Constructive Neutral Evolution is an evolutionary mechanism that can explain much molecular inter-dependence and organismal complexity without assuming positive selection favoring such dependency or complexity, either directly or as a byproduct of adaptation. It differs from but complements other non-selective explanations for complexity, such as genetic drift and the Zero Force Evolutionary Law, by being ratchet-like in character. With CNE, purifying selection maintains dependencies or complexities that were neutrally evolved. Preliminary treatments use it to explain specific genetic and molecular structures (...) or processes, such as retained gene duplications, the spliceosome, and RNA editing. Here we aim to expand the scope of such explanation beyond the molecular level, integrating CNE with Multi-Level Selection theory, and arguing that several popular higher-level selection scenarios are in fact instances of CNE. Suitably contextualized, CNE occurs at any level in the biological hierarchy at which natural selection as normally construed occurs. As examples, we focus on modularity in protein–protein interaction networks or “interactomes,” the origin of eukaryotic cells and the evolution of co-dependence in microbial communities—a variant of the “Black Queen Hypothesis” which we call the “Gray Queen Hypothesis”. (shrink)

Constructive Neutral Evolution theory provides selectively neutral explanations of the origin and maintenance of biological complexity. This essay provides an analysis of CNE as an explanatory strategy defined by a tripartite set of conditions, and shows how this applies to cases of the evolution of complexity at higher-levels of the biological hierarchy. CNE was initially deployed to help explain a variety of complex molecular structures and processes, including spliceosomal splicing, trypansomal pan-editing, scrambled genes in ciliates, duplicate gene retention and fungal (...) ATP synthetase structure. CNE has also been generalized to apply to phenomena at the cellular level, including protein-protein interaction network modularity, obligate microbial symbioses, eukaryogenesis and microbial unculturability. This essay further extends CNE to cases of complexity at levels of organization higher than the molecular and cellular. These are multicellular phenotypic complexity, multicellular ecological complexity and, some cases of cultural complexity. (shrink)

In their 2010 book, Biology’s First Law, D. McShea and R. Brandon present a principle that they call ‘‘ZFEL,’’ the zero force evolutionary law. ZFEL says (roughly) that when there are no evolutionary forces acting on a population, the population’s complexity (i.e., how diverse its member organisms are) will increase. Here we develop criticisms of ZFEL and describe a different law of evolution; it says that diversity and complexity do not change when there are no evolutionary causes.

Cultural evolutionary theory has been alternatively compared to a theory of forces, such as Newtonian mechanics, or the kinetic theory of gases. In this article, I clarify the scope and significance of these metatheoretical characterisations. First, I discuss the kinetic analogy, which has been recently put forward by Tim Lewens. According to it, cultural evolutionary theory is grounded on a bottom-up methodology, which highlights the additive effects of social learning biases on the emergence of large-scale cultural phenomena. Lewens supports this (...) claim by arguing that it is a consequence of cultural evolutionists’ widespread commitment to population thinking. While I concur with Lewens that cultural evolutionists often actually conceive cultural change in aggregative terms, I think that the kinetic framework does not properly account for the explanatory import of population-level descriptions in cultural evolutionary theory. Starting from a criticism of Lewens’ interpretation of population thinking, I argue that the explanatory role of such descriptions is best understood within a dynamical framework—that is, a framework according to which cultural evolutionary theory is a theory of forces. After having spelled out the main features of this alternative interpretation, I elucidate in which respects it helps to outline a more accurate characterisation of the overarching structure of cultural evolutionary theory. (shrink)

El proyecto del ‘Darwinismo Formal’ de Alan Grafen propone una formulación matemática de la teoría de Darwin que pretende demostrar que la selección natural moldea los rasgos fenotípicos a través de la maximización de la eficacia. El proyecto de Grafen reposa sobre tres premisas: la selección natural es la única fuerza que moldea los fenotipos; la eficacia es la única medida de le evolución; y el diseño biológico surge como resultado de un proceso de optimización selectiva. En este trabajo argumentamos (...) que estas tres premisas limitan la aplicabilidad del modelo a los hechos evolutivos más simples. Esto se debe a que Grafen implícitamente presupone un concepto de eficacia que es a la vez causa y efecto de las novedades fenotípicas; lo que, por un lado, vacía la teoría de Darwin de poder explicativo y, por el otro, lleva a ignorar la potencial contribución de fuerzas evolutivas no selectivas en la configuración de la complejidad biológica. Para superar estos escollos del proyecto de del Darwinismo Formal - que son además comunes a todos los modelos formales adaptacionistas - proponemos sustituir el concepto causal de eficacia por el de robustez, definiendo así el diseño biológico como forma y función en lugar de como maximización de la eficacia. (shrink)

