The Extended Evolutionary Synthesis (EES) debate is gaining ground in contemporary evolutionary biology. In parallel, a number of philosophical standpoints have emerged in an attempt to clarify what exactly is represented by the EES. For Massimo Pigliucci, we are in the wake of the newest instantiation of a persisting Kuhnian paradigm; in contrast, Telmo Pievani has contended that the transition to an EES could be best represented as a progressive reformation of a prior Lakatosian scientific research program, with the extension (...) of its Neo-Darwinian core and the addition of a brand-new protective belt of assumptions and auxiliary hypotheses. Here, we argue that those philosophical vantage points are not the only ways to interpret what current proposals to ‘extend’ the Modern Synthesis-derived ‘standard evolutionary theory’ (SET) entail in terms of theoretical change in evolutionary biology. We specifically propose the image of the emergent EES as a vast network of models and interweaved representations that, instantiated in diverse practices, are connected and related in multiple ways. Under that assumption, the EES could be articulated around a paraconsistent network of evolutionary theories (including some elements of the SET), as well as models, practices and representation systems of contemporary evolutionary biology, with edges and nodes that change their position and centrality as a consequence of the co-construction and stabilization of facts and historical discussions revolving around the epistemic goals of this area of the life sciences. We then critically examine the purported structure of the EES—published by Laland and collaborators in 2015—in light of our own network-based proposal. Finally, we consider which epistemic units of Evo-Devo are present or still missing from the EES, in preparation for further analyses of the topic of explanatory integration in this conceptual framework. (shrink)
In identifying intrinsic molecular chance and extrinsic adaptive pressures as the only causally relevant factors in the process of evolution, the theoretical perspective of the Modern Synthesis had a major impact on the perceived tenability of an ontology of dispositional properties. However, since the late 1970s, an increasing number of evolutionary biologists have challenged the descriptive and explanatory adequacy of this “chance alone, extrinsic only” understanding of evolutionary change. Because morphological studies of homology, convergence, and teratology have revealed a space (...) of possible forms and phylogenetic trajectories that is considerably more restricted than expected, evo-devo has focused on the causal contribution of intrinsic developmental processes to the course of evolution. Evo-devo’s investigation into the developmental structure of the modality of morphology – including both the possibility and impossibility of organismal form – has led to the utilisation of a number of dispositional concepts that emphasise the tendency of the evolutionary process to change along certain routes. In this sense, and in contrast to the perspective of the Modern Synthesis, evo-devo can be described as a “science of dispositions.” This chapter discusses the recent philosophical literature on dispositional properties in evo-devo, exploring debates about both the metaphysical and epistemological aspects of the central dispositional concepts utilised in contemporary evo-devo and addressing the epistemological question of how dispositional properties challenge existing explanatory models in evolutionary biology. (shrink)
Evolutionary developmental biology (Evo-devo) is a vibrant area of contemporary life science that should be (and is) increasingly incorporated into teaching curricula. Although the inclusion of this content is important for biological pedagogy at multiple levels of instruction, there are also philosophical lessons that can be drawn from the scientific practices found in Evo-devo. One feature of particular significance is the interdisciplinary nature of Evo-devo investigations and their resulting explanations. Instead of a single disciplinary approach being the (...) most explanatory or fundamental, different methodologies from biological disciplines must be synthesized to generate empirically adequate explanations. Thus, Evo-devo points toward a non-reductionist epistemology in biology. I review three areas where these synthetic efforts become manifest as a result of Evo-devo’s practices (form versus function reasoning styles; problem-structured investigations; idealizations related to studying model organisms), and then sketch some possible applications to teaching biology. These philosophical considerations provide resources for life science educators to address (and challenge) key aspects of the National Science Education Standards and Benchmarks for Scientific Literacy. (shrink)
Evolutionary developmental biology (Evo-devo) is a vibrant area of contemporary life science that should be (and is) increasingly incorporated into teaching curricula. Although the inclusion of this content is important for biological pedagogy at multiple levels of instruction, there are also philosophical lessons that can be drawn from the scientific practices found in Evo-devo. One feature of particular significance is the interdisciplinary nature of Evo-devo investigations and their resulting explanations. Instead of a single disciplinary approach being the (...) most explanatory or fundamental, different methodologies from biological disciplines must be synthesized to generate empirically adequate explanations. Thus, Evo-devo points toward a non-reductionist epistemology in biology. I review three areas where these synthetic efforts become manifest as a result of Evo-devo's practices (form versus function reasoning styles; problem-structured investigations; idealizations related to studying model organisms), and then sketch some possible applications to teaching biology. These philosophical considerations provide resources for life science educators to address (and challenge) key aspects of the National Science Education Standards and Benchmarks for Scientific Literacy. (shrink)
