preserve the diversity of biological data. Secondly, we have here a counterexample to the claim that divergence in motivation will lead to a corresponding separation in methodology. Pearson and Weldon, despite embracing biometry for different reasons, settled on precisely the same set of statistical tools for the investigation of evolution. (shrink)

Evo-Devo exhibits a plurality of scientific “cultures” of practice and theory. When are the cultures acting—individually or collectively—in ways that actually move research forward, empirically, theoretically, and ethically? When do they become imperialistic, in the sense of excluding and subordinating other cultures? This chapter identifies six cultures – three /styles/ (mathematical modeling, mechanism, and history) and three /paradigms/ (adaptationism, structuralism, and cladism). The key assumptions standing behind, under, or within each of these cultures are explored. Characterizing the internal structure of (...) the cultures is necessary for understanding how they collaborate or compete, and how they are fragmented or integrated, in the rich interdisciplinary /trading zone/ (Galison 1997) of Evo-Devo. Evo-Devo is an important example of how science can progress through a radical plurality of perspectives and cultures. (shrink)

The contents of the British Ornithologists' Union's journal, "The Ibis," during the first half of the 20th century illustrates some of the transformations that have taken place in the naturalist tradition. Although later generations of ornithologists described these changes as logical and progressive, their historical narratives had more to do with legitimizing the infiltration of the priorities of evolutionary theory, ecology, and ethology than analyzing the legacy of the naturalist tradition on its own terms. Despite ornithologists' claim that the journal's (...) increasing focus on "biology" represented a natural development after the preliminary phase of systematics and geographical ornithology, in fact a small group campaigned to bring the priorities of population ecology, behavior, and selection theory into the journal and British ornithology more generally. The problems involved in this transition highlight the importance of methodological and institutional context in determining and reinforcing appropriate research programs for ornithologists. Comparing the discipline-building rhetoric of moderns with the contents of the past illustrates how modern evaluations of 19th century research programs have been enmeshed in ornithologists' endeavors to forge new identities for traditional disciplines. (shrink)

It is an unfortunate fact of academic life that there is a sharp divide between science and philosophy, with scientists often being openly dismissive of philosophy, and philosophers being equally contemptuous of the naivete ́ of scientists when it comes to the philosophical underpinnings of their own discipline. In this paper I explore the possibility of reducing the distance between the two sides by introducing science students to some interesting philosophical aspects of research in evolutionary biology, using biological theories of (...) the origin of religion as an example. I show that philosophy is both a discipline in its own right as well as one that has interesting implications for the understanding and practice of science. While the goal is certainly not to turn science students into philoso- phers, the idea is that both disciplines cannot but benefit from a mutual dialogue that starts as soon as possible, in the classroom. (shrink)

The aim of the present volume is to draw the balance of naturalism's success so far.

The evolutionary synthesis of the 1930s and 1940s integrated the study of biological microevolution and biological macroevolution into the theoretically consistent and hugely productive field of evolutionary biology. A similar synthesis has yet to occur for the study of culture, and the social sciences remain fragmented and theoretically incompatible. Here, it is suggested that a Darwinian theory of cultural evolution can promote such a synthesis. Earlier non-Darwinian theories of cultural evolution, such as progress theories, lacked key elements of a Darwinian (...) theory of cultural evolution that are necessary to promote a synthesis, while other contemporary theories of cultural evolution, such as memetics, make too stringent neo-Darwinian assumptions that are inconsistent with evidence regarding cultural transmission. Several examples are given which indicate the beginnings of an evolutionary synthesis for culture, where patterns of cultural macroevolution have been explained in terms of underlying cultural microevolutionary forces. Finally, it is argued that experimental simulations of cultural evolution can play an important role in this emerging synthesis. (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)

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)

