In The Selfish Gene, Richard Dawkins crystallized the gene's eye view of evolution developed by W.D. Hamilton and others. The book provoked widespread and heated debate. Written in part as a response, The Extended Phenotype gave a deeper clarification of the central concept of the gene as the unit of selection; but it did much more besides. In it, Dawkins extended the gene's eye view to argue that the genes that sit within an organism have an influence that reaches out (...) beyond the visible traits in that body - the phenotype - to the wider environment, which can include other individuals. So, for instance, the genes of the beaver drive it to gather twigs to produce the substantial physical structure of a dam; and the genes of the cuckoo chick produce effects that manipulate the behaviour of the host bird, making it nurture the intruder as one of its own. This notion of the extended phenotype has proved to be highly influential in the way we understand evolution and the natural world. It represents a key scientific contribution to evolutionary biology, and it continues to play an important role in research in the life sciences. The Extended Phenotype is a conceptually deep book that forms important reading for biologists and students. But Dawkins' clear exposition is accessible to all who are prepared to put in a little effort. Oxford Landmark Science books are 'must-read' classics of modern science writing which have crystallized big ideas, and shaped the way we think. (shrink)

Since the introduction of mathematical population genetics, its machinery has shaped our fundamental understanding of natural selection. Selection is taken to occur when differential fitnesses produce differential rates of reproductive success, where fitnesses are understood as parameters in a population genetics model. To understand selection is to understand what these parameter values measure and how differences in them lead to frequency changes. I argue that this traditional view is mistaken. The descriptions of natural selection rendered by population genetics models are (...) in general neither predictive nor explanatory and introduce avoidable conceptual confusions. I conclude that a correct understanding of natural selection requires explicitly causal models of reproductive success. *Received May 2006; revised December 2006. †To contact the author, please write to: Department of Philosophy, Kansas State University, 201 Dickens Hall, Manhattan, KS 66506; e‐mail: [email protected] . (shrink)

Evolutionary biology is a field currently animated by much discussion concerning its conceptual foundations. On the one hand, we have supporters of a classical view of evolutionary theory, whose backbone is provided by population genetics and the so-called Modern Synthesis (MS). On the other hand, a number of researchers are calling for an Extended Synthe- sis (ES) that takes seriously both the limitations of the MS (such as its inability to incorporate developmental biology) and recent empirical and theoretical research on (...) issues such as evolvability, modularity, and self-organization. In this article, I engage in an in-depth commentary of an influential paper by population geneticist Michael Lynch, which I take to be the best defense of the MS-population genetics position published so far. I show why I think that Lynch’s arguments are wanting and propose a modification of evolutionary theory that retains but greatly expands on population genetics. (shrink)

There are two competing interpretations of the modern synthesis theory of evolution: the dynamical (also know as ‘traditional’) and the statistical. The dynamical interpretation maintains that explanations offered under the auspices of the modern synthesis theory articulate the causes of evolution. It interprets selection and drift as causes of population change. The statistical interpretation holds that modern synthesis explanations merely cite the statistical structure of populations. This paper offers a defense of statisticalism. It argues that a change in trait frequencies (...) in a population can be attributed only to selection or drift against the background of a particular statistical description of the population. The traditionalist supposition that selection and drift are description‐independent causes of population change leads the dynamical interpretation into a dilemma: it must face a contradiction or accept the loss of explanatory power. (shrink)

The ontological dependence of one domain on another is compatible with the explanatory autonomy of the less basic domain. That autonomy results from the fact that the relationship between two domains can be very complex. In this paper I distinguish two different types of complexity, two ways the relationship between domains can fail to be transparent, both of which are relevant to evolutionary biology. Sometimes high level explanations preserve a certain type of causal or counterfactual information which would be lost (...) at the lower level; I argue that this is central to the proper understanding of the adaptationist program. Sometimes high level kinds are multiply realised by lower level kinds: I argue that this is central to the understanding of macroevolution. (shrink)