Evolutionary developmental biology represents a paradigm shift in the understanding of the ontogenesis and evolutionary progression of the denizens of the natural world. Given the empirical successes of the evo-devo framework, and its now widespread acceptance, a timely and important task for the philosophy of biology is to critically discern the ontological commitments of that framework and assess whether and to what extent our current metaphysical models are able to accommodate them. In this paper, I argue that one particular (...) model is a natural fit: an ontology of dispositional properties coherently and adequately captures the crucial casual-cum-explanatory role that the fundamental elements of evo-devo play within that framework. (shrink)
A central reason that undergirds the significance of evo-devo is the claim that development was left out of the Modern synthesis. This claim turns out to be quite complicated, both in terms of whether development was genuinely excluded and how to understand the different kinds of embryological research that might have contributed. The present paper reevaluates this central claim by focusing on the practice of model organism choice. Through a survey of examples utilized in the literature of the Modern (...) synthesis, I identify a previously overlooked feature: exclusion of research on marine invertebrates. Understanding the import of this pattern requires interpreting it in terms of two epistemic values operating in biological research: theoretical generality and explanatory completeness. In tandem, these values clarify and enhance the significance of this exclusion. The absence of marine invertebrates implied both a lack of generality in the resulting theory and a lack of completeness with respect to particular evolutionary problems, such as evolvability and the origin of novelty. These problems were salient to embryological researchers aware of the variation and diversity of larval forms in marine invertebrates. In closing, I apply this analysis to model organism choice in evo-devo and discuss its relevance for an extended evolutionary synthesis. (shrink)
This paper addresses the role of time scales in conceptualizing biological hierarchies. So far, the concept of hierarchies in philosophy of science has been dominated by the idea of composition and parthood, respectively. However, this view does not exhaust the diversity of hierarchical descriptions in the biosciences. Therefore, we highlight a type of hierarchy usually overlooked by philosophers of science. It distinguishes processes based on the different time scales (i.e. rates, frequencies, and rhythms) on which they occur. These time scale (...) hierarchies often are connected with assumptions defended in process ontology. Due to their ability to describe interlevel dynamics of various kinds, we call these hierarchies ‘dynamic hierarchies.’ In order to highlight and discuss their organization, explanatory roles, and epistemic virtues we focus on dynamic hierarchies in developmental biology and evolutionary developmental biology (evo-devo). In these fields, dynamic hierarchies offer crucial complementary information to descriptions of compositional hierarchies. (shrink)
With Carl Gegenbaur and Ernst Haeckel, inspiredby Darwin and the cell theory, comparativeanatomy and embryology became established andflourished in Jena. This tradition wascontinued and developed further with new ideasand methods devised by some of Haeckelsstudents. This first period of innovative workin evolutionary morphology was followed byperiods of crisis and even a disintegration ofthe discipline in the early twentieth century.This stagnation was caused by a lack ofinterest among morphologists in Mendeliangenetics, and uncertainty about the mechanismsof evolution. Idealistic morphology was stillinfluental in (...) Germany, which prevented a fullappreciation of the importance of Darwinstheory of natural selection for comparativemorphology. Evolutionary morphology andembryology failed to contribute significantlyto the modern synthesis of evolutionarybiology, thereby probably delaying theintegration of developmental biology intomodern evolutionary biology. However, Haeckelsstudent Oscar Hertwig, as well as Victor Franzand Alexej N. Sewertzoff from a youngergeneration, all tried to forge their ownsynthetic approaches in which (inspired byHaeckels work) embryology played an importantrole. Important for all three researchers wereattempts to refine, and sometimes redefine, thebiogenetic law, and to find new scientificexplanations for it (and for the manyexceptions to it). Their research was latermore or less forgotten, and had littleinfluence on the architects of the modernsynthesis. As the relationship betweenevolutionary and developmental biology is nowagain rising in importance in the form ofEvo-Devo, we would like to draw attention tohow this earlier research tradition grappledwith similar questions to those now on theagenda, albeit from sometimes quite differentperspectives. (shrink)
Represents the most comprehensive and current survey of the various challenges to the Modern Synthesis theory of evolution. Incorporates a variety of theoretical and disciplinary perspectives, from evolutionary biologists, historians and philosophers of science. These essays constitute the state of the art in the current debate on the status of the Modern Synthesis.