Following Mayr (1961) evolutionary biologists often maintain that the hallmark of biology is its evolutionary perspective. In this view, biologists distinguish themselves from other natural scientists by their emphasis on why-questions. Why-questions are legitimate in biology but not in other natural sciences because of the selective character of the process by means of which living objects acquire their characteristics. For that reason, why-questions should be answered in terms of natural selection. Functional biology is seen as a reductionist science that applies (...) physics and chemistry to answer how-questions but lacks a biological point of view of its own. In this paper I dispute this image of functional biology. A close look at the kinds of issues studied in biology and at the way in which these issues are studied shows that functional biology employs a distinctive biological perspective that is not rooted in selection. This functional perspective is characterized by its concern with the requirements of the life-state and the way in which these are met. (shrink)

We suggest that human culture exhibits key Darwinian evolutionary properties, and argue that the structure of a science of cultural evolution should share fundamental features with the structure of the science of biological evolution. This latter claim is tested by outlining the methods and approaches employed by the principal subdisciplines of evolutionary biology and assessing whether there is an existing or potential corresponding approach to the study of cultural evolution. Existing approaches within anthropology and archaeology demonstrate a good match with (...) the macroevolutionary methods of systematics, paleobiology, and biogeography, whereas mathematical models derived from population genetics have been successfully developed to study cultural microevolution. Much potential exists for experimental simulations and field studies of cultural microevolution, where there are opportunities to borrow further methods and hypotheses from biology. Potential also exists for the cultural equivalent of molecular genetics in “social cognitive neuroscience,” although many fundamental issues have yet to be resolved. It is argued that studying culture within a unifying evolutionary framework has the potential to integrate a number of separate disciplines within the social sciences. (shrink)

A standard norm of reaction (NoR) is a graphical depiction of the phenotypic value of some trait of an individual genotype in a population as a function of an environmental parameter. NoRs thus depict the phenotypic plasticity of a trait. The topological properties of NoRs for sets of different genotypes can be used to infer the presence of (non-linear) genotype-environment interactions. While it is clear that many NoRs are adaptive, it is not yet settled whether their evolutionary etiology should be (...) explained by selection on the mean phenotypic trait values in different environments or whether there are specific genes conferring plasticity. If the second alternative is true the NoR is itself an object of selection. Generalized NoRs depict plasticity at the level of populations or subspecies within a species, species within a genus, or taxa at higher levels. Historically, generalized NoRs have routinely been drawn though rarely explicitly recognized as such. Such generalized NoRs can be used to make evolutionary inferences at higher taxonomic levels in a way analagous to how standard NoRs are used for microevolutionary inferences. (shrink)

The word ``deme'' was coined by the botanists J.S.L. Gilmour and J.W.Gregor in 1939, following the pattern of J.S. Huxley's ``cline''. Its purposewas not only to rationalize the plethora of terms describing chromosomaland genetic variation, but also to reduce hostility between traditionaltaxonomists and researchers on evolution, who sometimes scorned eachother's understanding of species. A multi-layered system of compoundterms based on deme was published by Gilmour and J. Heslop-Harrison in1954 but not widely used. Deme was adopted with a modified meaning byzoologists (...) leading the evolutionary synthesis – Huxley, Simpson, Wright,and Mayr. Connections are shown between Gilmour's ideas around definingthe deme, his role in founding the Systematics Association, and his chapter``Taxonomy and Philosophy'' in the book The New Systematics. Thishistorical episode raises questions about the role of carefully-definedwords in scientific practice. (shrink)

At the beginning of the 1950s most students of animal behavior in Britain saw the instinct concept developed by Konrad Lorenz in the 1930s as the central theoretical construct of the new ethology. In the mid 1950s J.B.S. Haldane made substantial efforts to undermine Lorenz''s status as the founder of the new discipline, challenging his priority on key ethological concepts. Haldane was also critical of Lorenz''s sharp distinction between instinctive and learnt behavior. This was inconsistent with Haldane''s account of the (...) evolution of language, and, according to Haldane, inconsistent with elementary genetics. British attitudes to the instinct concept changed dramatically in the wake of Daniel S. Lehraman''s 1953 critique of Lorenz, and by the 1960s Lorenz drew a clear distinction between his own views and those of the English-speaking ethologists. The inconsistencies between Lorenz''s ideas and the trends in contemporary evolutionary genetics that are reflected in Haldane''s critiques may help to explain why the Lorenzian instinct concept was unable to maintain itself in Britian. (shrink)

