Schaffner’s model of theory reduction has played an important role in philosophy of science and philosophy of biology. Here, the model is found to be problematic because of an internal tension. Indeed, standard antireductionist external criticisms concerning reduction functions and laws in biology do not provide a full picture of the limits of Schaffner’s model. However, despite the internal tension, his model usefully highlights the importance of regulative ideals associated with the search for derivational, and embedding, deductive relations among mathematical (...) structures in theoretical biology. A reconstructed Schaffnerian model could therefore shed light on mathematical theory development in the biological sciences and on the epistemology of mathematical practices more generally. *Received November 2006; revised March 2009. †To contact the author, please write to: Philosophy Department, University of California, Santa Cruz, 1156 High St., Santa Cruz, CA 95064; e‐mail: [email protected]. (shrink)

The dangers of character reification for cladistic inference are explored. The identification and analysis of characters always involves theory-laden abstraction—there is no theory-free “view from nowhere.” Given theory-ladenness, and given a real world with actual objects and processes, how can we separate robustly real biological characters from uncritically reified characters? One way to avoid reification is through the employment of objectivity criteria that give us good methods for identifying robust primary homology statements. I identify six such criteria and explore each (...) with examples. Ultimately, it is important to minimize character reification, because poor character analysis leads to dismal cladograms, even when proper phylogenetic analysis is employed. Given the deep and systemic problems associated with character reification, it is ironic that philosophers have focused almost entirely on phylogenetic analysis and neglected character analysis. (shrink)

This paper provides a philosophical analysis of the ongoing controversy surrounding R.A. Fisher's famous ‘fundamental theorem’ of natural selection. The difference between the ‘traditional’ and ‘modern’ interpretations of the theorem is explained. I argue that proponents of the modern interpretation have captured Fisher's intended meaning correctly and shown that the theorem is mathematically correct, pace the traditional consensus. However, whether the theorem has any real biological significance remains an unresolved issue. I argue that the answer depends on whether we accept (...) Fisher's non-standard notion of environmental change, on which the theorem rests; arguments for and against this notion are explored. I suggest that there is a close link between Fisher's fundamental theorem and the modern ‘gene's eye’ view of evolution. Introduction What Does the Fundamental Theorem Say? Key Concepts Explained Alleged Significance of the FTNS Traditional versus Modern Interpretations of the FTNS The Modern Interpretation Illustrated Fisher's Concept of ‘Environmental Change’ Causality and the Modern Interpretation The Significance of the FTNS Re-considered Appendix CiteULike Connotea Del.icio.us What's this? (shrink)

Darwinism designates a distinctive form of evolutionary explanation for the history and diversity of life on earth. Its original formulation is provided in the first edition of On the Origin of Species in 1859. This entry first formulates ‘Darwin's Darwinism’ in terms of five philosophically distinctive themes: (i) probability and chance, (ii) the nature, power and scope of selection, (iii) adaptation and teleology, (iv) nominalism vs. essentialism about species and (v) the tempo and mode of evolutionary change. Both Darwin and (...) his critics recognized that his approach to evolution was distinctive on each of these topics, and it remains true that, though Darwinism has developed in many ways unforeseen by Darwin, its proponents and critics continue to differentiate it from other approaches in evolutionary biology by focusing on these themes. This point is illustrated in the second half of the entry by looking at current debates in the philosophy of evolutionary biology on these five themes, with a special focus on Stephen Jay Gould's The Structure of Evolutionary Theory. (shrink)

In this paper, we aim to show that the framework of embedded, distributed, or extended cognition offers new perspectives on social cognition by applying it to one specific domain: the psychology of memory. In making our case, first we specify some key social dimensions of cognitive distribution and some basic distinctions between memory cases, and then describe stronger and weaker versions of distributed remembering in the general distributed cognition framework. Next, we examine studies of social influences on memory in cognitive (...) psychology, and identify the valuable concepts and methods to be extended and embedded in our framework; we focus in particular on three related paradigms: transactive memory, collaborative recall, and social contagion. Finally, we sketch our own early studies of individual and group memory developed within our framework of distributed cognition, on social contagion of autobiographical memories, collaborative flashbulb memories, and memories of high school at a high school reunion. We see two reciprocal benefits of this conceptual and empirical framework to social memory phenomena: that ideas about distributed cognition can be honed against and tested with the help of sophisticated methods in the social cognitive psychology of memory; and conversely, that a range of social memory phenomena that are as yet poorly understood can be approached afresh with theoretically motivated extensions of existing empirical paradigms. (shrink)

