By linking the concepts of homology and morphological organization to evolvability, this paper attempts to (1) bridge the gap between developmental and phylogenetic approaches to homology and to (2) show that developmental constraints and natural selection are compatible and in fact complementary. I conceive of a homologue as a unit of morphological evolvability, i.e., as a part of an organism that can exhibit heritable phenotypic variation independently of the organism’s other homologues. An account of homology therefore consists in explaining how (...) an organism’s developmental constitution results in different homologues/characters as units that can evolve independently of each other. The explanans of an account of homology is developmental, yet the very explanandum is an evolutionary phenomenon: evolvability in a character-by-character fashion, which manifests itself in phylogenetic patterns as recognized by phylogenetic approaches to homology. While developmental constraints and selection have often been viewed as antagonistic forces, I argue that both are complementary as they concern different parts of the evolutionary process. Developmental constraints, conceived of as the presence of the same set of homologues across phenotypic change, pertain to how heritable variation can be generated in the first place (evolvability), while natural selection operates subsequently on the produced variation. (shrink)

The discussion presents a framework of concepts that is intended to account for the rationality of semantic change and variation, suggesting that each scientific concept consists of three components of content: 1) reference, 2) inferential role, and 3) the epistemic goal pursued with the concept’s use. I argue that in the course of history a concept can change in any of these components, and that change in the concept’s inferential role and reference can be accounted for as being rational relative (...) to the third component, the concept’s epistemic goal. This framework is illustrated and defended by application to the history of the gene concept. It is explained how the molecular gene concept grew rationally out of the classical gene concept despite a change in reference, and why the use and reference of the contemporary molecular gene concept may legitimately vary from context to context. (shrink)

This introduction to the special section on integration in biology provides an overview of the different contributions. In addition to motivating the philosophical significance of analyzing integration and interdisciplinary research, I lay out common themes and novel insights found among the special section contributions, and indicate how they exhibit current trends in the philosophical study of integration. One upshot of the contributed papers is that there are different aspects to and kinds of integration, so that rather than attempting to offer (...) a universal construal of what integrations is, philosophers have to analyze in concrete cases in what respects particular aspects of scientific theorizing and/or practice are ‘integrative’ and how this instance of integration works and was achieved. (shrink)

The paper works towards an account of explanatory integration in biology, using as a case study explanations of the evolutionary origin of novelties-a problem requiring the integration of several biological fields and approaches. In contrast to the idea that fields studying lower level phenomena are always more fundamental in explanations, I argue that the particular combination of disciplines and theoretical approaches needed to address a complex biological problem and which among them is explanatorily more fundamental varies with the problem pursued. (...) Solving a complex problem need not require theoretical unification or the stable synthesis of different biological fields, as items of knowledge from traditional disciplines can be related solely for the purposes of a specific problem. Apart from the development of genuine interfield theories, successful integration can be effected by smaller epistemic units (concepts, methods, explanations) being linked. Unification or integration is not an aim in itself, but needed for the aim of solving a particular scientific problem, where the problem's nature determines the kind of intellectual integration required. (shrink)

This new work provides an approachable introduction to the complex system that Making It Explicit mapped out.

The practical criteria by which developmental biologists choose their model systems have evolutionary correlates. The result is a sample that is not merely small, but biased in particular ways, for example towards species with rapid, highly canalized development. These biases influence both data collection and interpretation, and our views of how development works and which aspects of it are important.

Many studies of the unification of science focus on the theories of different disciplines. The model for integration is the theory reduction model. This paper argues that the embodiment of theories in scientists, and the institutions in which scientists work and the instruments they employ, are critical to the sort of integration that actually occurs in science. This paper examines the integration of scientific endeavors that emerged in cell biology in the period after World War II when the development of (...) cell fractionation and electron microscopy made serious investigations of cell organelles possible. One surprising feature of such integration is that it generated further disintegration as the new institutions of cell biology separated the practitioners of the new discipline from other, closely related biological disciplines. (shrink)