I introduce a range of examples of different causal hypotheses about human mate selection. The hypotheses I focus on come from evolutionary psychology, fluctuating asymmetry research and chemical signaling research. I argue that a major obstacle facing an integrated biology of human behavior is the lack of a causal framework that shows how multiple proximate causal mechanisms can act together to produce components of our behavior.

The field of the philosophy of biology is flourishing in its aim to evaluate and rethink the view inherited from the previous century ---the Modern Synthesis. Different research areas and theories have come to the fore in the last decades in order to account for different biological phenomena that, in the first instance, fall beyond the explanatory scope of the Modern Synthesis. This thesis is anchored and motivated by this revolt in the philosophy of biology. -/- The central target in (...) this context is the possibility of naturalizing teleology, a classical nightmare for the history of biology itself. This requires, principally, understanding the causes of teleological explanations without assuming an unfashioned backward causation of sorts. As the riddles of teleological explanations are about their temporal dimension, I analyze different temporal scales of biological processes: evolutionary, developmental, and physiological. -/- The first one is the one defended in the context of the Modern Synthesis. As expected, one of the aims of this thesis is to evaluate the adequacy of an evolutionary account of teleological explanation. The scrutiny is negative. Evolutionary explanations in the context of the Modern Synthesis lack the necessary causal roots to naturalize teleology. Concerning the physiological scale, a long tradition pushed up by Kant and the organicist movement in the 20th century allows us to better understand how teleological explanations can be naturalized in physiological process. The key notions in this temporal scale are self-organization and the recursive, looped character of physiological process. While the physiological scale may be suitably accounted by contemporaries views, such as Autonomous Systems Theory, different central teleological phenomena remain unexplained from a purely physiological perspective. In particular, different issues concerning the (adaptive) construction of organism ---such as plasticity, robustness, variation, novelty, inheritance--- deserve an ontogenetic analysis. -/- The principal aim of this thesis is to provide a theory of teleological development that falls beyond the Modern Synthesis' framework and is prompted by different insights from the history of biology. I call it Agential Teleosemantics. It rests on two central pillars. First, that developmental processes, beyond any gene-centered stance, can be understood in informational terms; i.e. developmental processes are about the interaction of developmental resources conveying biological information. The second ingredient is agentivity, namely the idea that development is regulated by an agentive system according to the adaptiveness of the phenotypic outcomes produced. The role of agency in Agential Teleosemantic is equivalent to the role of genes in the Modern Synthesis: it is responsible for explaining the order and the adaptive complexity in the living realm. -/- The second target of this thesis regards the possibility of naturalizing intentional explanations in cognitive science. The central project involved in such an aim is known as teleosemantics. Classical teleosemantics however is etiological: it explains the teleofunctions of representational systems in terms of evolutionary processes. The different disputes in the contemporary philosophy of biology provide two insights to analyze teleosemantics in cognitive science. First, the challenges against the Modern Synthesis must be extended to the evolutionary approach of etiological teleosemantics. Second, as Agential Teleosemantics suggests an alternative source for teleofunctions ---ontogeny, I offer an attempt to integrate Agential Teleosemantics into cognitive science in order to provide an alternative teleosemantic project to understand intentional explanations in cognitive science. (shrink)

