It's been 41 years since the publication of Ernst Mayr's Cause and Effect in Biology wherein Mayr most clearly develops his version of the influential distinction between ultimate and proximate causes in biology. In critically assessing Mayr's essay I uncover false statements and red-herrings about biological explanation. Nevertheless, I argue to uphold an analogue of the ultimate/proximate distinction as it refers to two different kinds of explanations, one dynamical the other statistical.

The survival enhancing propensity (SEP) account has a crucial role to play in the analysis of proper function. However, a central feature of the account, its specification of the proper environment to which functions are relativized, is seriously underdeveloped. In this paper, I argue that existent accounts of proper environment fail because they either allow too many or too few characters to count as proper functions. While SEP accounts retain their promise, they are unworkable because of their inability to specify (...) this important feature. However, I suggest that this problem can be overcome by the application of a new strategy for specifying proper environment that is grounded in the operation of natural selection and I conclude by offering a first approximation of such an account. (shrink)

Selection explanations explain some non-accidental generalizations in virtue of a selection process. Such explanations are not particulaizable - they do not transfer as explanations of the instances of such generalizations. This is unlike many explanations in the physical sciences, where the explanation of the general fact also provides an explanation of its instances (i.e. standard D-N explanations). Are selection explanations (e.g. in biology) therefore a different kind of explanation? I argue that to understand this issue, we need to see that (...) a standard D-N explanation of some non-accidental generalization (al Fs are Gs) may also ipso facto explain its contrapositive (all non-Gs are non-Fs), but the explanation is particularizable with respect to the former but not to the latter. This can be seen by noting that the Raven Paradox counterexample to the H-D model of confirmation also generates a counterexample to the D-N model of explanation (all ravens are black does not explain why the non-black shoe is a non-raven). In such cases it is natural to take the generalization with the positive predicates to have a particularizable explanation. However, this need not be the case, and in selection explanations it is the generalization with the positive predicates whose explanation is no particularizable. Thus there is no need to suppose that selection explanations are fundamentally different. (shrink)

E. O. Wilson (1974: 54) describes the problem that social organisms pose: “On what bases do we distinguish the extremely modified members of an invertebrate colony from the organs of a metazoan animal?” This framing of the issue has inspired many to look more closely at how groups of organisms form and behave as emergent individuals. The possible existence of “superorganisms” test our best intuitions about what can count and act as genuine biological individuals and how we should study them. (...) As we will discuss, colonies of certain organisms display many of the properties that we usually reserve only to individual organisms. Although there is good reason to believe that many social insects form genuine emergent biological individuals, the conclusion offered here is of a slightly different sort. I will argue that to understand some social insects' interactions and the emergent traits they give rise to, it may be helpful to shift our understanding from a community-level approach to an ecosystem-level approach. I will argue that viewing certain insect colonies (termites) as parts of ecosystems allows us to better understand some of the adaptations that have emerged from their evolution. (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)

In this paper, using a multilevel approach, we defend the positive role of natural selection in the generation of organismal form. Despite the currently widespread opinion that natural selection only plays a negative role in the evolution of form, we argue, in contrast, that the Darwinian factor is a crucial (but not exclusive) factor in morphological organization. Analyzing some classic arguments, we propose incorporating the notion of ‘downward causation’ into the concept of ‘natural selection.’ In our opinion, this kind of (...) causation is fundamental to the operation of selection as a creative evolutionary process. (shrink)

The capacity to engage with art is a human universal present in all cultures and just about every individual human. This indicates that this capacity is evolved. In this Critical Notice of Denis Dutton's The Art Instinct, I discuss various evolutionary scenarios and their consequences. Dutton and I both reject the "spandrel" approach that originates from the work of Gould and Lewontin. Dutton proposes, following work of Geoffrey Miller, that art is sexually selected--that art-production is a sign of a fit (...) genome in males. I argue that while assortative mating may well have had a role in the evolution of "the art instinct", group selection is a better explanation. I also take issue with Dutton's "cluster concept" approach to defining art, and argue that it is a universal and essential characteristic of art that it is appreciated both for what it expresses and for the way that it expresses. It thus requires a reflexive capacity that is not operative in the appreciation of sport spectacles and pornography. (shrink)

The statistical interpretation of the Theory of Natural Selection claims that natural selection and drift are statistical features of mathematical aggregates of individual-level events. Natural selection and drift are not themselves causes. The statistical interpretation is motivated by a metaphysical conception of individual priority. Recently, Millstein, Skipper, and Dietrich (2009) have argued (a) that natural selection and drift are physical processes, and (b) that the statistical interpretation rests on a misconception of the role of mathematics in biology. Both theses are (...) contested. (shrink)

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: mohan.matthen@utoronto.ca. (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.