The assumption that natural selection alone is sufficient to explain not only which traits get fixed in a population/species, but also how they develop, has been questioned since Darwin’s times, and increasingly in the last decades. Alternative theories, linked to genetic and phenotypic processes, or to the theory of complex systems, have been proposed to explain the rise of the phenotypic variety upon which natural selection acts. In this article, we illustrate the current state of the issue and we propose (...) a logical space based on phenotypic robustness that allows a classification of evolutionary phenomena and can provide a framework for unifying all these accounts. (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)

A common belief is that evolution generally proceeds towards greater complexity at both the organismal and the genomic level, numerous examples of reductive evolution of parasites and symbionts notwithstanding. However, recent evolutionary reconstructions challenge this notion. Two notable examples are the reconstruction of the complex archaeal ancestor and the intron‐rich ancestor of eukaryotes. In both cases, evolution in most of the lineages was apparently dominated by extensive loss of genes and introns, respectively. These and many other cases of reductive evolution (...) are consistent with a general model composed of two distinct evolutionary phases: the short, explosive, innovation phase that leads to an abrupt increase in genome complexity, followed by a much longer reductive phase, which encompasses either a neutral ratchet of genetic material loss or adaptive genome streamlining. Quantitatively, the evolution of genomes appears to be dominated by reduction and simplification, punctuated by episodes of complexification. (shrink)

The aim of this contribution is to investigate certain selected parts of the extended evolutionary synthesis which all have a common denominator, namely evolution by meaning attribution. We start by arguing that living organisms can manipulate and interpret their genetic script via epigenetic modifications in a semiotic manner, that is, by meaning attribution. Genes do not build living beings to be transmitted to future generations. Genes have been shaped by evolution as a memory medium that is transmitted from one generation (...) to the next, but the actual reading of such scripts is modified by momentary contexts. Secondly, we show that phenotypic evolution variously re-uses already existing homologies which in evolving systems acquire a new meaning. We also suggest that the ways in which organisms perceive their environment and other living beings is an important but still largely neglected evolutionary force. Variations in perception influence the direction and intensity of sexual selection and some behaviourally mediated regimes of natural selection. Thirdly, we point out that especially if we want to study their evolution, living beings should not be considered in isolation but in their mutual coexistence, in their historically established being together. Recent attempts to view living beings as constructors of niches and holobionts seem compatible with the classical Umwelt theory. This approach seems capable of accounting for both competitiveness and cooperation, thus making the overall image of evolution more comprehensive. And finally, we argue that if we want to expand our understanding of biological evolution, in addition to variation, selection, and inheritance we also need to take into account processes which participate in meaning attribution. (shrink)

This paper proposes a sender-receiver model to explain two large-scale patterns observed in natural languages: Zipf’s inverse power law relating the frequency of word use and word rank, and the negative correlation between the frequency of word use and rate of lexical change. Computer simulations show that the model recreates Zipf’s inverse power law and the negative correlation between signal frequency and rate of change, provided that agents balance the rates with which they invent new signals and forget old ones. (...) Results are robust across a wide range of parameter values and structural assumptions, such as different forgetting rules and forgetting rates. Analysis of the model further suggests that Zipf’s law relating word frequency and rank arises because of language-external factors and that frequent signals change less because frequent signals are less subject to drift than rare ones. The paper concludes with some brief considerations on model-based and data-driven approaches in philosophy. (shrink)

The consensus among cultural evolutionists seems to be that human cultural evolution is cumulative, which is commonly understood in the specific sense that cultural traits, especially technological traits, increase in complexity over generations. Here we argue that there is insufficient credible evidence in favor of or against this technological complexity thesis. For one thing, the few datasets that are available hardly constitute a representative sample. For another, they substantiate very specific, and usually different versions of the complexity thesis or, even (...) worse, do not point to complexity increases. We highlight the problems our findings raise for current work in cultural-evolutionary theory, and present various suggestions for future research. (shrink)