Biologists explain organisms’ behavior not only as having been programmed by genes and shaped by natural selection, but also as the result of an organism’s agency: the capacity to react to environmental changes in goal-driven ways. The use of such ‘agential explanations’ reopens old questions about how justified it is to ascribe agency to entities like bacteria or plants that obviously lack rationality and even a nervous system. Is organismic agency genuinely ‘real’ or is it just a useful fiction? In (...) this paper we focus on two questions: whether agential explanations are to be interpreted ontically, and whether they can be reduced to non-agential explanations (thereby dispensing with agency). The Kantian approach we identify interprets agential explanations non-ontically, yet holds agency to be indispensable. Attributing agency to organisms is not to be taken literally in the way we attribute physical properties such as mass or acceleration, but nor is it a mere heuristic or predictive tool. Rather, it is an inevitable consequence of our own rational capacity: as long as we are rational agents ourselves, we cannot avoid seeing agency in organisms. (shrink)

The extended evolutionary synthesis intends to offer a new framework for understanding evolution based mainly on empirical and theoretical findings of current studies, including heredity and evolutionary developmental biology. In this essay, we present and develop the following objections about the terminology associated with the EES literature: despite using the term "extension," EES protagonists claim new evolutionary processes, reformulate conceptual networks, and modify central assumptions of the evolutionary synthesis. Therefore, the difference between ES and EES should not be described in (...) terms of a set-subset relationship ; despite using the term "synthesis," the EES leads, at least in the short term, to a pluralism of approaches in evolutionary biology. Thus, we argue that the EES should not be described as a synthesis, but as a broad and plural interpretative framework encompassing different approaches. These objections are not directed to the proposed changes, but to the interpretation of these changes as a mere expansion of the previous evolutionary framework, as well as to the interpretation of the EES as a synthesis. Based on these objections, we seek to present some explanations for the use of the term "extended synthesis" among its protagonists. (shrink)

In evolutionary theory, we generally believe that the evolution of life is from simple to complex, from single to diverse, and from lower to higher. Thus, the idea of evolutionary progress appears obvious. However, in contemporary academic circles, some biologists and philosophers challenge this idea. Among them, Gould is the most influential. This paper first describes Gould’s seven arguments against evolutionary progress, i.e., the human arrogance argument, anthropocentric argument, no inner thrust argument, no biological base argument, extreme contingency argument, statistical (...) error argument, and bacteria (other than human beings) ruling the earth argument. Gould’s arguments against evolutionary progress have great influence in contemporary evolutionary theory. Thus, it is necessary to conduct a careful philosophical analysis of each of Gould’s arguments to reveal his philosophical mistakes. This research contends that Gould’s arguments against evolutionary progress are invalid. (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)

Although there is no in-principle impediment to an EvoDevo of behavior, such an endeavor is not as straightforward as one might think; many of the key terms and concepts used in EvoDevo are tailored to suit its traditional focus on morphology, and are consequently difficult to apply to behavior. In this light, the application of the EvoDevo conceptual toolkit to the behavioral domain requires the establishment of a set of tractable concepts that are readily applicable to behavioral characters. Here, I (...) begin the type of theoretical work that needs to be undertaken in order to achieve this, focusing in particular on the key concept of “novelty.” Building on existing criteria used for the identification of behavioral homology from behavioral ecology, I develop a set of operational criteria for identifying novelty in the behavioral domain. These criteria provide a conceptual foundation for the study of novelty in behavioral traits. (shrink)

The so-called “Baldwin Effect” has been studied for years in the fields of Artificial Life, Cognitive Science, and Evolutionary Theory across disciplines. This idea is often conflated with genetic assimilation, and has raised controversy in trans-disciplinary scientific discourse due to the many interpretations it has. This paper revisits the “Baldwin Effect” in Baldwin’s original spirit from a joint historical, theoretical and experimental approach. Social Heredity – the inheritance of cultural knowledge via non-genetic means in Baldwin’s term – is also taken (...) into consideration. I shall argue that the Baldwin Effect can occur via social heredity without necessity for genetic assimilation. Computational experiments are carried out to show that when social heredity is permitted with high fidelity, there is no need for the assimilation of acquired characteristics; instead the Baldwin Effect occurs as promoting more plasticity to facilitate future intelligence. The role of mind and intelligence in evolution and its implications in an extended synthesis of evolution are briefly discussed. (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)

Recent debates between proponents of the modern evolutionary synthesis (the standard model in evolutionary biology) and those of a possible extended synthesis are a good example of the fascinating tangle among empirical, theoretical, and conceptual or philosophical matters that is the practice of evolutionary biology. In this essay, we briefly discuss two case studies from this debate, highlighting the relevance of philosophical thinking to evolutionary biologists in the hope of spurring further constructive cross-pollination between the two fields.

