The notion that natural selection is a process of fitness maximization gets a bad press in population genetics, yet in other areas of biology the view that organisms behave as if attempting to maximize their fitness remains widespread. Here I critically appraise the prospects for reconciliation. I first distinguish four varieties of fitness maximization. I then examine two recent developments that may appear to vindicate at least one of these varieties. The first is the ‘new’ interpretation of Fisher's (...) fundamental theorem of natural selection, on which the theorem is exactly true for any evolving population that satisfies some minimal assumptions. The second is the Formal Darwinism project, which forges links between gene frequency change and optimal strategy choice. In both cases, I argue that the results fail to establish a biologically significant maximization principle. I conclude that it may be a mistake to look for universal maximization principles justified by theory alone. A more promising approach may be to find maximization principles that apply conditionally and to show that the conditions were satisfied in the evolution of particular traits. (shrink)

In this paper, I am clarifying and defending my argument in favor of the claim that cumulative selection can explain adaptation provided that the environmental resources are limited. Further, elaborate on what this limitation of environmental resources means and why it is relevant for the explanatory power of natural selection.

Darwin provided us with a powerful theoretical framework to explain the evolution of living systems. Natural selection alone, however, has sometimes been seen as insufficient to explain the emergence of new levels of selection. The problem is one of “circularity” for evolutionary explanations: how to explain the origins of Darwinian properties without already invoking their presence at the level they emerge. That is, how does evolution by natural selection commence in the first place? Recent results (...) in experimental evolution suggest a way forward: Paul Rainey and his collaborators have shown that Darwinian properties could be exogenously imposed via what they call “ecological scaffolding.” This could solve the “black box” dilemma faced by Darwinian explanations of new levels of organization. Yet, despite “scaffolding” recently becoming a popular term in the study of cognition, culture, and evolution, the concept has suffered from vagueness and ambiguity. This article aims to show that scaffolding can be turned into a proper scientific concept able to do explanatory work within the context of the major evolutionary transitions. Doing so will allow us to once again extend the scope of the Darwinian model of evolution by natural selection. (shrink)

In their book What Darwin Got Wrong, Jerry Fodor and Massimo Piattelli-Palmarini construct an a priori philosophical argument and an empirical biological argument. The biological argument aims to show that natural selection is much less important in the evolutionary process than many biologists maintain. The a priori argument begins with the claim that there cannot be selection for one but not the other of two traits that are perfectly correlated in a population; it concludes that there cannot (...) be an evolutionary theory of adaptation. This article focuses mainly on the a priori argument. (shrink)

Many have argued that there is no reason why natural selection should cause directional increases in measures such as body size or complexity across evolutionary history as a whole. In this paper I argue that this conclusion does not hold for selection for adaptations to environmental variability, and that, given the inevitability of environmental variability, trends in adaptations to variability are an expected feature of evolution by natural selection. As a concrete instance of this causal (...) structure, I outline how this may be applied to a trend in phenotypic plasticity. (shrink)

Skipper and Millstein analyze natural selection and mechanism, concluding that natural selection is not a mechanism in the sense of the new mechanistic philosophy. Barros disagrees and provides his own account of natural selection as a mechanism. This discussion identifies a missing piece of Barros's account, attempts to fill in that piece, and reconsiders the revised account. Two principal objections are developed: one, the account does not characterize natural selection; two, the account (...) is not mechanistic. Extensive and persistent variability causes both of these difficulties, so further attempts to describe natural selection as a mechanism are also unlikely to succeed. (shrink)

Natural selection [Darwin 1859] is perhaps the most important component of evolutionary theory, since it is the only known process that can bring about the adaptation of living organisms to their environments [Gould 2002]. And yet, its study is conceptually and methodologically complex, and much attention needs to be paid to a variety of phenomena that can limit the efficacy of selection [Antonovics 1976; Pigliucci and Kaplan 2000]. In this essay, I will use examples of recent work (...) carried out in my laboratory to illustrate basic research on natural selection as conducted using a variety of approaches, including field work, laboratory experiments, and molecular genetics. I also discuss the application of this array of tools to questions pertinent to conservation biology, and in particular to the all-important problem of what makes invasive species so good at creating the sort of problems they are infamous for [Lee 2002]. (shrink)