Among the liveliest disputes in evolutionary biology today are disputes concerning the role of chance in evolution--more specifically, disputes concerning the relative evolutionary importance of natural selection vs. so-called "random drift". The following discussion is an attempt to sort out some of the broad issues involved in those disputes. In the first half of this paper, I try to explain the differences between evolution by natural selection and evolution by random drift. On some common construals of "natural selection", those two (...) modes of evolution are completely indistinguishable. Even on a proper construal of "natural selection", it is difficult to distinguish between the "improbable results of natural selection" and evolution by random drift. In the second half of this paper, I discuss the variety of positions taken by evolutionists with respect to the evolutionary importance of random drift vs. natural selection. I will then consider the variety of issues in question in terms of a conceptual distinction often used to describe the rise of probabilistic thinking in the sciences. I will argue, in particular, that what is going on here is not, as might appear at first sight, just another dispute about the desirability of "stochastic" vs. "deterministic" theories. Modern evolutionists do not argue so much about whether evolution is stochastic, but about how stochastic it is. (shrink)

This paper aims to identify the key characteristics of model organisms that make them a specific type of model within the contemporary life sciences: in particular, we argue that the term “model organism” does not apply to all organisms used for the purposes of experimental research. We explore the differences between experimental and model organisms in terms of their material and epistemic features, and argue that it is essential to distinguish between their representational scope and representational target. We also examine (...) the characteristics of the communities who use these two types of models, including their research goals, disciplinary affiliations, and preferred practices to show how these have contributed to the conceptualization of a model organism. We conclude that model organisms are a specific subgroup of organisms that have been standardized to fit an integrative and comparative mode of research, and that it must be clearly distinguished from the broader class of experimental organisms. In addition, we argue that model organisms are the key components of a unique and distinctively biological way of doing research using models.Keywords: Experimental organism; Genetics; Model organism; Modeling; Philosophy of biology; Representation. (shrink)

The so-called "adaptationism" of mainstream evolutionary biology has been criticized from a variety of sources. One, which has received relatively little philosophical attention, is developmental biology. Developmental constraints are said to be neglected by adaptationists. This paper explores the divergent methodological and explanatory interests that separate mainstream evolutionary biology from its embryological and developmental critics. It will focus on the concept of constraint itself; even this central concept is understood differently by the two sides of the dispute.

Evolutionary developmental biology (or developmental evolution) is in the middle stages of its “development.” Its early ontogeny cannot be traced back to fertilization but pivotal developmental events included Gould’s (1977) treatment of heterochrony, Riedl’s (1978) analysis of “burden”, the Dahlem conference of 1981, a British Society of Developmental Biologists Symposium, as well as books that incorporated developmental genetics into older comparative themes. A major inductive process began with the discovery of widespread phylogenetic conservation in homeobox-containing genes. One interpretation of these (...) early developmental dynamics holds that an increasingly prominent problem agenda (e.g., how do novel structures and functions originate?), inherited from long-standing research programs in comparative embryology, morphology, and paleontology, was handed a powerful set of experimental tools from the lineage of experimental embryology. This allowed for an experimental probing of development and its evolution in an unprecedented fashion. Although the relationship between evolution and development has exhibited a protracted and complex history, we are still in the process of comprehending these early stages of “modern” EvoDevo’s ontogeny. (shrink)

They are correct that punctuated equilibria apply to sexually reproducing organisms and that morphological evolutionary change is regarded as largely (if not exclusively) correlated with speciation events. However, they err in suggesting that we attribute stasis strictly to "developmental constraints," which represent only one of a set of possible mechanisms that we have suggested for the causes of stasis. Others include habitat tracking and the internal structure of species themselves [for example, (2)].

Reductionism encompasses a set of ontological, epistemological, and methodological claims about the relation of different scientific domains. The basic question of reduction is whether the properties, concepts, explanations, or methods from one scientific domain (typically at higher levels of organization) can be deduced from or explained by the properties, concepts, explanations, or methods from another domain of science (typically one about lower levels of organization). Reduction is germane to a variety of issues in philosophy of science, including the structure of (...) scientific theories, the relations between different scientific disciplines, the nature of explanation, the diversity of methodology, and the very idea of theoretical progress, as well as to numerous topics in metaphysics and philosophy of mind, such as emergence, mereology, and supervenience. (shrink)

The volume is a collection of essays devoted to the analysis of scientific change and stability. It explores the balance and tension that exist between commensurability and continuity on the one hand, and incommensurability and discontinuity on the other. Moreover, it discusses some central epistemological consequences regarding the nature of scientific progress, rationality and realism. In relation to these topics, it investigates a number of new avenues, and revisits some familiar issues, with a focus on the history and philosophy of (...) physics, and an emphasis on developments in cognitive sciences as well as on the claims of “new experimentalists”. The book constitutes fully revised versions of papers which were originally presented at the international colloquium held at the University of Nancy, France, in June 2004. (shrink)

ORIGIN OF SPECIES. INTRODUCTION. When on board HMS 'Beagle,' as naturalist, I was ranch struck with certain facts in the distribution of the organic beings ...