In recent years, many philosophers of science have attempted to articulate a theory of non-epistemic emergence that is compatible with mechanistic explanation and incompatible with reductionism. The 2005 account of Fred C. Boogerd et al. has been particularly influential. They argued that a systemic property was emergent if it could not be predicted from the behaviour of less complex systems. Here, I argue that Boogerd et al.'s attempt to ground emergence in complexity guarantees that we will see emergence, but at (...) the cost of rendering it either trivial or epistemic. There are three basic problems. First, neither the measures of complexity explicitly mentioned by Boogerd et al. nor the most popular measures in the literature can do the practical and theoretical work that they assign to complexity. Second, I argue that while the success of their view depends on restricting the base of information available to the reductionist, this cannot be done in a way that is metaphysically neutral with respect to emer.. (shrink)

Biology seems to present local and transitory regularities rather than immutable laws. To account for these historically constituted regularities and to distinguish them from mathematical invariants, Montévil and Mossio (Journal of Theoretical Biology 372:179–191, 2015) have proposed to speak of constraints. In this article we analyse the causal power of these constraints in the evolution of biodiversity, i.e., their positivity, but also the modality of their action on the directions taken by evolution. We argue that to fully account for the (...) causal power of these constraints on evolution, they must be thought of in terms of normativity. In this way, we want to highlight two characteristics of the evolutionary constraints. The first, already emphasised as reported by Gould (The structure of evolutionary theory, Harvard University Press, 2002), is that these constraints are both produced by and producing biological evolution and that this circular causation creates true novelties. The second is that this specific causality, which generates unpredictability in evolution, stems not only from the historicity of biological constraints, but also from their internalisation through the practices of living beings. (shrink)

Current Perspectives in Philosophy of Biology.

The propensity account of fitness intends to solve the classical tautologicity issue by identifying fitness with a disposition, the ability to survive and reproduce. As proponents recognized early on, this account requires operational independence from actual reproductive success to avoid circularity and vacuousness charges. They suggested that operational independence is achieved by measuring fitness values through optimality models. Our goal in this article is to develop this suggestion. We show that one plausible procedure by which these independent operationalizations could be (...) thought to take place, and which is in accordance with what is said in the optimality literature, is unsound. We provide two independent lines of reasoning to show this. We then provide a sketch of a more adequate account of the role of optimality models in evolutionary contexts and draw some consequences. (shrink)

En un libro reciente McShea y Brandon defienden que la diversidad y la complejidad de la vida se explican, principalmente, por la acción de un principio que llaman “la ley evolutiva de fuerzas cero” o “ZFEL”. Tal principio actuaría de un modo implícito por detrás de muchas explicaciones de la biología, pero nunca habría sido explicitado. Asumiendo que esta idea es interesante, y que los autores en cuestión tienen razón, discutiremos el modo metateórico en que presentan dicho principio, como siendo (...) parte de la teoría de la probabilidad. Esto permite a los autores afirmar que la teoría de la probabilidad brindaría la base reductiva para toda la biología evolutiva (dado que consideran que otros principios, como el de selección natural, también serían parte de la teoría de la probabilidad). Defenderemos que, efectivamente, ZFEL no es propio de la biología únicamente, pero no por formar parte de la teoría de la probabilidad, sino por tratarse de una versión específica del principio de causa común. -/- In a recent book, McShea and Brandon argue that the observed diversity and complexity of life are explainable by a principle they call the “zero-force evolutionary law” or “ZFEL”. Although this principle would be implicit in many explanations given by biologists, it would have never been made explicit. Assuming that this idea is interesting, and that the authors are right, we will discuss the metatheoretical way in which they present said principle, as being a part of probability theory. This allows the authors to claim that probability theory provides the reductive basis for all evolutionary biology. We will defend, in accordance with them, that ZFEL is not a solely biological principle, but not because it is a part of probability theory, but rather because it is a specific version of the principle of common cause. (shrink)