Life on Earth descends from a common ancestor. However, it is likely that there are other instances of life in the universe. If so, each abiogenesis event will have given rise to an independently originated life clade, of which Earth-life is an example. In this paper, I argue that the set of all IOLCs in the universe forms a Darwinian population subject to natural selection, with more widely dispersed IOLCs being less likely to face extinction. As a result, (...) we should expect that, over time, more planets will become inhabited by fewer, more successful IOLCs. (shrink)

True beliefs are better guides to the world than false ones. This is the common-sense assumption that undergirds theorizing in evolutionary epistemology. According to Alvin Plantinga, however, evolution by natural selection does not care about truth: it cares only about fitness. If our cognitive faculties are the products of blind evolution, we have no reason to trust them, anytime or anywhere. Evolutionary naturalism, consequently, is a self-defeating position. Following up on earlier objections, we uncover three additional flaws in (...) Plantinga's latest formulation of his argument: a failure to appreciate adaptive path dependency, an incoherent conception of content ascription, and a conflation of common-sense and scientific beliefs, which we diagnose as the ‘foundationalist fallacy’. More fundamentally, Plantinga's reductive formalism with respect to the issue of cognitive reliability is inadequate to deal with relevant empirical details. (shrink)

Natural selection is one of the fundamental keys of evolutionary biology, which is as much as saying of almost all life sciences. However, the effort to assess its true meaning has been involved in endless controversies almost from the beginning. The rethinking of natural selection through an abstract scheme with three ingredients - −population, environment and interaction between them− could significantly contribute to clarify this debate.

An influential argument due to Elliott Sober, subsequently strengthened by Denis Walsh and Joel Pust, moves from plausible premises to the bold conclusion that natural selection cannot explain the traits of individual organisms. If the argument were sound, the explanatory scope of selection would depend, surprisingly, on metaphysical considerations concerning origin essentialism. I show that the Sober-Walsh-Pust argument rests on a flawed counterfactual criterion for explanatory relevance. I further show that a more defensible criterion for explanatory relevance (...) recently proposed by Michael Strevens lends support to the view that natural selection can be relevant to the explanation of individual traits. (shrink)

David Hume's famous riddle of induction implies a second problem related to the question of whether the laws and principles of nature might change in the course of time. Claims have been made that modern developments in physics and astrophysics corroborate the translational invariance of the laws of physics in time. However, the appearance of a new general principle of nature, which might not be derivable from the known laws of physics, or that might actually be a non-physical one (this (...) means completely independent of physical science) supports the notion that the course of nature can change in time. Here it is argued that natural selection satisfies the criteria that identify a general principle of nature which so far, appears to be non-derivable from the known laws of physics and therefore, it is likely that it arose in the course of time, thus leaving open again the quest for a true solution to Hume's second problem. (shrink)

Accounts of the concepts of function and dysfunction have not adequately explained what factors determine the line between low‐normal function and dysfunction. I call the challenge of doing so the line‐drawing problem. Previous approaches emphasize facts involving the action of natural selection (Wakefield 1992a, 1999a, 1999b) or the statistical distribution of levels of functioning in the current population (Boorse 1977, 1997). I point out limitations of these two approaches and present a solution to the line‐drawing problem that builds (...) on the second one. (shrink)

In evolutionary biology and ecology, ontological and epistemological perspectives based on the replicator and the interactor have become the background that makes it possible to transcend traditional biological levels of organization and to achieve a unified view of evolution in which replication and interaction are fundamental operating processes. Using the transactional perspective proposed originally by John Dewey and Arthur Fisher Bentley, a new ontological and methodological category is proposed here: the transactor. The transactional perspective, based on the concept of the (...) transactor, bridges the dichotomy between organisms and environment that characterizes the interactional perspective on evolution and provides epistemological support for the emergentist, systemic view of evolutionary and developmental processes. (shrink)

I defend a radical interpretation of biological populations—what I call population pluralism—which holds that there are many ways that a particular grouping of individuals can be related such that the grouping satisfies the conditions necessary for those individuals to evolve together. More constraining accounts of biological populations face empirical counter-examples and conceptual difficulties. One of the most intuitive and frequently employed conditions, causal connectivity—itself beset with numerous difficulties—is best construed by considering the relevant causal relations as ‘thick’ causal concepts. I (...) argue that the fine-grained causal relations that could constitute membership in a biological population are huge in number and many are manifested by degree, and thus we can construe population membership as being defined by massively multidimensional constructs, the differences between which are largely arbitrary. I end by showing that positions in two recent debates in theoretical biology depend on a view of biological populations at odds with the pluralism defended here. (shrink)