How do novel scientific concepts arise? In Creating Scientific Concepts, Nancy Nersessian seeks to answer this central but virtually unasked question in the problem of conceptual change. She argues that the popular image of novel concepts and profound insight bursting forth in a blinding flash of inspiration is mistaken. Instead, novel concepts are shown to arise out of the interplay of three factors: an attempt to solve specific problems; the use of conceptual, analytical, and material resources provided by the cognitive-social-cultural (...) context of the problem; and dynamic processes of reasoning that extend ordinary cognition. Focusing on the third factor, Nersessian draws on cognitive science research and historical accounts of scientific practices to show how scientific and ordinary cognition lie on a continuum, and how problem-solving practices in one illuminate practices in the other. (shrink)

Continuing his exploration of the organization of complexity and the science of design, this new edition of Herbert Simon's classic work on artificial ...

In Darwinism's Struggle for Survival Jean Gayon offers a philosophical interpretation of the history of theoretical Darwinism. He begins by examining the different forms taken by the hypothesis of natural selection in the nineteenth century and the major difficulties which it encountered, particularly with regard to its compatibility with the theory of heredity. He then shows how these difficulties were overcome during the seventy years which followed the publication of Darwin's Origin of Species, and he concludes by analysing the major (...) features of the genetic theory of natural selection, as it developed from 1920 to 1960. This rich and wide-ranging study will appeal to philosophers and historians of science and to evolutionary biologists. (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)

Reconstructing the Past seeks to clarify and help resolve the vexing methodological issues that arise when biologists try to answer such questions as whether human beings are more closely related to chimps than they are to gorillas. It explores the case for considering the philosophical idea of simplicity/parsimony as a useful principle for evaluating taxonomic theories of evolutionary relationships. For the past two decades, evolutionists have been vigorously debating the appropriate methods that should be used in systematics, the field that (...) aims at reconstructing phylogenetic relationships among species. This debate over phylogenetic inference, Elliott Sober observes, raises broader questions of hypothesis testing and theory evaluation that run head on into long standing issues concerning simplicity/parsimony in the philosophy of science. Sober treats the problem of phylogenetic inference as a detailed case study in which the philosophical idea of simplicity/parsimony can be tested as a principle of theory evaluation. Bringing together philosophy and biology, as well as statistics, Sober builds a general framework for understanding the circumstances in which parsimony makes sense as a tool of phylogenetic inference. Along the way he provides a detailed critique of parsimony in the biological literature, exploring the strengths and limitations of both statistical and nonstatistical cladistic arguments. (shrink)

Stuart Kauffman here presents a brilliant new paradigm for evolutionary biology, one that extends the basic concepts of Darwinian evolution to accommodate recent findings and perspectives from the fields of biology, physics, chemistry and mathematics. The book drives to the heart of the exciting debate on the origins of life and maintenance of order in complex biological systems. It focuses on the concept of self-organization: the spontaneous emergence of order widely observed throughout nature. Kauffman here argues that self-organization plays an (...) important role in the emergence of life itself and may play as fundamental a role in shaping life's subsequent evolution as does the Darwinian process of natural selection. Yet until now no systematic effort has been made to incorporate the concept of self-organization into evolutionary theory. The construction requirements which permit complex systems to adapt remain poorly understood, as is the extent to which selection itself can yield systems able to adapt more successfully. This book explores these themes. It shows how complex systems, contrary to expectations, can spontaneously exhibit stunning degrees of order, and how this order, in turn, is essential for understanding the emergence and development of life on Earth. Topics include the new biotechnology of applied molecular evolution, with its important implications for developing new drugs and vaccines; the balance between order and chaos observed in many naturally occurring systems; new insights concerning the predictive power of statistical mechanics in biology; and other major issues. Indeed, the approaches investigated here may prove to be the new center around which biologicalscience itself will evolve. The work is written for all those interested in the cutting edge of research in the life sciences. (shrink)