Change is the fundamental idea of evolution. Explaining the extraordinary biological change we see written in the history of genomes and fossil beds is the primary occupation of the evolutionary biologist. Yet it is a surprising fact that for the majority of evolutionary research, we have rarely studied how evolution typically unfolds in nature, in changing ecological environments, over space and time. While ecology played a major role in the eventual acceptance of the population genetic viewpoint of evolution in the (...) synthetic era (circa 1918-1956), it held a lesser role in the development of evolutionary theory until the 1980s, when we began to systematically study the evolutionary dynamics of natural populations in space and time. As a result, early evolutionary theory was initially constructed in an abstract vacuum that was unrepresentative of evolution in nature. The subtle synthesis between ecology with evolutionary biology (eco-evo synthesis) over the past 40 years has progressed our knowledge of natural selection dynamics as they are found in nature, thus revealing how natural selection varies in strength, direction, form, and, more surprisingly, level of biological organization. Natural selection can no longer be reduced to lower levels of biological organization (i.e., individuals, selfish genes) over shorter timescales but should be expanded to include adaptation at higher levels and over longer timescales. Long-term and/or emergent evolutionary phenomena, such as multilevel selection or evolvability, have thus become tenable concepts within an evolutionary biology that embraces ecology and spatiotemporal change. Evolutionary biology is currently suspended at an intermediate stage of scientific progress that calls for the organization of all the recent knowledge revealed by the eco-evo synthesis into a coherent and unified theoretical framework. This is where philosophers of biology can be of particular use, acting as a bridge between the subdisciplines of biology and inventing new theoretical strategies to organize and accommodate the recent knowledge. Philosophers have recommended transitioning away from outdated philosophies that were originally derived from physics within the philosophical zeitgeist of logical positivism (i.e., monism, reductionism, and monocausation) and toward a distinct philosophy of biology that can capture the natural complexity of multifaceted biological systems within diverse ecosystems—one that embraces the emerging philosophies of pluralism, emergence, and multicausality. Therefore, I see recent advances in ecology, evolutionary biology, and the philosophy of biology as laying the groundwork for another major biological synthesis, what I refer to as the Second Synthesis because, in many respects, it is analogous to the aims and outcomes of the first major biological synthesis (but is notably distinct from the inorganic and contrived progressive movement known as the extended evolutionary synthesis). With the general development of a distinctive philosophy of science, biology has rightfully emerged as an autonomous science. Thus, while the first synthesis legitimized biology, the Second Synthesis autonomized biology and afforded biology its own philosophy, allowing biology to finally realize its full scientific potential. (shrink)

Why do we have social norms—of fairness, cooperation, trust, property, or gender? Modern-day Humeans, as I call them, believe these norms are best accounted for in cultural evolutionary terms, as adaptive solutions to recurrent problems of social interaction. In this paper, I discuss a challenge to this “Humean Program.” Social norms involve widespread behaviors, but also distinctive psychological attitudes and dispositions. According to the challenge, Humean accounts of norms leave their psychological side unexplained. They explain, say, why we share equally, (...) but not why we disapprove of those who don’t. I defend the Humean Program against this challenge. In particular, I suggest an idea for how to extend the Program to account for the psychological side of norms. Socially adaptive behaviors aren’t just likely to emerge in a group; They are also likely to be widely taught within it. The transmission of these behaviors through instruction explains why they are associated with distinctive normative attitudes and dispositions. These attitudes play a pedagogical role in helping transmit these behaviors to children and newcomers. (shrink)

Even after Willard Quine’s critique of the analytic-synthetic distinction in “Two Dogmas of Empiricism,” Wilfrid Sellars maintained some forms of analyticity or truth in virtue of meaning. In this article, I aim to reconstruct (a) his neglected account of the analytic-synthetic distinction and the revisability of analytic sentences, (b) its connection to his inferentialist account of meaning, and (c) his response to Quine. While Sellars’s account of the revisability of analytic sentences bears certain similarities to Carnap’s and Grice and Strawson’s (...) accounts, it is still striking in that it is part of his broader picture of how our language develops dynamically in our ongoing inquiry. I aim to show how it relates to his account of how the diachronic continuity in meaning, language, or conceptual framework can be preserved through a revision of analytic sentences and to his evolutionary account of the development of our language or conceptual framework. (shrink)

Multiple biological groups, such as ant colonies, appear to have a noteworthy inefficiency: they contain vast amounts of redundant members that are not strictly needed to maintain the group. Philosophers and biologists have proposed that such inefficiency is illusory because redundancy enhances the resilience of groups when living under harsh conditions. Still, this proposal is unsatisfactory in different respects. First, it is too vague to account for when redundancy is selectively advantageous. Furthermore, it overlooks cases in which redundancy fails to (...) increase the resilience of groups. This paper offers an account of group redundancy that addresses these difficulties. Specifically, it advances the hypothesis that the mere presence of harsh conditions is not what drives the evolution of redundancy; rather, it is the fact organisms are often unable to predict when they will face harsh conditions. Redundancy enables groups to properly respond to unpredictable circumstances without resorting to the unreliable detection systems of their members. A better understanding of the phenomenon of redundancy is likely to impact other key issues in philosophy of science, including the evolution of cooperation and transitions in individuality, and the role of redundancy in complex systems, be they biological or not. (shrink)