We argue that the fragility of contemporary marriages—and the corresponding high rates of divorce—can be explained (in large part) by a three-part mismatch: between our relationship values, our evolved psychobiological natures, and our modern social, physical, and technological environment. “Love drugs” could help address this mismatch by boosting our psychobiologies while keeping our values and our environment intact. While individual couples should be free to use pharmacological interventions to sustain and improve their romantic connection, we suggest that they may have (...) an obligation to do so as well, in certain cases. Specifically, we argue that couples with offspring may have a special responsibility to enhance their relationships for the sake of their children. We outline an evolutionarily informed research program for identifying promising biomedical enhancements of love and commitment. (shrink)

Life is supported by a myriad of chemical reactions. To describe the overall process we have formulated entropy for an open system undergoing chemical reactions. The entropy formula allows us to recognize various ways for the system to move towards more probable states. These correspond to the basic processes of life i.e. proliferation, differentiation, expansion, energy intake, adaptation and maturation. We propose that the rate of entropy production by various mechanisms is the fitness criterion of natural selection. The (...) quest for more probable states results in organization of matter in functional hierarchies. (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)

The modern concept of ideology was established by the liberal politician and philosopher Destutt de Tracy, with the objective of creating an all-embracing and general science of ideas, which followed the sensualist and empiricist trend initiated by Locke that culminated in the positivism of Comte. Natural selection and immunity are two key concepts in the history of biology that were strongly based on the Malthusian concept of struggle for existence. This concept wrongly assumed that population grew faster than (...) the means of existence. This “natural” law contained implicitly the idea that the poor and least gifted would not survive. This idea led to the progressive development of the concept of natural selection, whose definitive version was given by Darwin. Mechnikov took the concepts of struggle for existence and natural selection and conceived infectious diseases as a struggle between a host and its invader, the so-called phagocytosis theory. This theory created the necessity to possess mechanisms to discriminate between the own and the foreign, and led to the conception of the immune self. These concepts were not developed from ideas coming from perceptions or sensations, but from ideas coming from their values: individual interest, inevitable inequality, property, utility and profit. Values are ideals that constitute an ideological matrix which exerts a numinous activity and influence the development of our future actions. In consequence, science and its practice cannot avoid and ignore the values that drive them and impulse them towards certain directions. (shrink)

How do you put both physicists and biologists on their guard? Answer: propound a philosophical theory that ignores Darwin's demolition of essentialism in species and brands any physicist who denies your theory of natural kinds as an anti-realist. A traditional division in philosophy is between metaphysics (what sorts of things exist) and epistemology (what and how we know). Some think that the core of realism is the metaphysical assumption that there is a world independent of our minds. But this (...) core assumption is sometimes clothed in other assumptions, such as theories of truth, truth-likeness, meaning and knowledge. Scornful of what they see as an unnecessary retreat from a fully clothed realism to the naked postulate of a mind-independent reality, Harre, Aronson and Way present a realism that also embraces truth and truth-likeness, as well as their own conception of scientific method and the structure of the world. I argue that this approach, while intriguing and worth reading, leads to some insuperable difficulties. (shrink)

I argue that verbal models should be included in a philosophical account of the scientific practice of modelling. Weisberg (2013) has directly opposed this thesis on the grounds that verbal structures, if they are used in science, only merely describe models. I look at examples from Darwin's On the Origin of Species (1859) of verbally constructed narratives that I claim model the general phenomenon of evolution by natural selection. In each of the cases I look at, a particular (...) scenario is described that involves at least some fictitious elements but represents the salient causal components of natural selection. I pronounce the importance of prioritising observation of scientific practice for the philosophy of modelling and I suggest that there are other likely model types that are excluded from philosophical accounts. (shrink)

When natural selection theory was presented, much active philosophical debate, in which Darwin himself participated, centered on its hypothetical nature, its explanatory power, and Darwin's methodology. Upon first examination, Darwin's support of his theory seems to consist of a set of claims pertaining to various aspects of explanatory success. I analyze the support of his method and theory given in the Origin of Species and private correspondence, and conclude that an interpretation focusing on the explanatory strengths of (...) class='Hi'>natural selection theory accurately reflects neither Darwin's own self-consciously held views, nor the nature of his support. Darwin's methodological and philosophical arguments were at once consistently empiricist and more sophisticated than such interpretations credit to him. (shrink)