"Legend is overdue for replacement, and an adequate replacement must attend to the process of science as carefully as Hull has done. I share his vision of a serious account of the social and intellectual dynamics of science that will avoid both the rosy blur of Legend and the facile charms of relativism.... Because of [Hull's] deep concern with the ways in which research is actually done, Science as a Process begins an important project in the study of science. It (...) is one of a distinguished series of books, which Hull himself edits."—Philip Kitcher, Nature "In Science as a Process, [David Hull] argues that the tension between cooperation and competition is exactly what makes science so successful.... Hull takes an unusual approach to his subject. He applies the rules of evolution in nature to the evolution of science, arguing that the same kinds of forces responsible for shaping the rise and demise of species also act on the development of scientific ideas."—Natalie Angier, New York Times Book Review "By far the most professional and thorough case in favour of an evolutionary philosophy of science ever to have been made. It contains excellent short histories of evolutionary biology and of systematics (the science of classifying living things); an important and original account of modern systematic controversy; a counter-attack against the philosophical critics of evolutionary philosophy; social-psychological evidence, collected by Hull himself, to show that science does have the character demanded by his philosophy; and a philosophical analysis of evolution which is general enough to apply to both biological and historical change."—Mark Ridley, Times Literary Supplement "Hull is primarily interested in how social interactions within the scientific community can help or hinder the process by which new theories and techniques get accepted.... The claim that science is a process for selecting out the best new ideas is not a new one, but Hull tells us exactly how scientists go about it, and he is prepared to accept that at least to some extent, the social activities of the scientists promoting a new idea can affect its chances of being accepted."—Peter J. Bowler, Archives of Natural History "I have been doing philosophy of science now for twenty-five years, and whilst I would never have claimed that I knew everything, I felt that I had a really good handle on the nature of science, Again and again, Hull was able to show me just how incomplete my understanding was.... Moreover, [Science as a Process] is one of the most compulsively readable books that I have ever encountered."—Michael Ruse, Biology and Philosophy. (shrink)

Comprehensive evaluation and study of man's theories and knowledge of genetical characteristics and the evolutionary processes.

In the past century, nearly all of the biological sciences have been directly affected by discoveries and developments in genetics, a fast-evolving subject with important theoretical dimensions. In this rich and accessible book, Paul Griffiths and Karola Stotz show how the concept of the gene has evolved and diversified across the many fields that make up modern biology. By examining the molecular biology of the 'environment', they situate genetics in the developmental biology of whole organisms, and reveal how the molecular (...) biosciences have undermined the nature/nurture distinction. Their discussion gives full weight to the revolutionary impacts of molecular biology, while rejecting 'genocentrism' and 'reductionism', and brings the topic right up to date with the philosophical implications of the most recent developments in genetics. Their book will be invaluable for those studying the philosophy of biology, genetics and other life sciences. (shrink)

Argues for the primacy of the phylogenetic system as the general reference system in biology. This book, first published in 1966, generated significant controversy and opened possibilities for evolutionary biology.

Ideas about heredity and evolution are undergoing a revolutionary change. New findings in molecular biology challenge the gene-centered version of Darwinian theory according to which adaptation occurs only through natural selection of chance DNA variations. In Evolution in Four Dimensions, Eva Jablonka and Marion Lamb argue that there is more to heredity than genes. They trace four "dimensions" in evolution -- four inheritance systems that play a role in evolution: genetic, epigenetic, behavioral, and symbolic. These systems, they argue, can all (...) provide variations on which natural selection can act. Evolution in Four Dimensions offers a richer, more complex view of evolution than the gene-based, one-dimensional view held by many today. The new synthesis advanced by Jablonka and Lamb makes clear that induced and acquired changes also play a role in evolution.After discussing each of the four inheritance systems in detail, Jablonka and Lamb "put Humpty Dumpty together again" by showing how all of these systems interact. They consider how each may have originated and guided evolutionary history and they discuss the social and philosophical implications of the four-dimensional view of evolution. Each chapter ends with a dialogue in which the authors engage the contrarieties of the fictional "I.M.," or Ifcha Mistabra -- Aramaic for "the opposite conjecture" -- refining their arguments against I.M.'s vigorous counterarguments. The lucid and accessible text is accompanied by artist-physician Anna Zeligowski's lively drawings, which humorously and effectively illustrate the authors' points. (shrink)

