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. (...) (Gilbert, Opitz, and Raff 1996, 368). (shrink)

Philosophers of biology have been absorbed by the problem of defining evolutionary fitness since Darwin made it central to biological explanation. The apparent problem is obvious. Define fitness as some biologists implicitly do, in terms of actual survival and reproduction, and the principle of natural selection turns into an empty tautology: those organisms which survive and reproduce in larger numbers, survive and reproduce in larger numbers. Accordingly, many writers have sought to provide a definition for ‘fitness’ which avoid this outcome. (...) In particular the definition of fitness as a probabilistic propensity has been widely favored.1 Others, recognizing that no definition both correct and complete can actually be provided, have accepted the consequence that the leading principle of the theory is a definitional truth and attempted to mitigate the impact of this outcome for the empirical character of the theory.2 Still others have argued that ‘fitness’ is properly viewed as a term undefined in the theory of natural selection (on the model of mass—a term undefined in Newtonian mechanics).3 But few have contemplated the solution to this problem proposed by Mohan Matthen and André Ariew (hereafter, MA), in.. (shrink)

Many scientists and philosophers of science are troubled by the relative isolation of developmental from evolutionary biology. Reconciling the science of development with the science of heredity preoccupied a minority of biologists for much of the twentieth century, but these efforts were not corporately successful. Mainly in the past fifteen years, however, these previously dispersed integrating programmes have been themselves synthesized and so reinvigorated. Two of these more recent synthesizing endeavours are evolutionary developmental biology and developmental systems theory. While the (...) former is a bourgeoning and scientifically well-respected biological discipline, the same cannot be said of DST, which is virtually unknown among biologists. In this review, we provide overviews of DST and EDB, summarize their key tenets, examine how they relate to one another and to the study of epigenetics, and survey the impact that DST and EDB have had on biological theory and practice. (shrink)

Elliott Sober (1987, 1993) and Orzack and Sober (forthcoming) argue that adaptationism is a very general hypothesis that can be tested by testing various particular hypotheses that invoke natural selection to explain the presence of traits in populations of organisms. In this paper, I challenge Sobers claim that adaptationism is an hypothesis and I argue that it is best viewed as a heuristic (or research strategy). Biologists would still have good reasons for employing this research strategy even if it turns (...) out that natural selection is not the most important cause of evolution. (shrink)

In the context of scientists' reflections on genomics, we examine some fundamental issues in the emerging postgenomic discipline of systems biology. Systems biology is best understood as consisting of two streams. One, which we shall call ‘pragmatic systems biology’, emphasises large‐scale molecular interactions; the other, which we shall refer to as ‘systems‐theoretic biology’, emphasises system principles. Both are committed to mathematical modelling, and both lack a clear account of what biological systems are. We discuss the underlying issues in identifying systems (...) and how causality operates at different levels of organisation. We suggest that resolving such basic problems is a key task for successful systems biology, and that philosophers could contribute to its realisation. We conclude with an argument for more sociologically informed collaboration between scientists and philosophers. BioEssays 27:1270–1276, 2005. © 2005 Wiley Periodicals, Inc. (shrink)

The acronym Developmental systems theory (DST) has been introduced into the literature on development in at least three different contexts in recent years – twice for DST, and before that, for Dynamical Systems Theory – and in all cases, to designate a new perspective for understanding development. Subtle but significant differences in argument and aims distinguish these uses, and confound the difficulty of saying just what DST is. My aim in this paper is to disambiguate these different terms – both (...) to call attention to the difference of perspectives, and to carve out a conceptual space for the concrete issues at stake. (shrink)

Developmental systems theory is a general theoretical perspective on development, heredity and evolution. It is intended to facilitate the study of interactions between the many factors that influence development without reviving `dichotomous' debates over nature or nurture, gene or environment, biology or culture. Several recent papers have addressed the relationship between DST and the thriving new discipline of evolutionary developmental biology. The contributions to this literature by evolutionary developmental biologists contain three important misunderstandings of DST.