In many of his writings and lectures Ortega y Gasset criticized Darwinism on numbers of issues. In this paper I aim to reexamine his critique and to prove that in 1916 the harsh critique was designed to hide the similarity between his ideas and Darwin's idea of Natural Selection. The origins of Ortega's idea on the vocation of man can be traced in his dialogue with Darwin in the year 1916. In the historiography his Philosophy of Life is (...) conceived as a metaphysics that stands beyond biological and any other natural explanation. My contention on the other hand is that Ortega's concept of the vocation of man is an elaboration of Darwin's Natural Selection. I will try to show that in 1916 Ortega conceived the human character and the vocation of man as a product of Natural Selection. Instead of following Ortega's critiques on Darwin and his effort to show that there is no connection between his Philosophy of Life and Darwinism the article proposes to reconsider Ortega's thought as a constant dialogue with Darwin's theory of Natural Selection. The Change in Ortega's conception of Darwin's evolution came rather late in his thought and in the first section we will see how Ortega described technology in a manner that allows the human subject to change the environment in his favor. (shrink)

RESUMEN: El tema de este trabajo es la reconstrucción de la teoría de la selección natural darwiniana. Me propongo esbozar la ley fundamental de esta teoría de manera informal a partir de sus aplicaciones en El origen de las especies de Darwin y presentar sus conceptos fundamentales. Presentaré la red teórica de leyes especiales que surgen de la especialización de esta ley fundamental. Supondré el estructuralismo como marco metateórico. Señalaré también algunas consecuencias que mi propuesta tiene sobre ciertas discusiones (...) metateóricas en torno a la teoría y, finalmente, relacionaré mi reconstrucción con otras reconstrucciones disponibles.ABSTRACT: This paper is about the reconstruction of the Darwinian Theory of Natural Selection. My aim here is to outline the fundamental law of this theory in an informal way from its applications in The Origin of Species and to make explicit its fundamental concepts. I will introduce the theory-nets of special laws that arisefrom the specialization of the fundamental law. I will assume the metatheoretical structuralist frame. I will also point out many consequences that my proposal has about a few metatheoretical discussions around the theory and, finally, I will relate my propose to other reconstructions available. (shrink)

To understand Darwin’s concept of natural selection, we have to contrast it with his characterization of artificial selection, and then ask: what is natural in natural selection? While we do this, we develop two distinctions: one between ‘change by transformative action’ and ‘change by selection’, and another between ‘artificial selection’ and ‘natural selection’. The first distinction helps us understand evolution by selection and the second natural selection.

Neither Aristotle nor Aquinas assumes the reality of the evolution of species. Their systems of thought, however, remain open to the new data, offering an essential contribution to the ongoing debate between scientific, philosophical, and theological aspects of the theory of evolution. After discussing some key issues of substance metaphysics in its encounter with the theory of evolution (hylomorphism, transformism of species, teleology, chance, the principle of proportionate causation), I present a Thomistic response to its major hypotheses. Concerning the philosophy (...) of Aquinas, I trace what might be seen as a preliminary description of natural selection in his commentary on Aristotle's Physics. Turning toward theology, besides addressing the topics that were referred to in the past—such as: Aquinas’ reading of Genesis, his account of creation as dependence in being, secondary and instrumental causality, and univocal/equivocal predication of God—I bring into discussion Thomas’ concept of the perfection of the universe, which has been virtually unused in this context. (shrink)

Ever since the publication of Richard Dawkins' The Selfish Gene, a book for the lay reader that popularized the ideas of influential evolutionary biologists like William Hamilton and George Williams, there has been much discussion of so-called "universal Darwinism". Dawkins' dual aim was to reduce evolutionary phenomena to the level of the gene, while at the same time abstracting the Darwinian process of natural selection of "replicators" and making it into something that would apply beyond the domain of (...) biology. (shrink)