Exploring history pertinent to evolutionary developmental biology (hereafter, Evo-devo) is an exciting prospect given its current status as a cutting-edge field of research. The first and obvious question concerns where to begin searching for materials and sources. Since this new discipline adopts a moniker that intentionally juxtaposes ‘evolution’ and development’, individuals, disciplines, and institutional contexts relevant to the history of evolutionary studies and investigations of ontogeny prompt themselves. Each of these topics has received attention from historians and thus there (...) is both primary and secondary material from which to draw. For example, many historians have documented the historical trajectories of genetics and embryology, their split, and various relations (or lack thereof), especially in the first three decades of the 20th century. (shrink)
In this book Ron Amundson examines two hundred years of scientific views on the evolution-development relationship from the perspective of evolutionary developmental biology. This perspective challenges several popular views about the history of evolutionary thought by claiming that many earlier authors had made history come out right for the Evolutionary Synthesis. The book starts with a revised history of nineteenth-century evolutionary thought. It then investigates how development became irrelevant with the Evolutionary Synthesis. It concludes with an examination of the contrasts (...) that persist between mainstream evolutionary theory and evo-devo. This book will appeal to students and professionals in the philosophy and history of science, and biology. (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)
Explanation in terms of gene regulatory networks has become standard practice in evolutionary developmental biology. In this paper, we argue that GRNs fail to provide a robust, mechanistic, and dynamic understanding of the developmental processes underlying the genotype–phenotype map. Explanations based on GRNs are limited by three main problems: the problem of genetic determinism, the problem of correspondence between network structure and function, and the problem of diachronicity, as in the unfolding of causal interactions over time. Overcoming these problems requires (...) dynamic mechanistic explanations, which rely not only on mechanistic decomposition, but also on dynamic modeling to reconstitute the causal chain of events underlying the process of development. We illustrate the power and potential of this type of explanation with a number of biological case studies that integrate empirical investigations with mathematical modeling and analysis. We conclude with general considerations on the relation between mechanism and process in evo-devo. (shrink)
Although the role of morphology in evolutionary theory remains a subject of debate, assessing the contributions of morphological investigation to evolutionary developmental biology (Evo-devo) is a more circumscribed issue of direct relevance to ongoing research. Historical studies of morphologically oriented researchers and the formation of the Modern Synthesis in the Anglo-American context identify a recurring theme: the synthetic theory of evolution did not capture multiple levels of biological organization. When this feature is incorporated into a philosophical framework for explaining (...) the origin of evolutionary innovations and novelties (a core domain of inquiry in Evo-devo) two specific roles for morphology can be described: (1) the conceptualization and operational identification of the targets of explanation; and (2) the elucidation of causal interactions at higher levels of organization during ontogeny and through evolutionary time. These roles are critical components of any adequate explanation of innovation and novelty though not exhaustive of the parts played by morphology in evolutionary investigation. They also invite reflection on what counts as an evolutionary cause in contemporary evolutionary biology. (shrink)
The emerging discipline of evolutionary developmental biology has opened up many new lines of investigation into morphological evolution. Here I explore how two of the core theoretical concepts in ‘evo-devo’ – modularity and homology – apply to evolutionary psychology. I distinguish three sorts of module – developmental, functional and mental modules and argue that mental modules need only be ‘virtual’ functional modules. Evolutionary psychologists have argued that separate mental modules are solutions to separate evolutionary problems. I argue that the (...) structure of developmental modules in an organism helps determine what counts as a separate evolutionary problem for that organism. I suggest that homology as an organizing principle for research in evolutionary psychology, has been severely neglected in favor of analogy (adaptive function). I consider some arguments suggesting that determining homology is less epistemically demanding than determining adaptive function and argue that psychological categories defined by homology are, in fact, more suitable objects of psychological – and particularly neuropsychological – investigation than categories defined by analogy. (shrink)
The aim of this paper lies in characterizing the explanations and models used in the field of evolutionary developmental biology throughout its history. While manipulative experiments in controlled conditions have been useful to set the bases of the discipline and are still routinely performed, this approach supposes a tension between the reliability and the representativity of the conclusions. Given the recent changes in the understanding of evolutionary phenomena, different authors currently emphasize the need of avoiding excessive simplifications in experimental approaches, (...) incorporating the complexity of the analyzed systems as a relevant trait in the study of biological diversity. On the other hand, the fragile and contingent nature of evolutionary processes and the impossibility of intervening some of these phenomena underscore the importance of non-manipulative models that allow the elucidation of how-possibly mechanisms. We argue that a pluralism characterized by the integration of different kinds of explanations, models and metaphors used in evo-devo allows to exploit their respective advantages, favours interdisciplinarity and can contribute to form a representative description of evolutionary processes without relinquishing the local and detailed study of actual mechanisms. (shrink)