The themes, problems and challenges of developmental systems theory as described in Cycles of Contingency are discussed. We argue in favor of a robust approach to philosophical and scientific problems of extended heredity and the integration of behavior, development, inheritance, and evolution. Problems with Sterelny's proposal to evaluate inheritance systems using his `Hoyle criteria' are discussed and critically evaluated. Additional support for a developmental systems perspective is sought in evolutionary studies of performance and behavior modulation of fitness.

A framework is presented in which the role ofdevelopmental rules in phenotypic evolution canbe studied for some simple situations. Usingtwo different implicit models of development,characterized by different developmental mapsfrom genotypes to phenotypes, it is shown bysimulation that developmental rules and driftcan result in directional phenotypic evolutionwithout selection. For both models thesimulations show that the critical parameterthat drives the final phenotypic distributionis the cardinality of the set of genotypes thatmap to each phenotype. Details of thedevelopmental map do not matter. If phenotypesare (...) randomly assigned to genotypes, the lastresult can also be proved analytically. (shrink)

Developmental systems theory (DST) is a general theoretical perspective on development, heredity and evolution. It is intended to facilitate the study of interactions between the many factors that influence development without reviving `dichotomous' debates over nature or nurture, gene or environment, biology or culture. Several recent papers have addressed the relationship between DST and the thriving new discipline of evolutionary developmental biology (EDB). The contributions to this literature by evolutionary developmental biologists contain three important misunderstandings of DST.

We attempt a conclusive resolution of the debate over whether the principle of natural selection (PNS), especially conceived as the `principle' of the `survival of the fittest', is a tautology. This debate has been largely ignored for the past 15 years but not, we think, because it has actually been settled. We begin by describing the tautology objection, and situating the problem in the philosophical and biology literature. We then demonstrate the inadequacy of six prima facie plausible reasons for believing (...) that the tautology debate has been satisfactorily resolved (the PNS is strictly a methodological principle; scientific theories can contain tautologies; the scope of the PNS has been reduced; theories should be understood as models and not exceptionless laws; the widespread acceptance of the propensity interpretation of fitness; and the abandonment of operationalism and verificationism). We proceed to a detailed discussion of Brandon's law (D) describing the PNS, and show that law (D) seriously misrepresents the structure of evolution by natural selection. In the final sections, we provide and defend a novel reinterpretation of the structure of the principle (or, we prefer, model) of evolution by natural selection. (shrink)

We argue that a fashionable interpretation of the theory of natural selection as a claim exclusively about populations is mistaken. The interpretation rests on adopting an analysis of fitness as a probabilistic propensity which cannot be substantiated, draws parallels with thermodynamics which are without foundations, and fails to do justice to the fundamental distinction between drift and selection. This distinction requires a notion of fitness as a pairwise comparison between individuals taken two at a time, and so vitiates the interpretation (...) of the theory as one about populations exclusively. (shrink)

We will show that there is a strong form of emergence in cell biology. Beginning with C.D. Broad’s classic discussion of emergence, we distinguish two conditions sufficient for emergence. Emergence in biology must be compatible with the thought that all explanations of systemic properties are mechanistic explanations and with their sufficiency. Explanations of systemic properties are always in terms of the properties of the parts within the system. Nonetheless, systemic properties can still be emergent. If the properties of the components (...) within the system cannot be predicted, even in principle, from the behavior of the system’s parts within simpler wholes then there also will be systemic properties which cannot be predicted, even in principle, on basis of the behavior of these parts. We show in an explicit case study drawn from molecular cell physiology that biochemical networks display this kind of emergence, even though they deploy only mechanistic explanations. This illustrates emergence and its place in nature. (shrink)

We will show that there is a strong form of emergence in cell biology. Beginning with C.D. Broad's classic discussion of emergence, we distinguish two conditions sufficient for emergence. Emergence in biology must be compatible with the thought that all explanations of systemic properties are mechanistic explanations and with their sufficiency. Explanations of systemic properties are always in terms of the properties of the parts within the system. Nonetheless, systemic properties can still be emergent. If the properties of the components (...) within the system cannot be predicted, even in principle, from the behavior of the system's parts within simpler wholes then there also will be systemic properties which cannot be predicted, even in principle, on basis of the behavior of these parts. We show in an explicit case study drawn from molecular cell physiology that biochemical networks display this kind of emergence, even though they deploy only mechanistic explanations. This illustrates emergence and its place in nature. (shrink)