Recent discussion of mechanism has suggested new approaches to several issues in the philosophy of science, including theory structure, causal explanation, and reductionism. Here, I apply what I take to be the fruits of the 'new mechanical philosophy' to an analysis of a contemporary debate in evolutionary biology about the role of natural selection in speciation. Traditional accounts of that debate focus on the geographic context of genetic divergence--namely, whether divergence in the absence of geographic isolation is possible (...) (or significant). Those accounts are at best incomplete, I argue, because they ignore the mechanisms producing divergence and miss what is at stake in the biological debate. I argue that the biological debate instead concerns the scope of particular speciation mechanisms which assign different roles to natural selection at various stages of divergence. The upshot is a new interpretation of the crux of that debate--namely, whether divergence with gene flow is possible (or significant) and whether the isolating mechanisms producing it are adaptive. (shrink)

The eugenics movement was not the anomaly of just one country. In its day, it enamoured industrialized nations throughout the Western world. In the end, the eugenics movement ultimately did not recover from the stigma it sustained as a result of the Second World War. However, with the advancement of genetic engineering and the researches into embryonic stem cells, discussions about eugenics are becoming relevant once more, and it will be the responsibility of the informed (and not merely reactionary) to (...) cultivate reasoned argument about eugenics based on a sound knowledge of biology and ethics. The potential to rid humanity of so many hereditary ailments may rest on such a rational dialogue and, as such, we should hope to heed the lessons of history more carefully than perhaps we have. (shrink)

This is a more detailed version of my "On 'Cuteness'", which appeared in the British Journal of Aesthetics in April 1992. For John Morreall, cuteness is an abstract general attribute of infants that causes adults to want to care for them (or which is the reason, or at least important reason, for such solicitousness). I shall try to show, in what follows, that this is, if not an altogether fallacious way of explaining the matter, at least an extremely misleading one. (...) As it stands, in particular, it is too easy to infer from Morreall's line of reasoning 1) that infants in general might conceivably never have developed cuteness, and 2) that infants, because of this deficiency, would then not be cared for as adequately by their parents. An equally wrong further implication, which further helps to express my difficulty with Morreall's formulation of the matter, would be that if baby spiders (for example) had happened to have the abstract general characteristic called 'cuteness', while human children did not have it, then human adults would have been more inclined to care for baby spiders than for baby humans. It is to avoid such oddities as these that, it seems to me, a further consideration of the problem is warranted. (shrink)

As the principle of natural selection is generalized to explain (adaptive) patterns of human behavior, it becomes less clear what the selective environment empirically refers to. While the environment and individual are relatively separable in the non-human biological context, they are highly entangled in the context of moral, social, and institutional evolution. This chapter brings attention to the problem of generalizing the selective environment, and argues that it is ontologically disunified and definable only through its explanatory function. What (...) unifies the selective environment is that it explains adaptation in a non-agential way, by screening off various forms of agency, whether divine, organismic, or human. This explanatory function of the selective environment helps avoid some sources of confusion when the theory of natural selection is applied to humanities and social sciences. (shrink)

The Natural Problems, attributed to Aristotle, have gained much scholarly attention in the last decades, yet a systematic study of how the collection circulated in the Graeco-Roman Empire remains a blind spot in contemporary scholarship. Indeed, the Imperial Era is a seminal period for the history of the text, not just as a conduit between Aristotle and the Middle Ages – which in itself is essential for explaining the subsequent Arabic and Latin uptake of the Problems more clearly – (...) but also for the wealth of sources and testimonies it offers about the collection’s ancient readership and concrete use. The evidence shows that the collection sparked much debate among a range of ancient philosophers, doctors, sophists and scholars, both Greeks and Romans. This paper provides a selection of readings representative of the different socio-intellectual milieus in which the Problems circulated and the different agendas that it served. (shrink)

This brief paper summarizes a series of postal investigations on the acceptance of selection for X or Y spermatozoa. These were conducted mainly in Germany but also in the UK, the Netherlands and the US. Selected families were approached with a series of questions about their wish to use sperm selection, and their choice of boys or girls. In general, large majorities opposed this approach for family balancing or sex selection on the basis of cost and inconvenience (...) of the treatment. The view was expressed that its use in medicine was more acceptable. Care was taken to compose a series of open questions to particular sections of the population, e.g.men versus women, differing ages, and families with imbalanced sex ratios. More favourable responses arose, for example in pregnant women. Overall, all four nations had large majorities against the use of sex selection. (shrink)

