Evolutionary Theory and the Social uses of Biology

Stephen Jay Gould is rightly remembered for many different kinds of contributions to our intellectual life. I focus on his criticisms of uses of evolutionary ideas to defend inegalitarian doctrines and on his attempts to expand the framework of Darwinian evolutionary theory. I argue that his important successes in the former sphere are applications of the idea of local critique, grounded in careful attention to the details of the inegalitarian proposals. As he became more concerned with the second project, Gould was inclined to suggest that the abuses of evolutionary ideas rested on an insufficiently expanded Darwinism. I suggest that what is valuable in Gould's contribution to general evolutionary theory is the original claim about punctuated equilibrium (advanced, with Niles Eldredge in1972), and the careful defense of that claim through the accumulation of paleontological evidence. I try to show that the more ambitious program of a hierarchical expansion of neo-Darwinism is misguided, and that the endeavor to go beyond local critique fails.     

Gould on Laws in Biological Science

Are there laws in evolutionary biology? Stephen J. Gould has argued that there are factors unique to biological theorizing which prevent the formulation of laws in biology, in contradistinction to the case in physics and chemistry. Gould offers the problem of ’’complexity‘‘ as just such a fundamental barrier to biological laws in general, and to Dollo‘s Law in particular. But I argue that Gould fails to demonstrate: (1) that Dollo‘s Law is not law-like, (2) that the alleged failure of Dollo‘s Law demonstrates why there cannot be laws in biological science, and (3) that ’’complexity‘‘ is a fundamental barrier to nomologicality.     

Constraints and Spandrels in Gould's Structure of Evolutionary Theory

Gould's Structure ofEvolutionary Theory argues that Darwinism hasundergone significant revision. Although Gouldsucceeds in showing that hierarchicalapproaches have expanded Darwinism, hiscritique of adaptationism is less successful. Gould claims that the ubiquity of developmentalconstraints and spandrels has forced biologiststo soften their commitment to adaptationism. Iargue that Gould overstates his conclusion; hisprincipal claims are compatible with at leastsome versions of adaptationism. Despite thisweakness, Gould's discussion of adaptationism –particularly his discussions of the exaptivepool and cross-level spandrels – shouldprovoke new work in evolutionary theory and thephilosophy of biology.     

Organisms as natural purposes: the contemporary evolutionary perspective

Kant's conception of organisms as natural purposes raises a challenge to the adequacy of mechanistic explanation in biology. Certain features of organisms appear to be inexplicable by appeal to mechanical law alone. Some biological phenomena, it seems, can only be accounted for teleologically. Contemporary evolutionary biology has by and large ignored this challenge. It is widely held that Darwin's theory of natural selection gives us an adequate, wholly mechanical account of the nature of organisms. In contemporary biology, the category of the organism plays virtually no explanatory role. Contemporary evolutionary biology is a science of sub-organismal entities-replicators. I argue that recent advances in developmental biology demonstrate the inadequacy of sub-organismal mechanism. The category of the organism, construed as a 'natural purpose' should play an ineliminable role in explaining ontogenetic development and adaptive evolution. According to Kant the natural purposiveness of organisms cannot be demonstrated to be an objective principle in nature, nor can purposiveness figure in genuine explain. I attempt to argue, by appeal to recent work on self-organization, that the purposiveness of organisms is a natural phenomenon, and, by appeal to the apparatus of invariance explanation, that biological purposiveness provides genuine, ineliminable biological explanations.     

The Use and Abuse of Sir Karl Popper

Karl Popper has been one of the few philosophers of sciences who has influenced scientists. I evaluate Popper's influence on our understanding of evolutionary theory from his earliest publications to the present. Popper concluded that three sorts of statements in evolutionary biology are not genuine laws of nature. I take him to be right on this score. Popper's later distinction between evolutionary theory as a metaphysical research program and as a scientific theory led more than one scientist to misunderstand his position on evolutionary theory as a scientific theory. In his later work Popper also introduced what he took to be improvements of evolutionary theory. Thus far these improvements have had almost no influence on evolutionary biology. I conclude by examining the influence of Popper on the reception of cladistic analysis.     