The central point of this essay is to demonstrate the incommensurability of ‘Darwinian fitness’ with the numeric values associated with reproductive rates used in population genetics. While sometimes both are called ‘fitness’, they are distinct concepts coming from distinct explanatory schemes. Further, we try to outline a possible answer to the following question: from the natural properties of organisms and a knowledge of their environment, can we construct an algorithm for a particular kind of organismic life-history pattern that itself will (...) allow us to predict whether a type in the population will increase or decrease relative to other types? Introduction Darwinian fitness Reproductive fitness and genetical models of evolution The models of reproductive fitness 4.1 The Standard Viability Model 4.2 Frequency-dependent selection 4.3 Fertility models 4.4 Overlapping generations Fitness as outcome 5.1 Fitness as actual increase in type 5.2 Fitness as expected increase in type 5.2.1 Expected increase within a generation 5.2.2 Expected increase between generations 5.2.3 Postponed reproductive fitness effects The book-keeping problem Conclusion. (shrink)

The central point of this essay is to demonstrate the incommensurability of ‘Darwinian fitness’ with the numeric values associated with reproductive rates used in population genetics. While sometimes both are called ‘fitness’, they are distinct concepts coming from distinct explanatory schemes. Further, we try to outline a possible answer to the following question: from the natural properties of organisms and a knowledge of their environment, can we construct an algorithm for a particular kind of organismic life-history pattern that itself will (...) allow us to predict whether a type in the population will increase or decrease relative to other types? 1. Introduction2. Darwinian fitness3. Reproductive fitness and genetical models of evolution4. The models of reproductive fitness4.1 The Standard Viability Model4.2 Frequency-dependent selection4.3 Fertility models4.4 Overlapping generations5. Fitness as outcome5.1 Fitness as actual increase in type5.2 Fitness as expected increase in type5.2.1 Expected increase within a generation5.2.2 Expected increase between generations5.2.3 Postponed reproductive fitness effects6. The book-keeping problem7. Conclusion. (shrink)

In “Two Ways of Thinking About Fitness and Natural Selection” (Matthen and Ariew [2002]; henceforth “Two Ways”), we asked how one should think of the relationship between the various factors invoked to explain evolutionary change – selection, drift, genetic constraints, and so on. We suggested that these factors are not related to one another as “forces” are in classical mechanics. We think it incoherent, for instance, to think of natural selection and drift as separate and opposed “forces” in evolutionary change (...) – that it makes sense to say, for instance, that selection contributed 80% to the actual evolutionary history of the human eye, and drift only 20%. We proposed instead a statistical view of the Theory of Evolution, a view in which fitness is not a cause of evolution, but rather a measure of growth. We also argued for a “hierarchical realization model” for thinking about the relationship between evolutionary factors such as those mentioned above, and suggested that in a “fully specified model”, as we call it below, there is no distinction between natural selection and evolution. (shrink)

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)

The debate over the relative importance of natural selection as compared to other forces affecting the evolution of organisms is a long-standing and central controversy in evolutionary biology. The theory of adaptationism argues that natural selection contains sufficient explanatory power in itself to account for all evolution. However, there are differing views about the efficiency of the adaptation model of explanation. If the adaptationism theory is applied, are energy and resources being used to their optimum? This book presents an up-to-date (...) view of this controversy and reflects the dramatic changes in our understanding of evolution that have occurred in the last twenty years. The volume combines contributions from biologists and philosophers, and offers a systematic treatment of foundational, conceptual, and methodological issues surrounding the theory of adaptationism. The essays examine recent developments in topics such as phylogenetic analysis, the theory of optimality and ess models, and methods of testing models. (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)