I consider two attempts to combine realism with pluralism about the units of selection: Sober and Wilson’s combination of “model” and “unit” pluralism, and Sterelny and Griffiths’ “local pluralism”. I argue that both of these attempts fail to show that realism and pluralism are compatible. Sober and Wilson’s pluralism turns out, on closer inspection, to be a kind of monism in disguise, while Sterelny and Griffiths’ local pluralism involves a combination of realism and anti-realism about interactors, and the units of (...) selection, that is fundamentally unstable. My conclusion is that one must choose whether to be a realist or a pluralist in this area: one cannot be both. The question of which we should choose is a further question that I do not take a stand on. (shrink)

“Teleosemantic” or “biosemantic” theories form a strong naturalistic programme in the philosophy of mind and language. They seek to explain the nature of mind and language by recourse to a natural history of “proper functions” as selected-for effects of language- and thought-producing mechanisms. However, they remain vague with respect to the nature of the proposed analogy between selected-for effects on the biological level and phenomena that are not strictly biological, such as reproducible linguistic and cultural forms. This essay critically explores (...) various interpretations of this analogy. It suggests that these interpretations can be explicated by contrasting adaptationist with pluralist readings of the evolutionary concept of adaptation. Among the possible interpretations of the relations between biological adaptations and their analogues in language and culture, the two most relevant are a linear, hierarchical, signalling-based model that takes its cues from the evolution of co-operation and joint intentionality and a mutualistic, pluralist model that takes its cues from mimesis and symbolism in the evolution of human communication. Arguing for the merits of the mutualistic model, the present analysis indicates a path towards an evolutionary pluralist version of biosemantics that will align with theories of cognition as being environmentally “scaffolded”. Language and other cultural forms are partly independent reproducible structures that acquire proper functions of their own while being integrated with organism-based cognitive traits in co-evolutionary fashion. (shrink)

This special issue for the Journal for General Philosophy of Science is devoted to exploring the impact and many ramifications of current research in evolutionary epistemology. Evolutionary epistemology is an inter- and multidisciplinary area of research that can be divided into two ever-inclusive research avenues. One research avenue expands on the EEM program and investigates the epistemology of evolution. The other research avenue builds on the EET program and researches the evolution of epistemology. Since its conception, EE has developed three (...) schools of thought: adaptationist, non-adaptationist, and applied EE. Although diverse in outlook and theoretical background, these research avenues and schools share the same agenda of understanding how knowledge evolves, and how it relates to the world. In this paper, we first explain wherefrom evolutionary epistemological schools of thought developed, and then we highlight current debates in EE by briefly reviewing the papers that form part of this special issue. (shrink)

In this paper, I describe some strategies for teaching an introductory philosophy of science course to Science, Technology, Engineering, and Mathematics (STEM) students, with reference to my own experience teaching a philosophy of science course in the Fall of 2020. The most important strategy that I advocate is what I call the “Second Philosophy” approach, according to which instructors ought to emphasize that the problems that concern philosophers of science are not manufactured and imposed by philosophers from the outside, but (...) rather are ones that arise internally, during the practice of science itself. To justify this approach, I appeal to considerations from both educational research and the epistemology of testimony. In addition, I defend some distinctive learning goals that philosophy of science instructors ought to adopt when teaching STEM students, which include rectifying empirically well-documented shortcomings in students’ conceptions of the “scientific method” and the “nature of science.” Although my primary focus will be on teaching philosophy of science to STEM students, much of what I propose can be applied to non-philosophy majors generally. Ultimately, as I argue, a successful philosophy of science course for non-philosophy majors must be one that advances a student’s science education. The strategies that I describe and defend here are aimed at precisely that end. (shrink)

It is widely assumed that there is value in the biological tie between parent and child. An implication of this is that adoption is often considered a less desirable alternative to procreation. This paper offers a philosophical defence of adoptive parenthood as a valuable and authentic form of parenthood. While previous defences have suggested that society’s valorisation of the biological tie is unjustified, I argue herein that the conception of the biological tie that features in the normative discourse on parenthood (...) is too narrowly genocentric. Against this genocentric conception, recent work in the philosophy of biology has emphasised the roles of joint determination, dynamic construction, and extended inheritance in development, which I suggest can substantiate a more inclusive conception of the biological tie. Accordingly, I propose that adoptive parents form a rich variety of biological ties with their children, some of which are as heritable and formative as genetic relatedness. (shrink)

There is a certain metaphor that has enjoyed tremendous longevity in the evolution of ageing literature. According to this metaphor, nature has a certain goal or purpose, the perpetuation of the species, or, alternatively, the reproductive success of the individual. In relation to this goal, the individual organism has a function, job, or task, namely, to breed and, in some species, to raise its brood to maturity. On this picture, those who cannot, or can no longer, reproduce are somehow invisible (...) to, or even dispensable to, the evolutionary process. Here, I argue that the metaphor should be discarded, not on the grounds that it is a metaphor, but on the grounds that this particular metaphor distorts our understanding of the evolution of ageing. One reason the metaphor is problematic is that it frames senescence and death as nature’s verdict on the value of older individuals. Instead, we should explore a different metaphor: the lengthy post-reproductive period in humans and some other animals is not an accident of culture, but designed by nature for the purpose of supporting and guiding younger generations. On this alternate picture, different stages of life have their own evolutionary rationales, their distinctive design features, their special mandates. (shrink)

