Debates over adaptationism can be clarified and partially resolved by careful consideration of the ‘grain’ at which evolutionary processes are described. The framework of ‘adaptive landscapes’ can be used to illustrate and facilitate this investigation. We argue that natural selection may have special status at an intermediate grain of analysis of evolutionary processes. The cases of sickle-cell disease and genomic imprinting are used as case studies.

A phylogeny that allows for lateral gene transfer (LGT) can be thought of as a strictly branching tree (all of whose branches are vertical) to which lateral branches have been added. Given that the goal of phylogenetics is to depict evolutionary history, we should look for the best supported phylogenetic network and not restrict ourselves to considering trees. However, the obvious extensions of popular tree-based methods such as maximum parsimony and maximum likelihood face a serious problem—if we judge networks by (...) fit to data alone, networks that have lateral branches will always fit the data at least as well as any network that restricts itself to vertical branches. This is analogous to the well-studied problem of overfitting data in the curve-fitting problem. Analogous problems often have analogous solutions and we propose to treat network inference as a case of model selection and use the Akaike Information Criterion (AIC). Strictly tree-like networks are more parsimonious than those that postulate lateral as well as vertical branches. This leads to the conclusion that we should not always infer LGT events whenever it would improve our fit-to-data, but should do so only when the improved fit is larger than the penalty for adding extra lateral branches. (shrink)

Markov models of evolution describe changes in the probability distribution of the trait values a population might exhibit. In consequence, they also describe how entropy and conditional entropy values evolve, and how the mutual information that characterizes the relation between an earlier and a later moment in a lineage’s history depends on how much time separates them. These models therefore provide an interesting perspective on questions that usually are considered in the foundations of physics—when and why does entropy increase and (...) at what rates do changes in entropy take place? They also throw light on an important epistemological question: are there limits on what your observations of the present can tell you about the evolutionary past? (shrink)

Sewall Wright's adaptive landscape is the most influential heuristic in evolutionary biology. Wright's biographer, Provine, criticized Wright's adaptive landscape, claiming that its heuristic value is dubious because of deep flaws. Ruse has defended Wright against Provine. Ruse claims Provine has not shown Wright's use of the landscape is flawed, and that, even if it were, it is heuristically valuable. I argue that both Provine's and Ruse's analyses of the adaptive landscape are defective and suggest a more adequate understanding of it.

Stephen Jay Gould argued that replaying the “tape of life” would result in a radically different evolutionary outcome. Some biologists and philosophers, however, have pointed to convergent evolution as evidence for robust replicability in macroevolution. These authors interpret homoplasy, or the independent origination of similar biological forms, as evidence for the power of natural selection to guide form toward certain morphological attractors, notwithstanding the diversionary tendencies of drift and the constraints of phylogenetic inertia. In this paper, I consider the implications (...) of homoplasy for the debate over the nature of macroevolution. I argue that once the concepts of contingency and convergence are fleshed out, it becomes clear that many instances of homoplasy fail to negate Gould’s overarching thesis, and may in fact support a Gouldian view of life. My argument rests on the distinction between parallelism and convergence, which I defend against a recent challenge from developmental biology. I conclude that despite the difficulties in defining and identifying parallelism, the concept remains useful and relevant to the contingency controversy insofar as it underscores the common developmental origins of iterated evolution. (shrink)

The discussion of the adaptive landscape in the philosophical literature appears to be divided along the following lines. On the one hand, some claim that the adaptive landscape is either “uninterpretable” or incoherent. On the other hand, some argue that the adaptive landscape has been an important heuristic, or tool in the service of explaining, as well as proposing and testing hypotheses about evolutionary change. This paper attempts to reconcile these two views.

A hitherto neglected form of explanation is explored, especially its role in population genetics. “Statistically abstractive explanation” (SA explanation) mandates the suppression of factors probabilistically relevant to an explanandum when these factors are extraneous to the theoretical project being pursued. When these factors are suppressed, the explanandum is rendered uncertain. But this uncertainty traces to the theoretically constrained character of SA explanation, not to any real indeterminacy. Random genetic drift is an artifact of such uncertainty, and it is therefore wrong (...) to reify it as a cause of evolution or as a process in its own right. *Received July 2009. †To contact the author, please write to: Department of Philosophy, University of Toronto, 170 St. George St., Toronto, ON M5R 2M8, Canada; e‐mail: [email protected]. (shrink)

The concepts of adaptive/fitness landscapes and adaptive peaks are a central part of much of contemporary evolutionary biology; the concepts are introduced in introductory texts, developed in more detail in graduate-level treatments, and are used extensively in papers published in the major journals in the field. The appeal of visualizing the process of evolution in terms of the movement of populations on such landscapes is very strong; as one becomes familiar with the metaphor, one often develops the feeling that it (...) is possible to gain deep insights into evolution by thinking about the movement of populations on landscapes consisting of adaptive valleys and peaks. But, since Wright first introduced the metaphor in 1932, the metaphor has been the subject of persistent confusion, from equivocation over just what the features of the landscape are meant to represent to how we ought to expect the landscapes to look. Recent advances—conceptual, empirical, and computational—have pointed towards the inadequacy and indeed incoherence of the landscapes as usually pictured. I argue that attempts to reform the metaphor are misguided; it is time to give up the pictorial metaphor of the landscape entirely and rely instead on the results of formal modeling, however difficult such results are to understand in ‘intuitive’ terms. (shrink)

This paper explores an important type of biological explanation called ‘homology thinking.’ Homology thinking explains the properties of a homologue by citing the history of a homologue. Homology thinking is significant in several ways. First, it offers more detailed explanations of biological phenomena than corresponding analogy explanations. Second, it provides an important explanation of character similarity and difference. Third, homology thinking offers a promising account of multiple realizability in biology.