Historically, philosophers of biology have tended to sidestep the problem of development by focusing primarily on evolutionary biology and, more recently, on molecular biology and genetics. Quite often too, development has been misunderstood as simply, or even primarily, a matter of gene activation and regulation. Nowadays a growing number of philosophers of science are focusing their analyses on the complexities of development, and in Embryology, Epigenesis and Evolution Jason Scott Robert explores the nature of development against current trends in biological (...) theory and practice and looks at the interrelations between development and evolution, an area of resurgent biological interest. Clearly written, this book should be of interest to students and professionals in the philosophy of science and the philosophy of biology. (shrink)

With the advent of the Human Genome Project there have been many claims for the genetic origins of complex human behavior including insanity, criminality, and intelligence. But what does it really mean to call something 'genetic'? This is the fundamental question that Sahotra Sarkar's book addresses. The author analyses the nature of reductionism in classical and molecular genetics. He shows that there are two radically different kinds of reductionist explanation: genetic reduction (as found in classical genetics) and physical reduction (found (...) in molecular genetics). This important book clarifies the meaning of the term 'genetic', shows how molecular studies have affected genetics, and provides the philosophical background necessary to understand the debates over the Human Genome Project. It will be of particular interest to professionals and students in the philosophy of science, the history of science, and the social studies of science, medicine, and technology. (shrink)

This collection broadens the scientific discussion of modularity by bringing together experts from a variety of disciplines, including artificial life, ...

Is life different from the non-living? If so, how? And how, in that case, does biology as the study of living things differ from other sciences? These questions are traced through an exploration of episodes in the history of biology and philosophy. The book begins with Aristotle, then moves on to Descartes, comparing his position with that of Harvey. In the eighteenth century the authors consider Buffon and Kant. In the nineteenth century the authors examine the Cuvier-Geoffroy debate, pre-Darwinian geology (...) and natural theology, Darwin and the transition from Darwin to the revival of Mendelism. Two chapters deal with the evolutionary synthesis and such questions as the species problem, the reducibility or otherwise of biology to physics and chemistry, and the problem of biological explanation in terms of function and teleology. The final chapters reflect on the implications of the philosophy of biology for philosophy of science in general. (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)].

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)

Debate about adaptationism in biology continues, in part because within “the” problem of assessing adaptationism, three distinct problems are mixed together. The three problems concern the assessment of three distinct adaptationist positions, each of which asserts the central importance of adaptation and natural selection to the study of evolution, but conceives this importance in a different way. As there are three kinds of adaptationism, there are three distinct "anti-adaptationist" positions as well. Or putting it more formally, there are three different (...) dimensions here, and strongly adaptationist views, strongly anti-adaptationist views, and moderate views are possible for each dimension. (shrink)

The concept of fitness began its career in biology long before evolutionary theory was mathematized. Fitness was used to describe an organism’s vigor, or the degree to which organisms “fit” into their environments. An organism’s success in avoiding predators and in building a nest obviously contribute to its fitness and to the fitness of its offspring, but the peacock’s gaudy tail seemed to be in an entirely different line of work. Fitness, as a term in ordinary language (as in “physical (...) fitness”) and in its original biological meaning, applied to the survival of an organism and its offspring, not to sheer reproductive output (Paul ////; Cronin 1991). Darwin’s separation of natural from sexual selection may sound odd from a modern perspective, but it made sense from this earlier point of view. (shrink)

Darwin’s explanation of biological speciation in terms of variation and natural selection has revolutionised biological thought. However, while his principle of natural selection, the fitness principle, has shaped biology until the present, its interpretation changed more than once during the almost 150 years of its history. The most striking change of the status of the principle is that, in the middle of the 20th century, it transmutated from an often disputed, groundbreaking insight into a tautology. Moreover, not only the interpretation (...) of the fitness principle, but the whole body of biological knowledge was subjected to significant modifications. In this paper, I relate modifications of the fitness principle to those of the respective body of biological knowledge. This body of knowledge is conceived as a Quinean web of belief. After an exposition of Darwin’s conception of the principle, which equated fitness with adaptedness to the environment, several of its changes are analysed with respect to different webs of biological knowledge. It is concluded that the different interpretations and the reshaping of the fitness principle are rational responses to the modified systems of background knowledge, which saved the coherence of the web of biological knowledge in each single case. (shrink)