This article elaborates on McShea and Brandon’s idea that drift is unlike the rest of the evolutionary factors because it is constitutive rather than imposed on the evolutionary process. I show that the way they spelled out this idea renders it inadequate and is the reason why it received some objections. I propose a different way in which their point could be understood, that rests on two general distinctions. The first is a distinction between the underlying mathematical apparatus used to (...) formulate a theory and a concept proposed by that theory. With the aid of a formal reconstruction of a population genetic model, I show that drift belongs to the first category. That is, that drift is constitutive of population genetics in the same sense that multiplication is constitutive in classical mechanics, or that circle is constitutive in Ptolemaic astronomy. The second distinction is between eliminating a concept from a theory and setting its value to zero. I will show that even though drift can be set to zero just like the rest of the evolutionary factors, eliminating drift is much harder than eliminating those other factors, since it would require changing the entire mathematical apparatus of standard population genetic theory. I conclude by drawing some other implications from the proposed formal reconstruction. (shrink)

The adequacy of Elliott Sober’s analogy between classical mechanics and evolutionary theory—according to which both theories explain via a zero-force law and a set of forces that alter the zero-force state—has been criticized from various points of view. I focus here on McShea and Brandon’s claim that drift shouldn’t be considered a force because it is not directional. I argue that there are a number of different theses that could be meant by this, and show that one of those theses—the (...) idea that drift cannot bias populations to be taken somewhere in the evolutionary space from one generation to the next—is actually false. Not only has this thesis been implicitly assumed in the discussion of the force analogy thus far, but it is also commonly found in a wider range of philosophical and biological texts. I argue that correcting this view, and the usual images associated with it, will thereby bring heuristic benefits that impact the force analogy discussion, but that also go beyond it. (shrink)

Morphological EvoDevo is a field of biological inquiry in which explicit relations between evolutionary patterns and growth or morphogenetic processes are made. Historically, morphological EvoDevo results from the coming together of several traditions, notably Naturphilosophie, embryology, the study of heterochrony, and developmental constraints. A special feature binding different approaches to morphological EvoDevo is the use of formalisms and mathematical models. Here we will introduce anatomical network analysis, a new approach centered on connectivity patterns formed by anatomical parts, with its own (...) concepts and tools specifically designed for the study of morphological EvoDevo questions. Riedl’s concept of burden is tightly related to the use of anatomical networks, providing a nexus between the evolutionary patterns and the structural constraints that shape them. (shrink)

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

Cancer is typically spoken of as a “complex” disease. But, in what sense are cancers “complex”? Is there one sense, or several? What implications does this complexity have – both for how we study, and how we intervene upon cancers? The aim of this paper is first, to clarify the variety of senses in which cancer is spoken of as "complex" in the scientific literature, and second, to discover what explanatory and predictive roles such features play.

One hotly debated philosophical question in the analysis of evolutionary theory concerns whether or not evolution and the various factors which constitute it may profitably be considered as analogous to “forces” in the traditional, Newtonian sense. Several compelling arguments assert that the force picture is incoherent, due to the peculiar nature of genetic drift. I consider two of those arguments here – that drift lacks a predictable direction, and that drift is constitutive of evolutionary systems – and show that they (...) both fail to demonstrate that a view of genetic drift as a force is untenable. I go on to diagnose the reasons for the stubborn persistence of this problem, considering two open philosophical issues and offering some preliminary arguments in support of the force metaphor. (shrink)

No comprehensive theory of development is available yet. Traditionally, we regard the development of animals as a sequence of changes through which an adult multicellular animal is produced, starting from a single cell which is usually a fertilized egg, through increasingly complex stages. However, many phenomena that would not qualify as developmental according to these criteria would nevertheless qualify as developmental in that they imply nontrivial (e.g., non degenerative) changes of form, and/or substantial changes in gene expression. A broad, comparative (...) approach is badly needed. In the Cnidaria, for example, even the boundary between generations is problematic. Describing their life cycle in terms of metagenesis (alternation between polyp generation and medusa generation) or in terms of metamorphosis (polyp as larva or juvenile) are matters of semantics more than biology. The life cycle of other metazoans, described in textbooks in terms of larva-to-adult metamorphosis, is hardly different from a typical metagenetic life cycle of cnidarians. This applies to holometabolous insects and to marine invertebrates like sea urchins, where most of the larval cells are discarded at metamorphosis. The uncertain temporal and spatial boundaries of individual development are also shown by the widespread lack of a strict correspondence between adult and mature. A comprehensive theory of development should start with a zero principle of “developmental inertia,” corresponding to an indeterminate local self-perpetuation of cell-level dynamics. Indeterminate growth, scale-invariance, segmentation, and regeneration provide examples of developmental dynamics close to that. (shrink)