Evolution by natural selection has been extended to several supraorganismic levels, but whether it can apply to ecosystems remains controversial on two main counts. First, local ecosystems are loosely individuated, so that it is unclear how they manifest heredity and fitness. Second, even if they did, the meta-ecosystem formed by this population of local ecosystems will also suffer from a very low degree of cohesion, which will jeopardize any ENS. We suggest a way to overcome both issues, focusing on ecosystem (...) properties rather than on ecosystemic individuals. First, we follow recent theoretical developments which deny the centrality of reproduction in ENS. This leads us to merge heredity and fitness in a single process, in which a local ecosystem causally influences its neighbors, in a way responsible for their phenotypic similarity. Second, we suggest that the resulting meta-ecosystem need not meet the criteria of individuality. Instead, the evolving meta-ecosystem is best understood as a homeostatic property cluster, i.e., a set of phenotypes that together evolve by ENS at the ecosystem level. We illustrate this conceptual framework with a hypothetical example of local ecosystems that vary in terms of stability, productivity, diversity, and complementarity between species. Finally, we stress that our property-based account of ecosystemic ENS can help understand evolution at other biological levels, such as early life evolution and holobionts. (shrink)

Contemporary biological research has suggested that some host–microbiome multispecies systems (referred to as “holobionts”) can in certain circumstances evolve as unique biological individual, thus being a unit of selection in evolution. If this is so, then it is arguably the case that some biological adaptations have evolved at the level of the multispecies system, what we call hologenomic adaptations. However, no research has yet been devoted to investigating their nature, or how these adaptations can be distinguished from adaptations at the (...) species-level (genomic adaptations). In this paper, we cover this gap by investigating the nature of hologenomic adaptations. By drawing on the case of the evolution of sanguivory diet in vampire bats, we argue that a trait constitutes a hologenomic adaptation when its evolution can only be explained if the holobiont is considered the biological individual that manifests this adaptation, while the bacterial taxa that bear the trait are only opportunistic beneficiaries of it. We then use the philosophical notions of emergence and inter-identity to explain the nature of this form of individuality and argue why it is special of holobionts. Overall, our paper illustrates how the use of philosophical concepts can illuminate scientific discussions, in the trend of what has recently been called metaphysics of biology. (shrink)

Bourrat and Griffiths :33, 2018) have recently argued that most of the evidence presented by holobiont defenders to support the thesis that holobionts are evolutionary individuals is not to the point and is not even adequate to discriminate multispecies evolutionary individuals from other multispecies assemblages that would not be considered evolutionary individuals by most holobiont defenders. They further argue that an adequate criterion to distinguish the two categories is fitness alignment, presenting the notion of fitness boundedness as a criterion that (...) allows divorcing true multispecies evolutionary individuals from other multispecies assemblages and provides an adequate criterion to single out genuine evolutionary multispecies assemblages. A consequence of their criterion is that holobionts, as conventionally defined by hologenome defenders, are not evolutionary individuals except in very rare cases, and for very specific host-symbiont associations. This paper is a critical response to Bourrat and Griffiths’ arguments and a defence of the arguments presented by holobiont defenders. Drawing upon the case of the hologenomic basis of the evolution of sanguivory in vampire bats, I argue that Bourrat and Griffiths overlook some aspects of the biological nature of the microbiome that justifies the thesis that holobionts are evolutionarily different to other multispecies assemblages. I argue that the hologenome theory of evolution should not define the hologenome as a collection of genomes, but as the sum of the host genome plus some traits of the microbiome which together constitute an evolutionary individual, a conception I refer to as the stability of traits conception of the hologenome. Based on that conception I argue that the evidence presented by holobiont defenders is to the point, and supports the thesis that holobionts are evolutionary individuals. In this sense, the paper offers an account of the holobiont that aims to foster a dialogue between hologenome advocates and hologenome critics. (shrink)

Lucas Matthews and I substantially revised my SEP entry on Heritability. This version includes discussion of the missing heritability problem and other issues that arise from the use of Genome Wide Association Studies by Behavioral Geneticists.