It is nowadays a dominant opinion in a number of disciplines (anthropology, genetics, psychology, philosophy of science) that the taxonomy of human races does not make much biological sense. My aim is to challenge the arguments that are usually thought to invalidate the biological concept of race. I will try to show that the way “race” was defined by biologists several decades ago (by Dobzhansky and others) is in no way discredited by conceptual criticisms that are now fashionable and widely (...) regarded as cogent. These criticisms often arbitrarily burden the biological category of race with some implausible connotations, which then opens the path for a quick eliminative move. However, when properly understood, the biological notion of race proves remarkably resistant to these deconstructive attempts. Moreover, by analyzing statements of some leading contemporary scholars who support social constructivism about race, I hope to demonstrate that their eliminativist views are actually in conflict with what the best contemporary science tells us about human genetic variation. (shrink)

Evolutionary processes such as natural selection and random drift are commonly regarded as causes of population-level change. We respond to a recent challenge that drift and selection are best understood as statistical trends, not causes. Our reply appeals to manipulation as a strategy for uncovering causal relationships: if you can systematically manipulate variable A to bring about a change in variable B, then A is a cause of B. We argue that selection and drift can be systematically manipulated to produce (...) different kinds of population-level change. They should therefore be regarded as causes. (shrink)

In 1966, Richard Levins argued that there are different strategies in model building in population biology. In this paper, I reply to Orzack and Sober’s (1993) critiques of Levins, and argue that his views on modeling strategies apply also in the context of evolutionary genetics. In particular, I argue that there are different ways in which models are used to ask and answer questions about the dynamics of evolutionary change, prospectively and retrospectively, in classical versus molecular evolutionary genetics. Further, I (...) argue that robustness analysis is a tool for, if not confirmation, then something near enough, in this discipline. (shrink)

The causal nature of evolution is one of the central topics in the philosophy of biology. The issue concerns whether equations used in evolutionary genetics point to some causal processes or purely phenomenological patterns. To address this question the present article builds well-defined causal models that underlie standard equations in evolutionary genetics. These models are based on minimal and biologically plausible hypotheses about selection and reproduction, and generate statistics to predict evolutionary changes. The causal reconstruction of the evolutionary principles shows (...) adaptive evolution as a genuine causal process, where fitness and selection are both causes of evolution. 1 Introduction2 The Philosophical Puzzle3 Causal Models4 Causal Foundations of Evolutionary Genetics4.1 Univariate quantitative genetics model4.1.1 The causal graph4.1.2 Structural equations4.1.3 Deriving evolutionary transition functions4.2 One-locus population genetics model5 Evolution as a Causal Process5.1 Selection as a causal process5.2 Causes of evolutionary change6 Conclusion. (shrink)

Over the past decade philosophers of biology have discussed whether evolutionary theory is a causal theory or a phenomenological study of evolution based solely on the statistical features of a population. This article reviews this controversy from three aspects, respectively concerning the assumptions, applications, and explanations of evolutionary theory, with a view to arriving at a definite conclusion in each contention. In so doing I also argue that an implicit methodological assumption shared by both sides of the debate, namely the (...) overconfidence in conceptual analysis as a tool to understand the scientific theory, is the real culprit that has both generated the problem and precluded its solution for such a long time. (shrink)

Ernst Mayr has criticised the methodology of population genetics for being essentialist: interested only in “types” as opposed to individuals. In fact, he goes so far as to claim that “he who does not understand the uniqueness of individuals is unable to understand the working of natural selection” (1982, 47). This is a strong claim indeed especially since many responsible for the development of population genetics (especially Fisher, Haldane, and Wright) were avid Darwinians. In order to unravel this apparent incompatibility (...) I want to examine the possible sources and implications of essentialism in this context and show why the kind of mathematical analysis found in Fisher's work is better seen as responsible for extending the theory of natural selection to a broader context rather than inhibiting its applicability. (shrink)