It is an ongoing controversy whether natural selection is a cause of population change, or a mere statistical description of how individual births and deaths accumulate. In this paper I restate the problem in terms of the reference class problem, and propose how the structure of stable equilibrium can provide a solution in continuity with biological practice. Insofar natural selection can be understood as a tendency towards equilibrium, key statisticalist criticisms are avoided. Further, in a modification (...) of the Newtonian-force analogy, it can be suggested that a better metaphor for natural selection is that of an emergent force, similar in nature to entropic forces: with magnitude and direction, but lacking a spatiotemporal origin or point of application. (shrink)

A common misunderstanding of the selected effects theory of function is that natural selection operating over an evolutionary time scale is the only functionbestowing process in the natural world. This construal of the selected effects theory conflicts with the existence and ubiquity of neurobiological functions that are evolutionary novel, such as structures underlying reading ability. This conflict has suggested to some that, while the selected effects theory may be relevant to some areas of evolutionary biology, its relevance (...) to neuroscience is marginal. This line of reasoning, however, neglects the fact that synapses, entire neurons, and potentially groups of neurons can undergo a type of selection analogous to natural selection operating over an evolutionary time scale. In the following, I argue that neural selection should be construed, by the selected effect theorist, as a distinct type of function-bestowing process in addition to natural selection. After explicating a generalized selected effects theory of function and distinguishing it from similar attempts to extend the selected effects theory, I do four things. First, I show how it allows one to identify neural selection as a distinct function-bestowing process, in contrast to other forms of neural structure formation such as neural construction. Second, I defend the view from one major criticism, and in so doing I clarify the content of the view. Third, I examine drug addiction to show the potential relevance of neural selection to neuroscientific and psychological research. Finally, I endorse a modest pluralism of function concepts within biology. (shrink)

This paper critically reviews and characterizes the student's causal-explanatory understanding; this is done as a step toward explicating the problematic of evolution education as it concerns the cognitive difficulties in understanding Darwin's theory of natural selection. The review concludes that the student's understanding is fundamentally different from Darwin's, for the student understands evolutionary change as necessary individual transformation caused by the transformative action of various physical and behavioral factors. This is in complete contrast to Darwin's (and even the (...) Darwinian's, for that matter) understanding of evolutionary change as a change caused by accumulative selection. Hence, to understand natural selection, the student has to learn to “see” how the accumulative selection causes evolutionary change. (shrink)

A long-running dispute concerns which adaptation-related explananda natural selection can be said to explain. At issue are explananda of the form: why a given individual organism has a given adaptation rather than that same individual having another trait. It is broadly agreed that one must be ready to back up a “no” answer with an appropriate theory of trans-world identity for individuals. I argue, against the conventional wisdom, that the same is true for a “yes” answer. My conclusion (...) recasts the landscape and opens the door to a potential resolution. (shrink)

I defend a theory of mental representation that satisfies naturalistic constraints. Briefly, we begin by distinguishing (i) what makes something a representation from (ii) given that a thing is a representation, what determines what it represents. Representations are states of biological organisms, so we should expect a unified theoretical framework for explaining both what it is to be a representation as well as what it is to be a heart or a kidney. I follow Millikan in explaining (i) in terms (...) of teleofunction, explicated in terms of natural selection. -/- To explain (ii), we begin by recognizing that representational states do not have content, that is, they are neither true nor false except insofar as they both “point to” or “refer” to something, as well as “say” something regarding whatever it is they are about. To distinguish veridical from false representations, there must be a way for these separate aspects to come apart; hence, we explain (ii) by providing independent theories of what I call f-reference and f-predication (the ‘f’ simply connotes ‘fundamental’, to distinguish these things from their natural language counterparts). -/- Causal theories of representation typically founder on error, or on what Fodor has called the disjunction problem. Resemblance or isomorphism theories typically founder on what I’ve called the non-uniqueness problem, which is that isomorphisms and resemblance are practically unconstrained and so representational content cannot be uniquely determined. These traditional problems provide the motivation for my theory, the structural preservation theory, as follows. F-reference, like reference, is a specific, asymmetric relation, as is causation. F-predication, like predication, is a non-specific relation, as predicates typically apply to many things, just as many relational systems can be isomorphic to any given relational system. Putting these observations together, a promising strategy is to explain f-reference via causal history and f-predication via something like isomorphism between relational systems. -/- This dissertation should be conceptualized as having three parts. After motivating and characterizing the problem in chapter 1, the first part is the negative project, where I review and critique Dretske’s, Fodor’s, and Millikan’s theories in chapters 2-4. Second, I construct my theory about the nature of representation in chapter 5 and defend it from objections in chapter 6. In chapters 7-8, which constitute the third and final part, I address the question of how representation is implemented in biological systems. In chapter 7 I argue that single-cell intracortical recordings taken from awake Macaque monkeys performing a cognitive task provide empirical evidence for structural preservation theory, and in chapter 8 I use the empirical results to illustrate, clarify, and refine the theory. (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)