The religious phenomenon is a complex one in many respects. In recent years an increasing number of theories on the origin and evolution of religion have been put forward. Each one of these theories rests on a Darwinian framework but there is a lot of disagreement about which bits of the framework account best for the evolution of religion. Is religion primarily a by-product of some adaptation? Is it itself an adaptation, and if it is, does it benefi ciate individuals (...) or groups? In this chapter, I review a number of theories that link religion to cooperation and show that these theories, contrary to what is often suggested in the literature, are not mutually exclusive. As I present each theory, I delineate an integrative framework that allows distinguishing the explanandum of each theory. Once this is done, it becomes clear that some theories provide good explanations for the origin of religion but not so good explanations for its maintenance and vice versa. Similarly some explanations are good explanations for the evolution of religious individual level traits but not so good explanations for traits hard to defi ne at the individual level. I suggest that to fully understand the religious phenomenon, integrating in a systematic way the different theories and the data is a more successful approach. (shrink)

The impressive variation amongst biological individuals generates many complexities in addressing the simple-sounding question what is a biological individual? A distinction between evolutionary and physiological individuals is useful in thinking about biological individuals, as is attention to the kinds of groups, such as superorganisms and species, that have sometimes been thought of as biological individuals. More fully understanding the conceptual space that biological individuals occupy also involves considering a range of other concepts, such as life, reproduction, and agency. There has (...) been a focus in some recent discussions by both philosophers and biologists on how evolutionary individuals are created and regulated, as well as continuing work on the evolution of individuality. (shrink)

The aim of this paper is to provide a theoretical framework to understand how multicellular systems realize functionally integrated physiological entities by organizing their intercellular space. From a perspective centered on physiology and integration, biological systems are often characterized as organized in such a way that they realize metabolic self-production and self-maintenance. The existence and activity of their components rely on the network they realize and on the continuous management of the exchange of matter and energy with their environment. One (...) of the virtues of the organismic approach focused on organization is that it can provide an understanding of how biological systems are functionally integrated into coherent wholes. Organismic frameworks have been primarily developed by focusing on unicellular life. Multicellularity, however, presents additional challenges to our understanding of biological systems, related to how cells are capable to live together in higher-order entities, in such a way that some of their features and behaviors are constrained and controlled by the system they realize. Whereas most accounts of multicellularity focus on cell differentiation and increase in size as the main elements to understand biological systems at this level of organization, we argue that these factors are insufficient to provide an understanding of how cells are physically and functionally integrated in a coherent system. In this paper, we provide a new theoretical framework to understand multicellularity, capable to overcome these issues. Our thesis is that one of the fundamental theoretical principles to understand multicellularity, which is missing or underdeveloped in current accounts, is the functional organization of the intercellular space. In our view, the capability to be organized in space plays a central role in this context, as it enables (and allows to exploit all the implications of) cell differentiation and increase in size, and even specialized functions such as immunity. We argue that the extracellular matrix plays a crucial active role in this respect, as an evolutionary ancient and specific (non-cellular) control subsystem that contributes as a key actor to the functional specification of the multicellular space and to modulate cell fate and behavior. We also analyze how multicellular systems exert control upon internal movement and communication. Finally, we show how the organization of space is involved in some of the failures of multicellular organization, such as aging and cancer. (shrink)

Is the mammalian embryo/fetus a part of the organism that gestates it? According to the containment view, the fetus is not a part of, but merely contained within or surrounded by, the gestating organism. According to the parthood view, the fetus is a part of the gestating organism. This paper proceeds in two stages. First, I argue that the containment view is the received view; that it is generally assumed without good reason; and that it needs substantial support if it (...) is to be taken seriously. Second, I argue that the parthood view derives considerable support from a range of biological and physiological considerations. I tentatively conclude in favour of the parthood view, and end by identifying some of the interesting questions it raises. (shrink)

For decades Darwinian processes were framed in the form of the Lewontin conditions: reproduction, variation and differential reproductive success were taken to be sufficient and necessary. Since Buss and the work of Maynard Smith and Szathmary biologists were eager to explain the major transitions from individuals to groups forming new individuals subject to Darwinian mechanisms themselves. Explanations that seek to explain the emergence of a new level of selection, however, cannot employ properties that would already have to exist on that (...) level for selection to take place. Recently, Hammerschmidt et al. provided a ‘bottom-up’ experiment corroborating much of the theoretical work Paul Rainey has done since 2003 on how cheats can play an important role in the emergence of new Darwinian individuals on a multicellular level. The aims of this paper are twofold. First, I argue for a conceptual shift in perspective from seeing cheats as a ‘problem’ that needs to be solved for multi-cellularity to evolve to the very ‘key’ for the evolution of multicellularity. Secondly, I illustrate the consequences of this shift for both theoretical and experimental work, arguing for a more prominent role of ecology and the multi-level selection framework within the debate then they currently occupy. (shrink)

In this essay review of Samir Okasha's Agents and Goals in Evolution, I reflect on the rationale for agential thinking in biology, and consider whether the rationale is the same for genes as for organisms. I also discuss Okasha's ingenious examples of the evolution of irrational behaviour, and in particular the evolution of violations of the "independence axiom" of rational choice theory. These examples rely on a crucial distinction between aggregate and idiosyncratic risk.