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.

Recently, much philosophical discussion has centered on the best way to characterize the concepts of random drift1 and natural selection, and, in particular, whether selection and drift can be conceptually distinguished (Beatty, 1984; Brandon, 2005; Hodge, 1983, 1987; Millstein, 2002, 2005; PError: Illegal entry in bfchar block in ToUnicode CMapfeifer, 2005; Shanahan, 1992; Stephens, 2004).2 These authors all contend, to a greater or lesser degree, that their concepts make sense of biological practice. So it should be instructive to see how (...) the concepts of drift and selection were distinguished by the disputants in a high-profile debate; debates such as these often force biologists to take a more philosophical turn, discussing the concepts at issue in greater detail than usual. Moreover, it is important to consider a debate where the disputants are actually trying to apply the models of population genetics to natural populations; only then can their proper interpretations become fully apparent. (Indeed, I contend that some of the philosophical confusion has arisen because authors have considered only the models themselves, and not the phenomena that the models are attempting to represent). A prime candidate for just such a case study is what Provine (1986) has termed “The Great Snail Debate,” that is, the debates over the highly polymorphic land snails Cepaea nemoralis and C. hortensis in the 1950s and early 1960s. These studies represent one of the best, if not the best, of the early attempts to demonstrate drift in natural populations.3 Of course, some excellent historical accounts of the changes in the usages and meanings of random drift have been written already. Some of these histories of random drift have a broad scope (Beatty, 1992; Cain & Provine, 1991; Gigerenzer et al. (shrink)

Recent discussions in the philosophy of biology have brought into question some fundamental assumptions regarding evolutionary processes, natural selection in particular. Some authors argue that natural selection is nothing but a population-level, statistical consequence of lower-level events (Matthen and Ariew [2002]; Walsh et al. [2002]). On this view, natural selection itself does not involve forces. Other authors reject this purely statistical, population-level account for an individual-level, causal account of natural selection (Bouchard and Rosenberg [2004]). I argue that each of these (...) positions is right in one way, but wrong in another; natural selection indeed takes place at the level of populations, but it is a causal process nonetheless. Introduction A brief justification of population-level causality 2.1 Frequency-dependent selection 2.2 Accounts of causation The montane willow leaf beetle: a causal story The montane willow leaf beetle: a population-level story 4.1 Response to ‘naïve individualism’ 4.2 Response to ‘sophisticated individualism’ Conclusion. (shrink)

Two strong arguments have been given in favor of the claim that no selection process can play a role in explaining adaptations. According to the first argument, selection is a negative force; it may explain why the eliminated individuals are eliminated, but it does not explain why the ones that survived (or their offspring) have the traits they have. The second argument points out that the explanandum and the explanans are phenomena at different levels: selection is a population-level phenomenon, whereas (...) adaptation occurs on the individual level. Thus, selection can explain why individuals in a certain population have a certain trait, but it cannot explain why a certain indi- vidual has this trait. After pointing out that both arguments ignore the significance of the limitation of environmental resources, I will construe a positive argument for the claim that cumulative selection processes can, indeed, play a role in explaining adaptations. (shrink)

Instead of using only one notion of selection I argue for a broader typology of different types of selection. Three such types are differentiated, namely simple one-step selection, iterated one-step selection, and multi-step selection. It is argued that this more general and more inclusive typology might face more effectively the possible challenges of a general account of selection.