The latter half of the twentieth century has been marked by debates in evolutionary biology over the relative significance of natural selection and random drift: the so-called “neutralist/selectionist” debates. Yet John Beatty has argued that it is difficult, if not impossible, to distinguish the concept of random drift from the concept of natural selection, a claim that has been accepted by many philosophers of biology. If this claim is correct, then the neutralist/selectionist debates seem at best futile, and at worst, (...) meaningless. I reexamine the issues that Beatty raises, and argue that random drift and natural selection, conceived as processes, can be distinguished from one another. (shrink)

We have argued elsewhere that: (A) Natural selection is not a cause of evolution. (B) A resolution-of-forces (or vector addition) model does not provide us with a proper understanding of how natural selection combines with other evolutionary influences. These propositions have come in for criticism recently, and here we clarify and defend them. We do so within the broad framework of our own “hierarchical realization model” of how evolutionary influences combine.

Although there once was a general consensus among race scholars that applying race categories to humans is biologically illegitimate, this consensus has been erased over the past decade. This is largely due to advances in population genetics that allow biologists to pick out genetic population clusters that approximate some of our common sense racial categories. In this paper, I argue that this new ability really ought not undermine our confidence in the biological illegitimacy of the human races. Unfortunately, the claim (...) that races are biologically legitimate is ambiguous—the sense I will be concerned with here holds that the human species can be divided into biological subspecies. Sesardic and Andreasen have both argued that the cluster results are evidence for subspecies within the human population. I show the cluster results are in fact evidentially inert relative to each author’s preferred subspecies concept. I then sketch the kinds of biological facts that could be used to adjudicate whether human subspecies have existed in the past. (shrink)

In a small handful of papers in theoretical population genetics, John Gillespie (2000a, 2000b, 2001) argues that a new stochastic process he calls "genetic draft" is evolutionarily more significant than genetic drift. This case study of chance in evolution explores Gillespie's proposed stochastic evolutionary force and sketches the implications of Gillespie's argument for philosophers' explorations of genetic drift.

We outline three very different concepts of the gene—instrumental, nominal, and postgenomic. The instrumental gene has a critical role in the construction and interpretation of experiments in which the relationship between genotype and phenotype is explored via hybridization between organisms or directly between nucleic acid molecules. It also plays an important theoretical role in the foundations of disciplines such as quantitative genetics and population genetics. The nominal gene is a critical practical tool, allowing stable communication between bioscientists in a wide (...) range of fields grounded in well-defined sequences of nucleotides, but this concept does not embody major theoretical insights into genome structure or function. The post-genomic gene embodies the continuing project of understanding how genome structure supports genome function, but with a deflationary picture of the gene as a structural unit. This final concept of the gene poses a significant challenge to conventional assumptions about the relationship between genome structure and function, and between genotype and phenotype. (shrink)

The book examines issues related to the way modeling and simulation enable us to reconstruct aspects of the world we are investigating. It also investigates the processes by which we extract concrete knowledge from those reconstructions and how that knowledge is legitimated.

In this book Ron Amundson examines 200 years of scientific views on the evolution-development relationship from the perspective of evolutionary developmental biology (evo-devo). This new perspective challenges several popular views about the history of evolutionary thought by claiming that many earlier authors 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 to evolution with the Evolutionary Synthesis. It concludes with an examination of (...) the contrasts that persist between mainstream evolutionary theory and evo-devo. (shrink)

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)

Advances in molecular biological research in the latter half of the twentieth century have made the story of the gene vastly complicated: the more we learn about genes, the less sure we are of what a gene really is. Knowledge about the structure and functioning of genes abounds, but the gene has also become curiously intangible. This collection of essays renews the question: what are genes? Philosophers, historians and working scientists re-evaluate the question in this volume, treating the gene as (...) a focal point of interdisciplinary and international research. It will be of interest to professionals and students in the philosophy and history of science, genetics and molecular biology. (shrink)