Darwin hypothesized that some animals, when selecting sexual partners, possess a genuine “sense of beauty” that cannot be accounted for by the logic of natural selection. This hypothesis has been notoriously controversial. In this chapter I propose that the concept of agency can be useful to operationalize the “sense of beauty”, and can help identify the conditions under which one can infer that animals are acting as (aesthetic) agents. Focusing on a case study of the behavior of the (...) Pavo cristatus, I identify the types of evidence that would allow for the inference of agency through aesthetic choice. (shrink)

The epistemic status of Natural Selection has intrigued to biologists and philosophers since the very beginning of the theory to our present times. One prominent contemporary example is Elliott Sober, who claims that Natural Selection, and some other theories in biology, and maybe in economics, are peculiar in including explanatory models/conditionals that are a priori in a sense in which explanatory models/conditionals in Classical Mechanics and most other standard theories are not. In this paper, by analyzing (...) what we take to be the four possible interpretations of Sober’s claim, we argue that, terminological preferences aside, the possible senses in which explanatory models in Natural Selection can qualify, or include elements that can qualify, as a priori, also apply to Classical Mechanics and other standard, highly unified theories. (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)

Darwin’s claim that Natural Selection, through optimization of fitness, explains complex biological design has not yet been properly formalized. Alan Grafen’s Formal Darwinism Project aims at providing such a formalization and at demonstrating that fitness maximization is coherent with results from Population Genetics, usually interpreted as denying it. We suggest that Grafen’s proposal suffers from some limitations linked to its concept of design as optimized fitness. In order to overcome these limitations, we propose a classification of evolutionary facts (...) based on a bi-dimensional complexity space, which adds robustness to fitness as measure of the quality of a design. In this space, each point represents an organismic architecture, and movement between two points would be an evolutionary fact. Natural Selection explains movement along the fitness axis, while non-selective mechanisms explain movement on the robustness axis. Moreover, we propose a thermodynamic metaphor to draw parallelisms between notions of fitness and entropy, robustness and energy, and movement in this space and reversible and irreversible cycles. We argue that this metaphor illustrates different evolutionary processes usually overlooked by proposals of formalization of Darwinian theory, such Grafen’s Formal Darwinism Project. (shrink)

Environmental heterogeneity is invoked as a key explanatory factor in the adaptive evolution of a surprisingly wide range of phenomena. This article aims to analyze this explanatory scheme of categorizing traits or properties as adaptations to environmental heterogeneity. First it is suggested that this scheme can be understood as a reaction to how heterogeneity adaptations were discounted or ignored in the modern synthesis. Then a positive account is proposed, distinguishing between two broad categories of adaptation to environmental heterogeneity: properties selected (...) for by well-defined patterns of environmental heterogeneity, and properties that help organisms exploit novel patterns of environmental heterogeneity. (shrink)

Recently, the relationship between evolutionary ecology and perceptual science has received renewed attention under perception-mediated selection, a mode of natural selection linking perceptual saliency, rather than veridicality, to fitness. The Interface Theory of Perception (ITP) has been especially prominent in claiming that an organism’s perceptual interface is populated by icons, which arise as a function of evolved, species-specific perceptual interfaces that produce approximations of organisms’ environments through fitness-tuned perceptions. According to perception-mediated selection, perception and behavior calibrate (...) one another as organisms’ capacities to experience and know the objects and properties of their environments lead to responses highlighting certain environmental features selected for survival. We argue this occurs via the Umwelt/Umgebung distinction in ethology, demonstrating that organisms interact with their external environments (Umgebung) through constructed perceptual schema (Umwelt) that produce constrained representations of environmental objects and their properties. Following Peircean semiotics, we claim that ITP’s focus on icons as saliency-simplified markers corresponds to biosemiotics’ understanding of perceptual representations, which manifest as iconic (resembling objects), indexical (referring), or symbolic (arbitrary) modalities, which provide for organisms’ semiotic scaffolding. We argue that ITP provides the computational evidence for biosemiotics’ notion of iconicity, while biosemiotics provides explanation within ITP for how iconicity can build up into indices and symbols. The common contention of these separate frameworks that the process of perception tracks saliency rather than veridicality suggests that digital/dyadic perceptual strategies will be outcompeted by their semiotic/triadic counterparts. This carries implications for evolutionary theory as well as theories of cognition. (shrink)