This paper argues that the Extended Synthesis, ecological information, and biosemiotics are complementary approaches whose engagement will help us explain the organism-environment interaction at the cognitive level. The Extended Synthesis, through niche construction theory, can explain the organism-environment interaction at an evolutionary level because niche construction is a process guided by information. We believe that the best account that defines information at this level is the one offered by biosemiotics and, within all kinds of biosemiotic information available, we believe that (...) ecological information is the best candidate for making sense of the organism-environment relation at the cognitive level. This entanglement of biosemiotics, ecological information and the Extended Synthesis is promising for understanding the multidimensional character of the organism-environment reciprocity as well as the relation between evolution, cognition, and meaning. (shrink)

This Ph.D. thesis provides a pilosophical account of the structure of the evolutionary synthesis of the 1930s and 40s. The first, more historical part analyses how classical genetics came to be integrated into evolutionary thinking, highlighting in particular the importance of chromosomal mapping of Drosophila strains collected in the wild by Dobzansky, but also the work of Goldschmidt, Sumners, Timofeeff-Ressovsky and others. The second, more philosophical part attempts to answer the question wherein the unity of the synthesis consisted. I argue (...) that it exemplifies a weak of form of explanatory unification. (shrink)

Bergmann’s rule and Allen’s rule played important roles in mid-twentieth century discussions of adaptation, variation, and geographical distribution. Although inherited from the nineteenth-century natural history tradition these rules gained significance during the consolidation of the modern synthesis as evolutionary theorists focused attention on populations as units of evolution. For systematists, the rules provided a compelling rationale for identifying geographical races or subspecies, a function that was also picked up by some physical anthropologists. More generally, the rules provided strong evidence for (...) adaptation by natural selection. Supporters of the rules tacitly, or often explicitly, assumed that the clines described by the rules reflected adaptations for thermoregulation. This assumption was challenged by the physiologists Laurence Irving and Per Scholander based on their arctic research conducted after World War II. Their critique spurred a controversy played out in a series of articles in Evolution, in Ernst Mayr’s Animal Species and Evolution, and in the writings of other prominent evolutionary biologists and physical anthropologists. Considering this episode highlights the complexity and ambiguity of important biological concepts such as adaptation, homeostasis, and self-regulation. It also demonstrates how different disciplinary orientations and styles of scientific research influenced evolutionary explanations, and the consequent difficulties of constructing a truly synthetic evolutionary biology in the decades immediately following World War II. (shrink)

The dependence on history of both present and future dynamics of life is a common intuition in biology and in humanities. Historicity will be understood in terms of changes of the space of possibilities as well as by the role of diversity in life’s structural stability and of rare events in history formation. We hint to a rigorous analysis of “path dependence” in terms of invariants and invariance preserving transformations, as it may be found also in physics, while departing from (...) the physico-mathematical analyses. The idea is that the invariant traces of the past under organismal or ecosystemic transformations contribute to the understanding of present and future states of affairs. This yields a peculiar form of unpredictability in biology, at the core of novelty formation: the changes of observables and pertinent parameters may depend also on past events. In particular, in relation to the properties of synchronic measurement in physics, the relevance of diachronic measurement in biology is highlighted. This analysis may a fortiori apply to cognitive and historical human dynamics, while allowing to investigate some general properties of historicity in biology. (shrink)