In 1894 Pierre Curie announced what has come to be known as Curie's Principle: the asymmetry of effects must be found in their causes. In the same publication Curie discussed a key feature of what later came to be known as spontaneous symmetry breaking: the phenomena generally do not exhibit the symmetries of the laws that govern them. Philosophers have long been interested in the meaning and status of Curie's Principle. Only comparatively recently have they begun to delve into the (...) mysteries of spontaneous symmetry breaking. The present paper aims to advance the discussion of both of these twin topics by tracing their interaction in classical physics, ordinary quantum mechanics and quantum field theory. The features of spontaneous symmetry that are peculiar to quantum field theory have received scant attention in the philosophical literature. These features are highlighted here, along with an explanation of why Curie's Principle, though valid in quantum field theory, is nearly vacuous in that context. (shrink)

This essay examines the claim “path dependence entails irreversibility” from the point of view of evolutionary biology. I argue that evolutionary irreversibility possesses many faces, sometimes conflicting with path dependence. I propose an account of path dependence that does not rely on irreversibility and explains why it more naturally coexists with the notion of (contingent) irreversibility developed by the Belgian paleontologist Louis Dollo. However, I argue that we should not conceive of this relationship as necessary.

Biologists in the last 50 years have increasingly emphasized the role of historical contingency in explaining the distribution and dynamics of biological systems. However, recent work in philosophy of biology has shown that historical contingency carries various interpretations and that we are still lacking a general understanding of historicity, i.e., a framework from which to interpret why and to what extent history matters in biological processes. Building from examples and analyses of the long-term experimental evolution (LTEE) project, this paper argues (...) that historicity possess three essential conditions: (1) multiple possible pasts, (2) multiple possible outcomes at a given instant, and (3) a relationship of causal dependence between these two sets. These criteria can be further specified in two general forms of historicity: dependence on initial conditions and path dependence. More attention is devoted to developing a rigorous account of the latter, which captures the type of historicity displayed by stochastic processes. This paper also highlights that it is often more productive to adopt an instant-relative approach and think in terms of degree of historicity instead of trying to maintain a rigid and absolute dichotomy between historical and ahistorical (completely convergent) processes. (shrink)

This paper analyzes the relevance and interconnection of two forms of historicity in ecological restoration, namely historical fidelity and path dependence. Historical fidelity is the practice of attempting to restore an ecological system to some sort of idealized past condition. Path dependence occurs when a system can evolve in alternative local equilibria, and that the order and timing of the events that follow from the initial state influence which equilibrium is reached. Using theoretical examples and case studies, the following analysis (...) shows that path dependence can seriously impinge the feasibility of historical fidelity, thus reinforcing the idea that restoration ecology needs to move away from a rigid reliance on historical fidelity and is sometimes justified to endorse an interventionist agenda. Yet, contra recent criticisms, I argue that ecological restoration, directed by prudential and evidence-based reasoning, should maintain historical fidelity as one of its guiding objectives. (shrink)

In recent years, the concept of evolvability has been gaining in prominence both within evolutionary developmental biology (evo-devo) and the broader field of evolutionary biology. Despite this, there remains considerable disagreement about what evolvability is. This article offers a solution to this problem. I argue that, in focusing too closely on the role played by evolvability as an explanandum in evo-devo, existing philosophical attempts to clarify the evolvability concept have been overly narrow. Within evolutionary biology more broadly, evolvability offers a (...) robust explanation for the evolutionary trajectories of populations. Evolvability is an abstract, robust, dispositional property of populations, which captures the joint causal influence of their internal features on the outcomes of evolution (as opposed to the causal influence of selection, which is often characterized as external). When considering the nature of the physical basis of this disposition, it becomes clear that the many existing definitions of evolvability at play within evo-devo should be understood as capturing only aspects of a much broader phenomenon. 1 Introduction2 The Problem of Evolvability3 The Theoretical Role of Evolvability in Evolutionary Biology3.1 The explanatory targets of evolutionary biology3.2 Selection-based explanations3.3 Lineage explanations3.4 Evolvability-based explanations3.5 What properties must evolvability have?4 What Evolvability Really Is4.1 Making sense of ft4.2 Making sense of x and b5 What of the Limbs? The Power of E6 Conclusion. (shrink)