In recent years, biologists have increasingly been asking whether the ability to evolve — the evolvability — of biological systems, itself evolves, and whether this phenomenon is the result of natural selection or a by-product of other evolutionary processes. The concept of evolvability, and the increasing theoretical and empirical literature that refers to it, may constitute one of several pillars on which an extended evolutionary synthesis will take shape during the next few years, although much work remains to be done (...) on how evolvability comes about. (shrink)

The study of phenotypic plasticity has progressed significantly over the past few decades. We have moved from variation for plasticity being considered as a nuisance in evolutionary studies to it being the primary target of investigations that use an array of methods, including quantitative and molecular genetics, as well as of several approaches that model the evolution of plastic responses. Here, I consider some of the major aspects of research on phenotypic plasticity, assessing where progress has been made and where (...) additional effort is required. I suggest that some areas of research, such the study of the quantitative genetic underpinning of plasticity, have been either settled in broad outline or superseded by new approaches and questions. Other issues, such as the costs of plasticity are currently at the forefront of research in this field, and are likely to be areas of major future development. (shrink)

The idea of genetic assimilation, that environmentally induced phenotypes may become genetically fixed and no longer require the original environmental stimulus, has had varied success through time in evolutionary biology research. Proposed by Waddington in the 1940s, it became an area of active empirical research mostly thanks to the efforts of its inventor and his collaborators. It was then attacked as of minor importance during the ‘‘hardening’’ of the neo-Darwinian synthesis and was relegated to a secondary role for decades. Recently, (...) several papers have appeared, mostly independently of each other, to explore the likelihood of genetic assimilation as a biological phenomenon and its potential importance to our understanding of evolution. In this article we briefly trace the history of the concept and then discuss theoretical models that have newly employed genetic assimilation in a variety of contexts. We propose a typical scenario of evolution of genetic assimilation via an intermediate stage of phenotypic plasticity and present potential examples of the same. We also discuss a conceptual map of current and future lines of research aimed at exploring the actual relevance of genetic assimilation for evolutionary biology. (shrink)

The idea of genetic assimilation, that environmentally induced phenotypes may become genetically fixed and no longer require the original environmental stimulus, has had varied success through time in evolutionary biology research. Proposed by Waddington in the 1940s, it became an area of active empirical research mostly thanks to the efforts of its inventor and his collaborators. It was then attacked as of minor importance during the ‘‘hardening’’ of the neo-Darwinian synthesis and was relegated to a secondary role for decades. Recently, (...) several papers have appeared, mostly independently of each other, to explore the likelihood of genetic assimilation as a biological phenomenon and its potential importance to our understanding of evolution. In this article we briefly trace the history of the concept and then discuss theoretical models that have newly employed genetic assimilation in a variety of contexts. We propose a typical scenario of evolution of genetic assimilation via an intermediate stage of phenotypic plasticity and present potential examples of the same. We also discuss a conceptual map of current and future lines of research aimed at exploring the actual relevance of genetic assimilation for evolutionary biology. (shrink)

In addition to considerable debate in the recent evolutionary literature about the limits of the Modern Synthesis of the 1930s and 1940s, there has also been theoretical and empirical interest in a variety of new and not so new concepts such as phenotypic plasticity, genetic assimilation and phenotypic accommodation. Here we consider examples of the arguments and counter- arguments that have shaped this discussion. We suggest that much of the controversy hinges on several misunderstandings, including unwarranted fears of a general (...) attempt at overthrowing the Modern Synthesis paradigm, and some fundamental conceptual confusion about the proper roles of phenotypic plasticity and natural selection within evolutionary theory. (shrink)

I have recently argued that origin essentialism regarding individual organisms entails that natural selection does not explain why individual organisms have the traits that they do. This paper defends this and related theses against Mohan Matthen's recent objections.

Does natural selection explain why individual organisms have the traits that they do? According to "the Negative View," natural selection does not explain why any individual organism has the traits that it does. According to "the Positive View," natural selection at least sometimes does explain why an individual organism has the traits that it does. In this paper, I argue that recent arguments for the Positive View fail in virtue of running afoul of the doctrine of origin essentialism and I (...) demonstrate that other recent defenses of the Negative View depend upon my own for their plausibility. (shrink)