A crucial question for a process view of life is how to identify a process and how to follow it through time. The genidentity view can contribute decisively to this project. It says that the identity through time of an entity X is given by a well-identified series of continuous states of affairs. Genidentity helps address the problem of diachronic identity in the living world. This chapter describes the centrality of the concept of genidentity for David Hull and proposes an (...) extension of Hull’s view to the ubiquitous phenomenon of symbiosis. Finally, using immunology as a key example, it shows that the genidentity view suggests that the main interest of a process approach is epistemological rather than ontological and that its principal claim is one of priority, namely that processes precede and define things, and not vice versa. (shrink)

There is an active research program currently underway, which treats cancer progression as an evolutionary process. This contribution investigates the ways that cancer progression is like and unlike evolution in other contexts. The aim is to take a multi-level perspective on cancer, investigating the levels at which selection may be acting, the unit or target of selection, the relative roles of selection and drift, and the idea that cancer progression may be a by-product of selection at other levels of organization.

In multicellular organisms, cells are frequently programmed to die. This makes good sense: cells that fail to, or are no longer playing important roles are eliminated. From the cell’s perspective, this also makes sense, since somatic cells in multicellular organisms require the cooperation of clonal relatives. In unicellular organisms, however, programmed cell death poses a difficult and unresolved evolutionary problem. The empirical evidence for PCD in diverse microbial taxa has spurred debates about what precisely PCD means in the case of (...) unicellular organisms. In this article, we survey the concepts of PCD in the literature and the selective pressures associated with its evolution. We show that definitions of PCD have been almost entirely mechanistic and fail to separate questions concerning what PCD fundamentally is from questions about the kinds of mechanisms that realize PCD. We conclude that an evolutionary definition is best able to distinguish PCD from closely related phenomena. Specifically, we define “true” PCD as an adaptation for death triggered by abiotic or biotic environmental stresses. True PCD is thus not only an evolutionary product but must also have been a target of selection. Apparent PCD resulting from pleiotropy, genetic drift, or trade-offs is not true PCD. We call this “ersatz PCD.”. (shrink)

Recent debates between proponents of the modern evolutionary synthesis (the standard model in evolutionary biology) and those of a possible extended synthesis are a good example of the fascinating tangle among empirical, theoretical, and conceptual or philosophical matters that is the practice of evolutionary biology. In this essay, we briefly discuss two case studies from this debate, highlighting the relevance of philosophical thinking to evolutionary biologists in the hope of spurring further constructive cross-pollination between the two fields.

Sharon Street defines her constructivism about practical reasons as the view that whether something is a reason to do a certain thing for a given agent depends on that agent’s normative point of view. However, Street has also maintained that there is a judgment about practical reasons which is true relative to every possible normative point of view, namely constructivism itself. I show that the latter thesis is inconsistent with Street’s own constructivism about epistemic reasons and discuss some consequences of (...) this incompatibility. (shrink)

Evolutionary biology is striking for its ability to explain a large and diverse range of empirical phenomena on the basis of a few general theoretical principles. This article offers a philosophical perspective on the way that evolutionary biology has come to achieve such impressive generality, by focusing on “the strategy of endogenization”. This strategy involves devising evolutionary explanations for biological features that were originally part of the background conditions, or scaffolding, against which such explanations take place. Where successful, the strategy (...) moves biology a step closer to the ideal of explaining as much as possible from evolutionary first principles. The strategy of endogenization is illustrated through a series of biological examples, historical and recent, and its philosophical implications are explored. (shrink)

We assess the arguments for recognising functionally integrated multispecies consortia as genuine biological individuals, including cases of so-called ‘holobionts’. We provide two examples in which the same core biochemical processes that sustain life are distributed across a consortium of individuals of different species. Although the same chemistry features in both examples, proponents of the holobiont as unit of evolution would recognize one of the two cases as a multispecies individual whilst they would consider the other as a compelling case of (...) ecological dependence between separate individuals. Some widely used arguments in support of the ‘holobiont’ concept apply equally to both cases, suggesting that those arguments have misidentified what is at stake when seeking to identify a new level of biological individuality. One important aspect of biological individuality is evolutionary individuality. In line with other work on the evolution of individuality, we show that our cases can be distinguished by focusing on the fitness alignment between the partners of the consortia. We conclude that much of the evidence currently presented for the ubiquity and importance of multi-species individuals is simply not to the point, at least unless the issue of biological individuality is firmly divorced from the question of evolutionary individuality. (shrink)

Levels of organization are structures in nature, usually defined by part-whole relationships, with things at higher levels being composed of things at the next lower level. Typical levels of organization that one finds in the literature include the atomic, molecular, cellular, tissue, organ, organismal, group, population, community, ecosystem, landscape, and biosphere levels. References to levels of organization and related hierarchical depictions of nature are prominent in the life sciences and their philosophical study, and appear not only in introductory textbooks and (...) lectures, but also in cutting-edge research articles and reviews. In philosophy, perennial debates such as reduction, emergence, mechanistic explanation, interdisciplinary relations, natural selection, and many other topics, also rely substantially on the notion. -/- Yet, in spite of the ubiquity of the notion, levels of organization have received little explicit attention in biology or its philosophy. Usually they appear in the background as an implicit conceptual framework that is associated with vague intuitions. Attempts at providing general and broadly applicable definitions of levels of organization have not met wide acceptance. In recent years, several authors have put forward localized and minimalistic accounts of levels, and others have raised doubts about the usefulness of the notion as a whole. -/- There are many kinds of ‘levels’ that one may find in philosophy, science, and everyday life—the term is notoriously ambiguous. Besides levels of organization, there are levels of abstraction, realization, being, analysis, processing, theory, science, complexity, and many others. In this article, the focus will be on levels of organization and debates associated with them, and other kinds of levels will only be discussed when they are relevant to this main topic. (shrink)