Why don't zebras have machine guns? Adaptation, selection, and constraints in evolutionary theory

In an influential paper, Stephen Jay Gould and Richard Lewontin (1979) contrasted selection-driven adaptation with phylogenetic, architectural, and developmental constraints as distinct causes of phenotypic evolution. In subsequent publications Gould (e.g., 1997a,b, 2002) has elaborated this distinction into one between a narrow "Darwinian Fundamentalist" emphasis on "external functionalist" processes, and a more inclusive "pluralist" emphasis on "internal structuralist" principles. Although theoretical integration of functionalist and structuralist explanations is the ultimate aim, natural selection and internal constraints are treated as distinct causes of evolutionary change. This distinction is now routinely taken for granted in the literature in evolutionary biology. I argue that this distinction is problematic because the effects attributed to non-selective constraints are more parsimoniously explained as the ordinary effects of selection itself. Although it may still be a useful shorthand to speak of phylogenetic, architectural, and developmental constraints on phenotypic evolution, it is important to understand that such "constraints" do not constitute an alternative set of causes of evolutionary change. The result of this analysis is a clearer understanding of the relationship between adaptation, selection and constraints as explanatory concepts in evolutionary theory.     

ADAPTIONISM—30 YEARS AFTER GOULD AND LEWONTIN

Gould and Lewontin's 30-year-old critique of adaptionism fundamentally changed the discourse of evolutionary biology. However, with the influx of new ideas and scientific traditions from genomics into evolutionary biology, the old adaptionist controversies are being recycled in a new context. The insight gained by evolutionary biologists, that functional differences cannot be equated to adaptive changes, has at times not been appreciated by the genomics community. In this comment, I argue that even in the presence of both functional data and evidence for selection from DNA sequence data, it is still difficult to construct strong arguments in favor of adaptation. However, despite the difficulties in establishing scientific arguments in favor of specific historic evolutionary events, there is still much to learn about evolution from genomic data.     

San Marco and evolutionary biology

Gould and Lewontin use San Marco, Venice, to criticise the adaptationist program in biology. Following their lead, the architectural term “spandrel” is now widely used in biology to denote a feature that is a necessary byproduct of other aspects of the organism. I review the debate over San Marco and argue that the spandrels are not necessary in the sense originally used by Gould and Lewontin. I conclude that almost all the claims that Gould makes about San Marco are wrong and that it is reasonable to view the architectural spandrel as an adaptation. The spandrels example has not provided a good illustration of why adaptive explanations should be avoided. In fact, it can be used as an example of how adaptive explanations can be dismissed even when there is evidence in their favour. I also discuss the use of the concept of a spandrel in biology.     

Le dossier Vavilov

Vavilov’s dossier. Gould revived the memory of N.I. Vavilov, a victim of the Stalinian system and misjudged among occidental evolutionists. His contribution is impressive in applied research (phytogeography, his list of world-wide plant resources, a unique collection of germplasms intact and always available) as well as in theoretical research work on artificial selection, immunitary relationships between parasite and plant, the bases of his Law of homologous series in hereditary variations, and centers of origin of cultivated plants. Darwinian concept of natural selection were essential for him, but he considered that the evolutionary changes were not only produced by random variations, but by preset channels, recognising the internal constraints of heredity. His heritage has always been maintained in his Institutes. His Evolutionary theories are now confirmed by molecular genetics and systematics. S.J. Gould was the first to revive Vavilov’s memory and scientific importance. During his studies on the gastropod Cerion Gould recognised the balance between external and internal constraints in Evolution. To cite this article: M. Debrenne, F. Debrenne, C. R. Palevol 2 (2003).     

Urban evolutionary ecology brings exaptation back into focus

The contribution of pre-existing phenotypic variation to evolution in novel environments has long been appreciated. Nevertheless, evolutionary ecologists have struggled with communicating these aspects of the adaptive process. In 1982, Gould and Vrba proposed terminology to distinguish character states shaped via natural selection for the roles they currently serve (‘adaptations’) from those shaped under preceding selective regimes (‘exaptations’), with the intention of replacing the inaccurate ‘preadaptation’. Forty years later, we revisit Gould and Vrba’s ideas which, while often controversial, continue to be widely debated and highly cited. We use the recent emergence of urban evolutionary ecology as a timely opportunity to reintroduce the ideas of Gould and Vrba as an integrated framework to understand contemporary evolution in novel environments.     

A Philosophical Evaluation of Adaptationism as a Heuristic Strategy

Adaptationism has prompted many a debate in philosophy of biology but the focus is usually on empirical and explanatory issues rather than methodological adaptationism (MA). Likewise, the context of evolutionary biology has provided the grounding for most discussions of the heuristic role of adaptationism. This paper extends the debate by drawing on case studies from physiology and systems biology to discuss the productive and problematic aspects of adaptationism in functional as well as evolutionary studies at different levels of biological organization. Gould and Lewontin’s Spandrels-paper famously criticized adaptationist methodology for implying a risk of generating ‘blind spots’ with respect to non-selective effects on evolution. Some have claimed that this bias can be accommodated through the testing of evolutionary hypotheses. Although this is an important aspect of overcoming the pitfalls of adaptationism, I argue that the issue of methodological biases is broader than the question of testability. I demonstrate the productivity of adaptationist heuristics but also discuss the deeper problematic aspects associated with the imperialistic tendencies of the strong account of MA.     