The 1981 Dahlem conference was a catalyst for contemporary evolutionary developmental biology (Evo-devo). This introductory chapter rehearses some of the details of the history surrounding the original conference and its associated edited volume, explicates the philosophical problem of conceptual change that provided the rationale for a workshop devoted to evaluating the epistemic revisions and transformations that occurred in the interim, explores conceptual change with respect to the concept of evolutionary novelty, and highlights some of the themes and patterns in the (...) different contributions to the present volume, Conceptual Change in Biology: Scientific and Philosophical Perspectives on Evolution and Development. (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)

The concept of adaptation is employed in many fields such as biology, psychology, cognitive sciences, robotics, social sciences, even literacy and art,1 and its meaning varies quite evidently according to the particular research context in which it is applied. We expect to find a particularly rich catalogue of meanings within evolutionary biology, where adaptation has held a particularly central role since Darwin’s The Origin of Species (1859) throughout important epistemological shifts and scientific findings that enriched and diversified the concept. Accordingly, (...) a conceptual taxonomy of adaptation in evolutionary biology may help to disambiguate it. Interdisciplinary researches focused on adaptation would benefit from such a result. In the present work we recognize and define seven different meanings of adaptation: (1) individual fitness; (2) adaptation of a population; (3) adaptation as the process of natural selection; (4) adaptive traits; (5) molecular adaptation; (6) adaptation as structural tinkering; (7) plasticity. For convenience here, we refer to them as W-, P-, NS-, T-, M-, S- and PL-ADAPTATION. We present the seven meanings in some detail, hinting at their respective origins and conceptual developments in the history of evolutionary thought (references are offered for further deepening). However, it is important to point out that evolution researchers seldom if ever refer to a single meaning purified from the others. This applies also to the authors we cite as representatives of one of the seven meanings. In Discussion and Conclusion draw from our work some future perspectives for adaptation within evolutionary biology. (shrink)

Comparative biology is afield that deals with morphology. Johann Wolfgang von Goethe recognised comparative biology, not as a passive science obsessed with counting similarities as it is today, but as an active field wherein he sought to perceive the inter-relationships of individual organisms to the organic whole, which he termed the archetype. I submit that Goethe's archetype and his application of a technique termed the Anschauung are rigorous and significant ways to conduct delicate empiricism in comparative biology. The future of (...) comparative biology lies in the use of the Anschauung to communicate the archetype as a set of inter-relationships of homologues that we perceive intuitively. In this essay I present how the extension of our own intuitive perception forms the foundations of a method for seeing and discovering the archetype in comparative biology. (shrink)

In recent years, several prominent biologists have pointed to the relatively new field of evolutionary developmental biology (EvoDevo) as evidence of an Extended Synthesis in evolutionary biology. More particularly, these biologists claim that theoretical and empirical EvoDevo research is extending the Modern Synthesis framework of evolutionary theory through investigation of evolutionarily important concepts that are not part of the framework developed during the 20th century. To describe the current changes in evolutionary biology as an Extended Synthesis, however, is incorrect. Through (...) review of Extended Synthesis arguments and analysis of the same biological concepts used to support these arguments, I argue that the foundation of the Modern Synthesis framework, theoretical population genetics, faces significant, perhaps insurmountable challenges from the concepts highlighted by EvoDevo research. As the foundation of the Modern Synthesis framework will require considerable remodeling—if possible—in light of the concepts emphasized by EvoDevo, it is incorrect to describe the ongoing changes in evolutionary biology as an Extended Synthesis. (shrink)