Phenotypic Evolution explicitly recognizes organisms as complex genetic-epigenetic systems developing in response to changing internal and external environments. As a key to a better understanding of how phenotypes evolve, the authors have developed a framework that centers on the concept of the Developmental Reaction Norm. This encompasses their views: (1) that organisms are better considered as integrated units than as disconnected parts (allometry and phenotypic integration); (2) that an understanding of ontogeny is vital for evaluating evolution of adult forms (ontogenetic (...) trajectories, epigenetics, and constraints); and (3) that environmental heterogeneity is ubiquitous and must be acknowledged for its pervasive role in phenotypic expression. (shrink)

"This is a wonderful book! In "How Scientists Explain Disease," Paul Thagard offers us a delightful essay combining science, its history, philosophy, and sociology.

In the six decades since the publication of Julian Huxley's Evolution: The Modern Synthesis, spectacular empirical advances in the biological sciences have been accompanied by equally significant developments within the core theoretical framework of the discipline. As a result, evolutionary theory today includes concepts and even entire new fields that were not part of the foundational structure of the Modern Synthesis. In this volume, sixteen leading evolutionary biologists and philosophers of science survey the conceptual changes that have emerged since Huxley's (...) landmark publication, not only in such traditional domains of evolutionary biology as quantitative genetics and paleontology but also in such new fields of research as genomics and EvoDevo. Most of the contributors to Evolution—The Extended Synthesis accept many of the tenets of the classical framework but want to relax some of its assumptions and introduce significant conceptual augmentations of the basic Modern Synthesis structure—just as the architects of the Modern Synthesis themselves expanded and modulated previous versions of Darwinism. This continuing revision of a theoretical edifice the foundations of which were laid in the middle of the nineteenth century—the reexamination of old ideas, proposals of new ones, and the synthesis of the most suitable—shows us how science works, and how scientists have painstakingly built a solid set of explanations for what Darwin called the "grandeur" of life. (shrink)

In this path-breaking work, Paul Thagard draws on history and philosophy of science, cognitive psychology, and the field of artificial intelligence to develop a ...

A comprehensive introduction to current research on the psychology of concept formation and use.

_This sweeping discussion of the philosophy of evolutionary biology is based on the revolutionary idea that species are not kinds of organisms but wholes composed of organisms._.

This book presents a critical analysis of the semantic incommensurability thesis of Thomas Kuhn and Paul Feyerabend. In putting forward the thesis of incommensurability, Kuhn and Feyerabend drew attention to complex issues concerning the phenomenon of conceptual change in science. They raised serious problems about the semantic and logical relations between the content of theories which deploy unlike systems of concepts. Yet few of the more extreme claims associated with incommensurability stand scrutiny. The argument of this book is as follows. (...) The terms of theories with different concepts diverge semantically and the vocabularies of such theories may not be fully intertranslatable. Given the possibility of referential overlap between the terms of conceptually divergent theories, it does not follow that the content of such theories is incomparable. Nor, given the distinction between understanding a language and translation from one language into another, does it follow that the proponents of mutually untranslatable theories are unable to communicate. Neither is it the case that the shift between conceptually divergent theories involves a discontinuous transition between theories which have no common reference or that refer to distinct worlds of their own making. The semantic differences resulting from conceptual variance may be embraced within a thoroughly realist framework on which they are construed as diverse linguistic relations to a fixed and independent reality. -/- The book, which was originally published in 1994, has now been re-issued in the Routledge Revivals series. -/- . (shrink)

Only human beings have a rich conceptual repertoire with concepts like tort, entropy, Abelian group, mannerism, icon and deconstruction. How have humans constructed these concepts? And once they have been constructed by adults, how do children acquire them? While primarily focusing on the second question, in The Origin of Concepts , Susan Carey shows that the answers to both overlap substantially. Carey begins by characterizing the innate starting point for conceptual development, namely systems of core cognition. Representations of core cognition (...) are the output of dedicated input analyzers, as with perceptual representations, but these core representations differ from perceptual representations in having more abstract contents and richer functional roles. Carey argues that the key to understanding cognitive development lies in recognizing conceptual discontinuities in which new representational systems emerge that have more expressive power than core cognition and are also incommensurate with core cognition and other earlier representational systems. Finally, Carey fleshes out Quinian bootstrapping, a learning mechanism that has been repeatedly sketched in the literature on the history and philosophy of science. She demonstrates that Quinian bootstrapping is a major mechanism in the construction of new representational resources over the course of childrens cognitive development. Carey shows how developmental cognitive science resolves aspects of long-standing philosophical debates about the existence, nature, content, and format of innate knowledge. She also shows that understanding the processes of conceptual development in children illuminates the historical process by which concepts are constructed, and transforms the way we think about philosophical problems about the nature of concepts and the relations between language and thought. (shrink)