The traditional practice of establishing morphological types and investigating morphological organization has found new support from evolutionary developmental biology (evo-devo), especially with respect to the notion of body plans. Despite recurring claims that typology is at odds with evolutionary thinking, evo-devo offers mechanistic explanations of the evolutionary origin, transformation, and evolvability of morphological organization. In parallel, philosophers have developed non-essentialist conceptions of natural kinds that permit kinds to exhibit variation and undergo change. This not only facilitates a construal (...) of species and higher taxa as natural kinds, but also broadens our perspective on the diversity of kinds found in biology. There are many different natural kinds relevant to the investigative and explanatory aims of evo-devo, including homologues and developmental modules. (shrink)
In the evolutionary biology of the Modern Synthesis the study of patterns refers to how to identify and systematise order in lineages, looking for hierarchies or for branching/splitting events in the tree of life, whereas the resulting order is supposed to be due to underlying processes or mechanisms. But patterns and processes play distinct roles in evo-devo: four different views on the role of patterns and processes in descriptions and explanations of development and evolution: A) transformational; B) generative; C) (...) processual; and D) complex are reviewed in this paper. Then, this discussion is related to two issues in evo-devo: homology and variation. (shrink)
The study of evolutionary developmental biology (“evo‐devo”) has recently experienced a dramatic surge in popularity among researchers and theorists concerned with evolution. However, some biologists and philosophers remain skeptical of the claims of evo‐devo. This paper discusses and responds to the recent high profile criticisms of evo‐devo presented by biologists Hopi E. Hoekstra and Jerry A. Coyne. I argue that their objections are unconvincing. Indeed, empirical research supports the main tenets of evo‐devo, including the claim that morphological evolution is the (...) result of cis ‐regulatory change and the distinction that evo‐devo draws between morphological and physiological traits. *Received January 2008; revised March 2009. †To contact the author, please write to: Department of Philosophy, University of Cincinnati, Cincinnati, OH 45221; e‐mail: firstname.lastname@example.org. (shrink)
Evolutionary developmental biology (“evo-devo”) may provide insights and new methods for studies of cognition and cultural evolution. For example, I propose using cultural selection and individual learning to examine constraints on cultural evolution. Modularity, the idea that traits vary independently, can facilitate evolution (increase “evolvability”), because evolution can act on one trait without disrupting another. I explore links between cognitive modularity, evolutionary modularity, and cultural evolvability. (Published Online November 9 2006).
While the notion of chance has been central in discussions over the probabilistic nature of natural selection and genetic drift, its role in the production of variants on which populational sampling takes place has received much less philosophical attention. This article discusses the concept of chance in evolution in the light of contemporary work in evo-devo. We distinguish different levels at which randomness and chance can be defined in this context, and argue that recent research on variability and evolvability (...) demands a causal understanding of variational probabilities under which development acquires a creative, rather than a constraining role in evolution. We then provide a propensity interpretation of variational probabilities that solves a conceptual confusion between causal properties, variational probabilities and extant variation present in the literature, and explore some metaphysical consequences that follow from our interpretation, specifically with regards to the nature of developmental types. (shrink)
Although the construction of neo-Darwinism grew out of Thomas Hunt Morgan's melding of Darwinism and Mendelism, his evidence did not soley support a model of gradual change. To the contrary, he was confronted with observations that could have led him to a more "evo-devo" understanding of the emergence of novel features. Indeed, since Morgan was an embryologist before he became a fruit-fly geneticist, one would have predicted that the combination of these two lines of research would have resulted in (...) early formulations of concepts relevant to evolutionary developmental biology. It is thus of interest to review Morgan's thought processes and arguments for at first rejecting both Darwinism and Mendelism, and then for later dismissing data that would have yielded a model of rapid morphological change in favor of a model of change based on the accumulation of minor mutations and their morphological consequences. (shrink)
This article reviews the recent reissuing of Richard Owen’s On the Nature of Limbs and its three novel, introductory essays. These essays make Owen’s 1849 text very accessible by discussing the historical context of his work and explaining how Owen’s ideas relate to his larger intellectual framework. In addition to the ways in which the essays point to Owen’s relevance for contemporary biology, I discuss how Owen’s unity of type theory and his homology claims about fins and limbs compare with (...) modern views. While the phenomena studied by Owen are nowadays of major interest to evolutionary developmental biology, research in evo-devo has largely shifted from homology (which was Owen’s concern) towards evolutionary novelty, e.g., accounting for fins as a novelty. Still, I argue that questions about homology are important and raise challenges even for explanations of novelty. (shrink)