Starting from the early decades of the twentieth century, evolutionary biology began to acquire mathematical overtones. This took place via the development of a set of models in which the Darwinian picture of evolution was shown to be consistent with the laws of heredity discovered by Mendel. The models, which came to be elaborated over the years, define a field of study known as population genetics. Population genetics is generally looked upon as an essential component of modern evolutionary theory. This (...) article deals with a famous dispute between J. B. S. Haldane, one of the founders of population genetics, and Ernst Mayr, a major contributor to the way we understand evolution. The philosophical undercurrents of the dispute remain relevant today. Mayr and Haldane agreed that genetics provided a broad explanatory framework for explaining how evolution took place but differed over the relevance of the mathematical models that sought to underpin that framework. The dispute began with a fundamental issue raised by Mayr in 1959: in terms of understanding evolution, did population genetics contribute anything beyond the obvious? Haldane's response came just before his death in 1964. It contained a spirited defense, not just of population genetics, but also of the motivations that lie behind mathematical modelling in biology. While the difference of opinion persisted and was not glossed over, the two continued to maintain cordial personal relations. (shrink)

The explanatory role of natural selection is one of the long-term debates in evolutionary biology. Nevertheless, the consensus has been slippery because conceptual confusions and the absence of a unified, formal causal model that integrates different explanatory scopes of natural selection. In this study we attempt to examine two questions: (i) What can the theory of natural selection explain? and (ii) Is there a causal or explanatory model that integrates all natural selection explananda? For the first question, we argue that (...) five explananda have been assigned to the theory of natural selection and that four of them may be actually considered explananda of natural selection. For the second question, we claim that a probabilistic conception of causality and the statistical relevance concept of explanation are both good models for understanding the explanatory role of natural selection. We review the biological and philosophical disputes about the explanatory role of natural selection and formalize some explananda in probabilistic terms using classical results from population genetics. Most of these explananda have been discussed in philosophical terms but some of them have been mixed up and confused. We analyze and set the limits of these problems. (shrink)

Darwin proposed that evolutionary novelties are environmentally induced in organisms “constitutionally” sensitive to environmental change, with selection effective owing to the inheritance of constitutional responses. A molecular theory of inheritance, pangenesis , explained the cross‐generational transmission of environmentally induced traits, as required for evolution by natural selection. The twentieth‐century evolutionary synthesis featured mutation as the source of novelty, neglecting the role of environmental induction. But current knowledge of environmentally sensitive gene expression, combined with the idea of genetic accommodation of mutationally (...) and environmentally induced change, supports a revival of Darwin's original theory that is consistent with modern molecular and population genetics. †To contact the author, please write to: Smithsonian Tropical Research Institute, c/o Escuela de Biología, Universidad de Costa Rica, Costa Rica; e‐mail: [email protected]. (shrink)

Organic selection (the Baldwin Effect) by which an environmentally elicitedphenotypic adaptation comes under genotypic control following selectionwas proposed independently in 1896 by the psychologists James Baldwinand Conwy Lloyd Morgan and by the paleontologist Henry Fairfield Osborn.Modified forms of organic selection were proposed as autonomization bySchmalhausen in 1938, as genetic assimilation by Waddington in 1942, andas an explanation for evolution in changing environments or for speciationby Matsuda and West-Eberhard in the 1980s. Organic selection as amechanism mediating proximate environmental effects on the (...) evolution ofmorphology and behaviour is the topic of this essay. Discussion includesthe context in which organic selection was proposed, Lamarckian or neo-Lamarckian implications of organic selection, Waddingtons experimentalstudies demonstrating the existence and efficacy of genetic assimilation,stabilizing selection and norms of reaction favoured by Schmalhausen, andMatsudas search for a mechanism of organic selection in endocrine changesand in heterochrony. (shrink)

This paper analyzes the development of evolutionary theory in the period from 1918 to 1932. It argues that: (i) Fisher's work in 1918 constituted a not fully satisfactory reduction of biometry to Mendelism; (ii) there was a synthesis in the 1920s but that this synthesis was mainly one of classical genetics with population genetics, with Haldane's The Causes of Evolution being its founding document; (iii) the most important achievement of the models of theoretical population genetics was to show that natural (...) selection sufficed as a mechanism for evolution; and (iv) Haldane formulated a prospective evolutionary theory in the 1920s whereas Fisher and Wright formulated retrospective theories of evolutionary history. (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)