The operation of developing a concept is a common procedure in mathematics and in natural science, but has traditionally seemed much less possible to philosophers and, especially, logicians. Meir Buzaglo's innovative study proposes a way of expanding logic to include the stretching of concepts, while modifying the principles which block this possibility. He offers stimulating discussions of the idea of conceptual expansion as a normative process, and of the relation of conceptual expansion to truth, meaning, reference, ontology and paradox, and (...) analyzes the views of Kant, Wittgenstein, Godel, and others, paying especially close attention to Frege. His book will be of interest to a wide range of readers, from philosophers to logicians, mathematicians, linguists, and cognitive scientists. (shrink)

New concepts are constantly being introduced into our thinking. Conceptual Systems explores how these new concepts are entered into our systems along with sufficient continuity with older ideas to ensure understanding. The encyclopaedic breadth of this text highlights the many different aspects and disciplines that together present an insightful view into the various theories of concepts. Harold Brown, a reputable author in the philosophy of science examines several historically influential theories of concepts as well as providing a clear view on (...) the general theory of conceptual change. Interesting case studies examine examples of conceptual change in the history of physics including the move in seventeenth century physics from Galileo to Descates to Newton; and the conceptual framework of the "standard model" in the late twentieth century high- energy physics. The key central themes in the philosophy of science that are explored in- depth in this enormous book make it an essential read for academics in this field. (shrink)

The present paper analyzes the use and understanding of the homology concept across different biological disciplines. It is argued that in its history, the homology concept underwent a sort of adaptive radiation. Once it migrated from comparative anatomy into new biological fields, the homology concept changed in accordance with the theoretical aims and interests of these disciplines. The paper gives a case study of the theoretical role that homology plays in comparative and evolutionary biology, in molecular biology, and in evolutionary (...) developmental biology. It is shown that the concept or variant of homology preferred by a particular biological field is used to bring about items of biological knowledge that are characteristic for this field. A particular branch of biology uses its homology concept to pursue its specific theoretical goals. (shrink)

How do fitness and natural selection relate to other evolutionary factors like architectural constraint, mode of reproduction, and drift? In one way of thinking, drawn from Newtonian dynamics, fitness is one force driving evolutionary change and added to other factors. In another, drawn from statistical thermodynamics, it is a statistical trend that manifests itself in natural selection histories. It is argued that the first model is incoherent, the second appropriate; a hierarchical realization model is proposed as a basis for a (...) statistical treatment. It emerges that natural selection does not cause evolution; it just is evolution. The theory incorporates relations of statistical correlation, but not the kind of causation found in fundamental physical processes. (shrink)

Comprehending the origin of marine invertebrate larvae remains a key domain of research for evolutionary biologists, including the repeated origin of direct developmental modes in echinoids. In order to address the latter question, we surveyed existing evidence on relationships of homology between the ectoderm territories of two closely related sea urchin species in the genus Heliocidaris that differ in their developmental mode. Additionally, we explored a recently articulated idea about homology called ‘organizational homology’ (Muller 2003. In: Muller GB, Newman SA, (...) editors. Origination of organismal form: beyond the gene in developmental and evolutionary biology. Cambridge, MA: A Bradford Book, The MIT Press. p 51–69. ) in the context of this specific empirical case study. Applying the perspective of organizational homology to our experimental system of congeneric echinoids has led us to a new hypothesis concerning the ectoderm evolution in these species. The extravestibular ectoderm of the direct developer Heliocidaris erythrogramma is a novel developmental territory that arose as a fusion of the oral and aboral ectoderm territories found in indirect developing echinoids such as Heliocidaris tuberculata. This hypothesis instantiates a theoretical principle concerning the origin of developmental modules, ‘integration’, which has been neglected because the opposite theoretical principle, ‘parcellation’, is more readily observable in events such as gene duplication and divergence (Wagner 1996. Am Zool 36:36–43). J. Exp. Zool. (Mol. Dev. Evol.) 306B:18– 34, 2006. (shrink)