The proximate/ultimate distinction in the multiple careers of Ernst Mayr

Ernst Mayr's distinction between “ultimate” and “proximate” causes is justly considered a major contribution to philosophy of biology. But how did Mayr come to this “philosophical” distinction, and what role did it play in his earlier “scientific” work? I address these issues by dividing Mayr's work into three careers or phases: 1) Mayr the naturalist/researcher, 2) Mayr the representative of and spokesman for evolutionary biology and systematics, and more recently 3) Mayr the historian and philosopher of biology. If we want to understand the role of the proximate/ultimate distinction in Mayr's more recent career as a philosopher and historian, then it helps to consider hisearlier use of the distinction, in the course of his research, and in his promotion of the professions of evolutionary biology and systematics. I believe that this approach would also shed light on some other important “philosophical” positions that Mayr has defended, including the distinction between “essentialism: and “population thinking.”     

Darwinism after Mendelism: the case of Sewall Wright's intellectual synthesis in his shifting balance theory of evolution (1931)

Historians of science have long been agreeing: what many textbooks of evolutionary biology say, about the histories of Darwinism and the New Synthesis, is just too simple to do justice to the complexities revealed to critical scholarship and historiography. There is no current consensus, however, on what grand narratives should replace those textbook histories. The present paper does not offer to contribute directly to any grand, consensual, narrational goals; but it does seek to do so indirectly by showing how, in just one individual case, details of intellectual biography connect with big picture issues. To this end, I examine here how very diverse scientific and metaphysical commitments were integrated in Sewall Wright's own personal synthesis of biology and philosophy. Taking as the decisive text the short final section of Wright's long 1931 paper on 'Evolution in Mendelian populations,' I examine how his shifting balance theory (SBT) related to his optimum breeding strategy research, his physiological genetics, his general theory of homogenising and heterogenesing causation and his panpsychist view of mind and matter; and I discuss how understanding these relations can clarify Wright's place in the longue durée of evolutionary thought.     

'Molecules and monkeys': George Gaylord Simpson and the challenge of molecular evolution

In this paper, I analyze George Gaylord Simpson's response to the molecularization of evolutionary biology from his unique perspective as a paleontologist. I do so by exploring his views on early attempts to reconstruct phylogenetic relationships among primates using molecular data. Particular attention is paid to Simpson's role in the evolutionary synthesis of the 1930s and 1940s, as well as his concerns about the rise of molecular biology as a powerful discipline and world-view in the 1960s. I argue that Simpson's belief in the supremacy of natural selection as the primary driving force of evolution, as well as his view that biology was a historical science that seeks ultimate causes and highlights contingency, prevented him from acknowledging that the study of molecular evolution was an inherently valuable part of the life sciences.     

Thirty years of <Emphasis Type="Italic">Biology &amp; Philosophy:</Emphasis> philosophy of which biology?

Which domains of biology do philosophers of biology primarily study? The fact that philosophy of biology has been dominated by an interest for evolutionary biology is widely admitted, but it has not been strictly demonstrated. Here I analyse the topics of all the papers published in Biology & Philosophy, just as the journal celebrates its thirtieth anniversary. I then compare the distribution of biological topics in Biology & Philosophy with that of the scientific journal Proceedings of the National Academy of Science of the USA, focusing on the recent period 2003–2015. This comparison reveals a significant mismatch between the distributions of these topics. I examine plausible explanations for that mismatch. Finally, I argue that many biological topics underrepresented in philosophy of biology raise important philosophical issues and should therefore play a more central role in future philosophy of biology.     

Richard Owen, Morphology and Evolution

Richard Owen has been condemned by Darwinians as an anti-evolutionist and an essentialist. In recent years he has been the object of a revisionist analysis intended to uncover evolutionary elements in his scientific enterprise. In this paper I will examine Owen's evolutionary hypothesis and its connections with von Baer's idea of divergent development. To give appropriate importance to Owen's evolutionism is the first condition to develop an up-to-date understanding of his scientific enterprise, that is to disentagle Owen's contribution to the modernization of typology and morphology. I will argue that Owen's Platonic essentialism is rhetorical and incongruous. On the contrary, an interpretation of the archetype based on Aristotle's biological works makes possible a new conception of type, based on a homeostatic mechanism of stability. The renewal of morphology hinges on homological correspondences and a homeostatic process is also the origin of serial and special homology. I will argue that special homology shows an evolutionary orientation insofar as it is a typically inter-specific character while serial homology is determined through an elementary usage of the categories of developmental morphology. This revised version was published online in July 2006 with corrections to the Cover Date.     