"Modern History" versions of the etiological theory claim that in order for a trait X to have the proper function F, individuals with X must have been recently favored by natural selection for doing F (Godfrey-Smith 1994; Griffiths 1992, 1993). For many traits with prototypical proper functions, however, such recent selection may not have occurred: traits may have been maintained due to lack of variation or due to selection for other effects. I examine this flaw in (...) Modern History accounts and offer an alternative etiological theory, the Continuing Usefulness account, which appears to avoid such problems. (shrink)

Review of Javier Suárez and Elisabeth A. Lloyd, _Units of Selection_. Cambridge: Cambridge University Press, 2023, 75 pp, £49.99 HB.

Resumen -/- El objetivo de este trabajo consiste en analizar las relaciones entre los modelos de optimalidad y la selección natural. Defenderemos que esas relaciones pueden dividirse en dos tipos, en tanto hay dos tipos de explicaciones seleccionistas, que llamaremos “históricas” y “ahistóricas”. Las explicaciones históricas revelan como una población dada adquiere un rasgo que es adaptativo en ese ambiente e involucran muchas generaciones, variación, etc. Las explicaciones ahistóricas, explican por qué, en determinado momento, ciertos tipos de organismos tienen (...) un mayor éxito reproductivo que otros. Mostraremos que los modelos de optimalidad pueden jugar un rol en la determinación del explanandum de las explicaciones histórica seleccionistas, esto es, que ayudan a reconocer qué rasgos son adaptativos. Por otra parte, mostramos que los modelos de optimalidad nos permiten determinar a veces la parte del explanans de las explicaciones ahistóricas (particularmente, el concepto de fitness). -/- Abstract -/- The goal of this paper is to analyze the relations between optimality models and natural selection. We contend that these relationships can be divided into two kinds, as there are two kinds of natural selection explanations, which we call “historical” and “ahistorical”. Historical explanations reveal how a given population acquires a trait which is adaptive in its environment, and involves many generations, variations, etc. Ahistorical ones, explain why, at a given moment, certain kinds of organisms have a greater reproductive success than others. We show that optimality models can play a role in determining the explanandum of historical selectionist explanations, that is, they help us to recognize which traits are adaptive. And, on the other hand, that optimality models sometimes allow us to determine part of the explanans of ahistorical explanations (particularly, the concept of fitness). (shrink)

The selected effects or ‘etiological’ theory of Proper function is a naturalistic and realist account of biological teleology. It is used to analyse normativity in philosophy of language, philosophy of mind, philosophy of medicine and elsewhere. The theory has been developed with a simple and intuitive view of natural selection. Traits are selected because of their positive effects on the fitness of the organisms that have them. These ‘selected effects’ are the Proper functions of the traits. Proponents argue (...) that this analysis of biological teleology has the unique advantage that the selected effect function of a trait is also a causal explanation of the trait: the trait exists because it performs this function. We show, however, that selected effect functions as currently defined explain the existence of traits only under highly restrictive assumptions about evolutionary dynamics. In many common scenarios in which traits evolve by natural selection, selected effect functions do not explain those traits. This is because definitions of selected effect function extract from any evolutionary scenario only the information that would be explanatorily relevant in the simple evolutionary scenario implicit in those definitions. When applied to more complex scenarios selected effect functions omit the key information that is explanatorily relevant in those scenarios. The assumptions required for selected effect functions to be explanatory are particularly unlikely to hold in the domain that its proponents care most about - the evolution of representation. A more adequate selected effects theory of Proper functions may be possible, but will require much greater attention to the structure of actual evolutionary explanations. (shrink)