Individuality is an important concept in biology, yet there are many non-equivalent criteria of individuality expressed in different kinds of biological individuals. This article evaluates these different kinds in terms of their capacity to support explanatory generalizations over the systems they individuate. Viewing the problem of individuality from this perspective promotes a splitting strategy in which different kinds make different epistemic trade-offs that suit them for different explanatory roles. I argue that evolutionary individuals, interpreted as forming a functional kind, face (...) difficulties of individuation and explanatory power that are mitigated by relying on more structurally based properties and non-evolutionary kinds. 1Introduction2Kinds of Biological Individuals3Evolutionary Individuals and Functional Kinds4Evolutionary versus Non-evolutionary Kinds of Individuals 4.1Physiological individuality4.2Ecological individuality4.3Developmental individuality5Conclusion. (shrink)

A darwinian evolutionary approach can contribute to reassess philosophical problems in different fields, including ethics and moral theory. Sociobiology and evolutionary psychology address these issues by presupposing mechanisms such as kin selection and reciprocal altruism. However, these mechanisms can’t account for cooperation in the human species. Dual inheritance theory addresses human cooperation differently, by taking into account the above-mentioned classical biological mechanisms without ignoring, however, relevant knowledge produced by social scientists. According to this approach, human social psychology comprises tribal social (...) instincts and symbolic markers. One implication of this approach is that there are innate and universal moral principles hardwired in the human mind-brain, which where selected through an evolutionary process that makes life possible in large, structured social groups. Although innate, these principles are plastically shaped to meet the demands of different cultural niches in particular societies. (shrink)

Evolutionary, ecological and ethical studies are, at the same time, specific scientific disciplines and, from an historical point of view, structurally linked domains of research. In a context of environmental crisis, the need is increasingly emerging for a connecting epistemological framework able to express a common or convergent tendency of thought and practice aimed at building, among other things, an environmental policy management respectful of the planet’s biodiversity and its evolutionary potential.

This paper attempts to reconcile critics and defenders of inclusive fitness by constructing a synthesis that does justice to the insights of both. I argue that criticisms of the regression-based version of Hamilton’s rule, although they undermine its use for predictive purposes, do not undermine its use as an organizing framework for social evolution research. I argue that the assumptions underlying the concept of inclusive fitness, conceived as a causal property of an individual organism, are unlikely to be exactly true (...) in real populations, but they are approximately true given a specific type of weak selection that Hamilton took, on independent grounds, to be responsible for the cumulative assembly of complex adaptation. Finally, I reflect on the uses and limitations of “design thinking” in social evolution research. The debate about the foundations of inclusive fitness theory that has followed in the wake of Nowak, Tarnita and Wilson’s (1) critique has been remarkably polarizing. After several rounds of rebuttals and replies, there is still little evidence of any serious reconciliation between the theory’s critics (2-10) and its defenders (11-24). It doesn’t have to be this way. I believe that, on the main points of disagreement, it is possible to find a way forward that does justice to the insights of both camps. My aim in this paper is to find that way forward. (shrink)

With an increasing number of ways to ‘assist’ reproduction, some bioethicists have started to wonder what it takes to become a genetic parent. It is widely agreed that sharing genes is not enough to substantiate the parent–offspring relation, but what is? Without a better understanding of the concept of reproduction, our thinking about parent–offspring relations and the ethical issues surrounding them risk being unprincipled. Here, I address that problem by offering a principled account of reproduction—the Overlap, Development and Persistence account—which (...) I believe best captures the meaning of ‘genetic parenthood’. (shrink)

This paper addresses the foundations of Teleological Individualism, the view that organisms, even non-sentient organisms, are goal-oriented systems while biological collectives, such as ecosystems or conspecific groups, are mere assemblages of organisms. Typical defenses of Teleological Individualism ground the teleological organization of organisms in the workings of natural selection. This paper shows that grounding teleological organization in natural selection is antithetical to Teleological Individualism because such views assume a view about the units of selection on which it is only individual (...) organisms that are units of selection. However, none of the Conventionalist, Reductionist, or Multi-Level Realist theories serve to justify such an assumption. Thus, Teleological Individualism cannot be grounded in natural selection. (shrink)