Evolutionary convergences and parallelisms: &;their theoretical differences and the difficulty &;of discriminating them in a practical &;phylogenetic context

The importance of evolutionary parallelisms and their differences from evolutionary convergences have been historically underappreciated, as recently noticed in Gould's last book `The structure of evolutionary history'. In that book, Gould make an effort to distinguish and to reinterpret these concepts in the light of the new discoveries of the last decades on developmental biology and genetics, presenting the elegant metaphor of `Pharaonic bricks versus Corinthian columns'. In this paper I will briefly discuss these concepts, and will argue that, despite the advances that have been made to define them in theory, it is rather hard to differentiate them in a practical phylogenetic context. In order to do so, I will provide some few examples from my own empirical studies on the last years of one of the most morphologically and taxonomically diverse groups of Vertebrates, the catfishes.     

From the "Modern Synthesis" to cybernetics: Ivan Ivanovich Schmalhausen (1884-1963) and his research program for a synthesis of evolutionary and developmental biology

Ivan I. Schmalhausen was one of the central figures in the Russian development of the "Modern Synthesis" in evolutionary biology. He is widely cited internationally even today. Schmalhausen developed the main principles of his theory facing the danger of death in the totalitarian Soviet Union. His great services to evolutionary and theoretical biology are indisputable. However, the received view of Schmalhausen's contributions to evolutionary biology makes an unbiased reading of his texts difficult. Here we show that taking all of his works into consideration (including those only available in Russian) paints a much more dynamic and exciting picture of what he tried to achieve. Schmalhausen pioneered the integration of a developmental perspective into evolutionary thinking. A main tool for achieving this was his approach to living objects as complex multi-level self-regulating systems. Schmalhausen put enormous effort into bringing this idea into fruition during the final stages of his career by combining evolutionary theory with cybernetics. His results and ideas remain thought-provoking, and his texts are of more than just historical interest.     

How developmental is evolutionary developmental biology?

Evolutionary developmental biology (evo-devo) offers both an account of developmental processes and also new integrative frameworks for analyzing interactions between development and evolution. Biologists and philosophers are keen on evo-devo in part because it appears to offer a comfort zone between, on the one hand, what some take to be the relative inability of mainstream evolutionary biology to integrate a developmental perspective; and, on the other hand, what some take to be more intractable syntheses of development and evolution. In this article, I outline core concerns of evo-devo, distinguish theoretical and practical variants, and counter Sterelny's recent argument that evo-devo's attention to development, while important, offers no significant challenge to evolutionary theory as we know it. This revised version was published online in July 2006 with corrections to the Cover Date.     

Evolutionary Anatomy of the Primate Cerebral Cortex, 2001 et S.J. Gould : regards croisés

Evolutionary Anatomy of the Primate Cerebral Cortex, 2001 and S.J. Gould: crossed paths. The book Evolutionary Anatomy of the Primate Cerebral Cortex (Falk and Gibson, 2001) is a report of the congress in honor of H.J. Jerison held in USA (1998). Gould wrote the foreword of that book and Jerison its epilogue. For some participants, Gould’s successive works on organisms' evolution have given an important background to their own research. Jerison and Gould met for the first time something like 40 years ago and all along they have shared common scientific interests. In this article, we shall attempt an overview of Evolutionary Anatomy of the Primate Cerebral Cortex, one of the themes close to an ‘influential’ (fide Gould) book entitled: Evolution of the Brain and Intelligence (Jerison 1973). Tendencies in the organization of the brain volume of Vertebrates, and especially in still-living or fossils mammals, is a major question (cf. recent publications and the debate in Nature). If Jerison was a leader for the encephalization concept (sensu 1D, 2D), Gould clearly introduced modern research (nD) on encephalization. From him, the representation through hyperspace could describe the complexity of the paths followed by Vertebrate brain evolution. From this, the new concept of neurotaxonomy (see Thireau et al., Bull. Soc. Zool. France 122 (4) (1997) 393–411) relates encephalic organisation to species taxonomy. So, in the future, paleoneurologists should be using some results from the neurotaxonomy of existing species, for help in completing interpretation of endocranian casts. Throughout this article, special attention is given to Gould’s arguments, proposed in his foreword to Falk et Gibson, 2001. Each of these deserves to be discussed, thus benefiting evolutive neurobiology, and should be completed by the reading of his major contribution to modern biology: The structure of Evolutionary Theory (Gould 2002). To cite this article: L. Thireau, J.-C. Doré, C. R. Palevol 2 (2003).