In recent years, the concept of evolvability has been gaining in prominence both within evolutionary developmental biology (evo-devo) and the broader field of evolutionary biology. Despite this, there remains considerable disagreement about what evolvability is. This article offers a solution to this problem. I argue that, in focusing too closely on the role played by evolvability as an explanandum in evo-devo, existing philosophical attempts to clarify the evolvability concept have been overly narrow. Within evolutionary biology more broadly, evolvability offers a (...) robust explanation for the evolutionary trajectories of populations. Evolvability is an abstract, robust, dispositional property of populations, which captures the joint causal influence of their internal features on the outcomes of evolution (as opposed to the causal influence of selection, which is often characterized as external). When considering the nature of the physical basis of this disposition, it becomes clear that the many existing definitions of evolvability at play within evo-devo should be understood as capturing only aspects of a much broader phenomenon. 1 Introduction2 The Problem of Evolvability3 The Theoretical Role of Evolvability in Evolutionary Biology 3.1 The explanatory targets of evolutionary biology 3.2 Selection-based explanations 3.3 Lineage explanations 3.4 Evolvability-based explanations 3.5 What properties must evolvability have?4 What Evolvability Really Is 4.1 Making sense of ft 4.2 Making sense of x and b5 What of the Limbs? The Power of E6 Conclusion. (shrink)

The paper provides a new characterization of the concepts of necessity and contingency as they should be used in the historical context. The idea is that contingency (necessity) increases in direct (reverse) proportion to sensitivity to initial conditions. The merits of this suggestion are that it avoids the conflation of causality and necessity (or contingency and chance), that it enables the bracketing of the problem of free will while maintaining the concept of human action making a difference, that it sanctions (...) tendencies without recourse to teleology, and that it recasts the controversy between historicists and anti‐historicists in less dogmatic language. (shrink)

In “Spandrels,” Gould and Lewontin criticized what they took to be an all-too-common conviction, namely, that adaptation to current environments determines organic form. They stressed instead the importance of history. In this paper, we elaborate upon their concerns by appealing to other writings in which those issues are treated in greater detail. Gould and Lewontin’s combined emphasis on history was three-fold. First, evolution by natural selection does not start from scratch, but always refashions preexisting forms. Second, preexisting forms are refashioned (...) by the selection of whatever mutational variations happen to arise: the historical order of mutations needs to be taken into account. Third, the order of environments and selection pressures also needs to be taken into account. (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)

Causality offers the first comprehensive coverage of causal analysis in many sciences, including recent advances using graphical methods. Pearl presents a unified account of the probabilistic, manipulative, counterfactual and structural approaches to causation, and devises simple mathematical tools for analyzing the relationships between causal connections, statistical associations, actions and observations. The book will open the way for including causal analysis in the standard curriculum of statistics, artificial intelligence, business, epidemiology, social science and economics.

Making Sense of Evolution explores contemporary evolutionary biology, focusing on the elements of theories—selection, adaptation, and species—that are complex and open to multiple possible interpretations, many of which are incompatible with one another and with other accepted practices in the discipline. Particular experimental methods, for example, may demand one understanding of “selection,” while the application of the same concept to another area of evolutionary biology could necessitate a very different definition.

Highlighting main issues and controversies, this book brings together current philosophical discussions of symmetry in physics to provide an introduction to the subject for physicists and philosophers. The contributors cover all the fundamental symmetries of modern physics, such as CPT and permutation symmetry, as well as discussing symmetry-breaking and general interpretational issues. Classic texts are followed by new review articles and shorter commentaries for each topic. Suitable for courses on the foundations of physics, philosophy of physics and philosophy of science, (...) the volume is a valuable reference for students and researchers. (shrink)

Symmetry, intended as invariance with respect to a transformation (more precisely, with respect to a transformation group), has acquired more and more importance in modern physics. This Chapter explores in 8 Sections the meaning, application and interpretation of symmetry in classical physics. This is done both in general, and with attention to specific topics. The general topics include illustration of the distinctions between symmetries of objects and of laws, and between symmetry principles and symmetry arguments (such as Curie's principle), and (...) reviewing the meaning and various types of symmetry that may be found in classical physics, along with different interpretative strategies that may be adopted. Specific topics discussed include the historical path by which group theory entered classical physics, transformation theory in classical mechanics, the relativity principle in Einstein's Special Theory of Relativity, general covariance in his General Theory of Relativity, and Noether's theorems. In bringing these diverse materials together in a single Chapter, we display the pervasive and powerful influence of symmetry in classical physics, and offer a possible framework for the further philosophical investigation of this topic. (shrink)

Symmetries play a major role in physics, in particular since the work by E. Noether and H. Weyl in the first half of last century. Herein, we briefly review their role by recalling how symmetry changes allow to conceptually move from classical to relativistic and quantum physics. We then introduce our ongoing theoretical analysis in biology and show that symmetries play a radically different role in this discipline, when compared to those in current physics. By this comparison, we stress that (...) symmetries must be understood in relation to conservation and stability properties, as represented in the theories. We posit that the dynamics of biological organisms, in their various levels of organization, are not just processes, but permanent (extended, in our terminology) critical transitions and, thus, symmetry changes. Within the limits of a relative structural stability (or interval of viability), variability is at the core of these transitions. (shrink)