Various results show the ‘formal equivalence’ of kin and group selectionist methodologies, but this does not preclude there being a real and useful distinction between kin and group selection processes. I distinguish individual- and population-centred approaches to drawing such a distinction, and I proceed to develop the latter. On the account I advance, the differences between kin and group selection are differences of degree in the structural properties of populations. A spatial metaphor provides a useful framework for thinking about these (...) differences: kin and group selection may be conceptualized as large, overlapping regions of K - G space. I then consider some implications of the account, defend it from possible objections, and further argue that the structural features characteristic of both kin and group selection may recur at multiple levels of biological organization. (shrink)

Biological individuality is a major topic of discussion in biology and philosophy of biology. Recently, several objections have been raised against traditional accounts of biological individuality, including the objections of monism, theory-centrism, ahistoricity, disciplinary isolationism, and the multiplication of conceptual uncertainties. In this introduction, I will examine the current philosophical landscape about biological individuality, and show how the contributions gathered in this special issue address these five objections. Overall, the aim of this issue is to offer a more diverse, unifying, (...) and scientifically informed conception of what a biological individual is. (shrink)

I spell out and update the individuality thesis, that species are individuals, and not classes, sets, or kinds. I offer three complementary presentations of this thesis. First, as a way of resolving an inconsistent triad about natural kinds; second, as a phylogenetic systematics theoretical perspective; and, finally, as a novel recursive account of an evolved character. These approaches do different sorts of work, serving different interests. Presenting them together produces a taxonomy of the debates over the thesis, and isolates ways (...) it has been productive. This goes to the larger point of this paper: a defense of the individuality thesis in terms of its utility, and an update of it in light of recent theoretical developments and empirical work in biology. (shrink)

This paper provides a philosophical analysis of the Price equation and its role in evolutionary theory. Traditional models in population genetics postulate simplifying assumptions in order to make the models mathematically tractable. On the contrary, the Price equation implies a very specific way of theorizing, starting with assumptions that we think are true and then deriving from them the mathematical rules of the system. I argue that the Price equation is a generalization-sketch, whose main purpose is to provide a unifying (...) framework for researchers, helping them to develop specific models. The Price equation plays this role because, like other scientific principles, shows features as abstractness, unification and invariance. By underwriting this special role for the Price equation some recent disputes about it could be diverted. (shrink)

Clade selection is unpopular with philosophers who otherwise accept multilevel selection theory. Clades cannot reproduce, and reproduction is widely thought necessary for evolution by natural selection, especially of complex adaptations. Using microbial evolutionary processes as heuristics, I argue contrariwise, that (1) clade growth (proliferation of contained species) substitutes for clade reproduction in the evolution of complex adaptation, (2) clade-level properties favoring persistence – species richness, dispersal, divergence, and possibly intraclade cooperation – are not collapsible into species-level traits, (3) such properties (...) can be maintained by selection on clades, and (4) clade selection extends the explanatory power of the theory of evolution. (shrink)

This paper presents a modified random network model to illustrate how groups can form in the absence of evolutionary forces, assuming groups are collections of entities at any level of organization. This model is inspired by the Zero Force Evolutionary Law, which states that there is always a tendency for diversity and complexity to increase in any evolutionary system containing variation and heredity. That is, in the absence of evolutionary forces, the expectation is a continual increase in diversity and complexity (...) at any level of biological hierarchy. I show that, when modeled, this expectation of increasing variation results not only in the formation of groups, but also in a higher probability of group formation than is found in a model that is purely random. (shrink)

In this contribution, I comment on Raffaella Campaner’s defense of explanatory pluralism in psychiatry (in this volume). In her paper, Campaner focuses primarily on explanatory pluralism in contrast to explanatory reductionism. Furthermore, she distinguishes between pluralists who consider pluralism to be a temporary state on the one hand and pluralists who consider it to be a persisting state on the other hand. I suggest that it would be helpful to distinguish more than those two versions of pluralism – different understandings (...) of explanatory pluralism both within philosophy of science and psychiatry – namely moderate/temporary pluralism, anything goes pluralism, isolationist pluralism, integrative pluralism and interactive pluralism. Next, I discuss the pros and cons of these different understandings of explanatory pluralism. Finally, I raise the question of how to implement or operationalize explanatory pluralism in scientific practice; how to structure the “genuine dialogue” or shape “the pluralistic attitude” Campaner is referring to. As tentative answers, I explore a question-based framework for explanatory pluralism as well as social-epistemological procedures for interaction among competing approaches and explanations. (shrink)

Ernst Mayr’s classical work on the nature of causation in biology has had a huge influence on biologists as well as philosophers. Although his distinction between proximate and ultimate causation recently came under criticism from those who emphasize the role of development in evolutionary processes, the formal relationship between these two notions remains elusive. Using causal graph theory, this paper offers a unified framework to systematically translate a given “proximate” causal structure into an “ultimate” evolutionary response, and illustrates evolutionary implications (...) of various kinds of causal mechanisms including epigenetic inheritance, maternal effects, and niche construction. These results not only reveal the essential interplay between proximate and ultimate causation in the study of evolution, but also provide a formal method to evaluate or discover non-standard or yet unknown evolutionary phenomena. (shrink)