The explanatory role of natural selection is one of the long-term debates in evolutionary biology. Nevertheless, the consensus has been slippery because conceptual confusions and the absence of a unified, formal causal model that integrates different explanatory scopes of natural selection. In this study we attempt to examine two questions: (i) What can the theory of natural selection explain? and (ii) Is there a causal or explanatory model that integrates all natural selection explananda? For the first question, we argue that (...) five explananda have been assigned to the theory of natural selection and that four of them may be actually considered explananda of natural selection. For the second question, we claim that a probabilistic conception of causality and the statistical relevance concept of explanation are both good models for understanding the explanatory role of natural selection. We review the biological and philosophical disputes about the explanatory role of natural selection and formalize some explananda in probabilistic terms using classical results from population genetics. Most of these explananda have been discussed in philosophical terms but some of them have been mixed up and confused. We analyze and set the limits of these problems. (shrink)

This paper explores whether natural selection, a putative evolutionary mechanism, and a main one at that, can be characterized on either of the two dominant conceptions of mechanism, due to Glennan and the team of Machamer, Darden, and Craver, that constitute the “new mechanistic philosophy.” The results of the analysis are that neither of the dominant conceptions of mechanism adequately captures natural selection. Nevertheless, the new mechanistic philosophy possesses the resources for an understanding of natural selection under the rubric.

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)

We distinguish dynamical and statistical interpretations of evolutionary theory. We argue that only the statistical interpretation preserves the presumed relation between natural selection and drift. On these grounds we claim that the dynamical conception of evolutionary theory as a theory of forces is mistaken. Selection and drift are not forces. Nor do selection and drift explanations appeal to the (sub-population-level) causes of population level change. Instead they explain by appeal to the statistical structure of populations. We briefly discuss the implications (...) of the statistical interpretation of selection for various debates within the philosophy of biologythe `explananda of selection' debate and the `units of selection' debate. (shrink)

Few problems in the philosophy of evolutionary biology are more widely disseminated and discussed than the charge of Darwinian evolution being a tautology. The history is long and complex, and the issues are many, and despite the problem routinely being dismissed as an introductory-level issue, based on misunderstandings of evolution, it seems that few agree on what exactly these misunderstandings consist of. In this paper, I will try to comprehensively review the history and the issues. Then, I will try to (...) present the following “solution”, or, one might say, “dissolution”, of the problem, and consider the wider implications of formal, or schematic, explanations in science: yes, the principle of natural selection is a tautology, and so what? It is a promissory note for actual, physical, explanations in particular cases, and is none the worse for that. This is not a new argument, of course, but it does point up the importance of formal schematic models in science. (shrink)

Darwin’s pluralism, then and now Content Type Journal Article Pages 1-5 DOI 10.1007/s11016-011-9578-5 Authors Rasmus Grønfeldt Winther, Philosophy Department, University of California, Santa Cruz, 1156 High St., Santa Cruz, CA 95064, USA Journal Metascience Online ISSN 1467-9981 Print ISSN 0815-0796.

Selectionist evolutionary theory has often been faulted for not making novel predictions that are surprising, risky, and correct. I argue that it in fact exhibits the theoretical virtue of predictive capacity in addition to two other virtues: explanatory unification and model fitting. Two case studies show the predictive capacity of selectionist evolutionary theory: parallel evolutionary change in E. coli and the origin of eukaryotic cells through endosymbiosis. †To contact the author, please write to: Philosophy Department, University of California, Santa Cruz, (...) 1156 High St., Santa Cruz, CA 95064; e‐mail: rgw@ucsc.edu ; rgwinther@gmail.com. (shrink)

I investigate how theoretical assumptions, pertinent to different perspectives and operative during the modeling process, are central in determining how nature is actually taken to be. I explore two different models by Michael Turelli and Steve Frank of the evolution of parasite-mediated cytoplasmic incompatility, guided, respectively, by Fisherian and Wrightian perspectives. Since the two models can be shown to be commensurable both with respect to mathematics and data, I argue that the differences between them in the (1) mathematical presentation of (...) the models, (2) explanations, and (3) objectified ontologies stem neither from differences in mathematical method nor the employed data, but from differences in the theoretical assumptions, especially regarding ontology, already present in the respective perspectives. I use my "set up, mathematically manipulate, explain, and objectify" (SMEO) account of the modeling process to track the model-mediated imposition of theoretical assumptions. I conclude with a discussion of the general implications of my analysis of these models for the controversy between Fisherian and Wrightian perspectives. (shrink)