El objetivo del presente trabajo es indagar la naturaleza integradora de la biología evolutiva del desarrollo. En particular analizamos las características temporales de los procesos estudiados por diferentes programas de investigación de evo-devo y las comparamos con aquellas propias de los procesos macroevolutivos, microevolutivos y del desarrollo de los organismos. Encontramos que en los principales programas de investigación de evo-devo se recuperan principalmente características propias de los fenómenos macroevolutivos, mientras que en la sub-área de eco-evo-devo se recuperan (...) principalmente características propias de la microevolución.The aim of this study is to investigate the integrative nature of evolutionary developmental biology. We particularly analyzed the temporal characteristics of the processes studied by different research programs of evo-devo and we compared them with those of macroevolution, microevolution and developmentalprocesses of organisms. We found that in the major research programs of evo-devo, macroevolutionary phenomena characteristics are mainly recovered, while in the sub-area of eco-evo-devo microevolutionary characteristics are mainly recovered. (shrink)
This article reviews the recent reissuing of Richard Owen’s On the Nature of Limbs and its three novel, introductory essays. These essays make Owen’s 1849 text very accessible by discussing the historical context of his work and explaining how Owen’s ideas relate to his larger intellectual framework. In addition to the ways in which the essays point to Owen’s relevance for contemporary biology, I discuss how Owen’s unity of type theory and his homology claims about ﬁns and limbs compare with (...) modern views. While the phenomena studied by Owen are nowadays of major interest to evolutionary developmental biology, research in evo-devo has largely shifted from homology (which was Owen’s concern) towards evolutionary novelty, e.g., accounting for ﬁns as a novelty. Still, I argue that questions about homology are important and raise challenges even for explanations of novelty. (shrink)
Sound comparative psychology and modern evolutionary and developmental biology (often called evo-devo) emphasize powerful effects of developmental conditions on the expression of genetic endowment. Both demand that evolutionary theorists recognize these effects. Instead, Tomasello et al. compares studies of normal human children with studies of chimpanzees reared and maintained in cognitively deprived conditions, while ignoring studies of chimpanzees in cognitively appropriate environments.
Abstract A lot of science has passed under the bridge since the classic definition of evolution as a change in gene frequencies in a population became common. Much knowledge has accumulated since then about evolution, heredity, ecology, development, phenotypic plasticity, niche construction and genetic drift. Building on Van Valen’s description of evolution as “the control of development by ecology,” it is suggested that the classic definition be replaced by a updated ‘eco‐evo-evo’ definition of evolution by natural selection which acknowledges this (...) accumulation. (shrink)
Accounting for the evolutionary origins of morphological novelty is one of the core challenges of contemporary evolutionary biology. A successful explanatory framework requires the integration of different biological disciplines, but the relationships between developmental biology and standard evolutionary biology remain contested. There is also disagreement about how to define the concept of evolutionary novelty. These issues were the subjects of a workshop held in November 2009 at the University of Alberta. We report on the discussion and results of this workshop, (...) addressing questions about (i) how to define evolutionary novelty and understand its significance, (ii) how to interpret evolutionary developmental biology as a synthesis and its relation to neo-Darwinian evolutionary theory, and (iii) how to integrate disparate biological approaches in general. (shrink)
The relatively new field of evolutionary developmental biology continues to attract considerable attention from biologists, philosophers, and historians, in part, because work in this field demonstrates that important changes are underway within biology. Though studies of development and evolution were closely connected during the 19th century, continued work in genetics fostered a general split between the two during the first decades of the twentieth century (e.g., Allen 1978; Gilbert 1978; Mayr and Provine 1980; Gilbert, Opitz and..
The homologies of process within morphogenetic fields provide some of the best evidence for evolution—just as skeletal and organ homologies did earlier. Thus, the evidence for evolution is better than ever. The role of natural selection in evolution, how–ever, is seen to play less an important role. It is merely a filter for unsuccessful morphologies generated by development. Population genetics is destined to change if it is not to become as irrelevant to evolution as Newtonian mechanics is to contempo–rary physics.
The homologies of process within morphogenetic fields provide some of the best evidence for evolution—just as skeletal and organ homologies did earlier. Thus, the evidence for evolution is better than ever. The role of natural selection in evolution, however, is seen to play less an important role. It is merely a filter for unsuccessful morphologies generated by development. Population genetics is destined to change if it is not to become as irrelevant to evolution as Newtonian mechanics is to contemporary physics.