Our understanding of the universe has grown rapidly in recent decades. We’ve discovered evidence of water in nearby planets, discovered planets outside our solar system, mapped the genomes of thousands of organisms, and probed the very origins and limits of life. The scientific perspective of life-as-it-could-be has expanded in part by research in astrobiology, synthetic biology, and artificial life. In the face of such scientific developments, we argue there is an ever-growing need for universal biology, life-as-it-must-be, the multidisciplinary study of (...) non-contingent aspects of life as guided by biological theory and constrained by the universe. We present three distinct but connected ways of universalizing biology—with respect to characterizing aspects of life everywhere, with respect to the explanatory scope of biological theory, and with respect to extending biological insights to the structure of nonbiological entities. For each of these, we sketch the theoretical goals and challenges, as well as give examples of current research that might be labeled universal biology. (shrink)

Due to its high degree of complexity and its historical nature, evolutionary biology has been traditionally portrayed as a messy science. According to the supporters of such a view, evolutionary biology would be unable to formulate laws and robust theories, instead just delivering coherent narratives and local models. In this article, our aim is to challenge this view by showing how the Price equation can work as the core of a general theoretical framework for evolutionary phenomena. To support this claim, (...) we outline some unnoticed structural similarities between physical theories and evolutionary biology. More specifically, we shall argue that the Price equation, in the same way as fundamental formalisms in physics, can serve as a heuristic principle to formulate and systematise different theories and models in evolutionary biology. (shrink)

In this paper, I present a new framework supporting the claim that some elements in science play a constitutive function, with the aim of overcoming some limitations of Friedman's (2001) account. More precisely, I focus on what I consider to be the gradualism implicit in Friedman's interpretation of the constitutive a priori, that is, the fact that it seems to allow for degrees of 'constitutivity'. I tease out such gradualism by showing that the constitutive character Friedman aims to track can (...) be captured by three features - namely, quasi-axiomaticity (QA), generative potential (GP), and empirical shielding (ES) - which are exhibited to a maximal degree by the examples Friedman deploys, particularly in his analysis of Newtonian mechanics. I argue that not all varieties of 'constitutivity' can be captured by the kind of gradualism implicit in Friedman's view, although developing the gradualism itself might provide useful insights. To show this, I analyse the function of the Hardy-Weinberg principle (HWP) in population genetics in terms of its QA, GP, and ES. Whereas the HWP does not count as constitutive in classical philosophical interpretations (Sober 1984), nor does it within Friedman's framework, it does nonetheless perform a minimally constitutive function. By means of historical details and considerations on the prospects of replacing the HWP, I show that the HWP is minimally constitutive by being a counterfactual instantiation of a paradigmatically constitutive stability principle, where the latter might itself be regarded as an enabling condition for a variety of modelling practices across the sciences. (shrink)

The pervasive use of computer simulations in the sciences brings novel epistemological issues discussed in the philosophy of science literature since about a decade. Evolutionary biology strongly relies on such simulations, and in relation to it there exists a research program (Artificial Life) that mainly studies simulations themselves. This paper addresses the specificity of computer simulations in evolutionary biology, in the context (described in Sect. 1) of a set of questions about their scope as explanations, the nature of validation processes (...) and the relation between simulations and true experiments or mathematical models. After making distinctions, especially between a weak use where simulations test hypotheses about the world, and a strong use where they allow one to explore sets of evolutionary dynamics not necessarily extant in our world, I argue in Sect. 2 that (weak) simulations are likely to represent in virtue of the fact that they instantiate specific features of causal processes that may be isomorphic to features of some causal processes in the world, though the latter are always intertwined with a myriad of different processes and hence unlikely to be directly manipulated and studied. I therefore argue that these simulations are merely able to provide candidate explanations for real patterns. Section 3 ends up by placing strong and weak simulations in Levins’ triangle, that conceives of simulations as devices trying to fulfil one or two among three incompatible epistemic values (precision, realism, genericity). (shrink)

Following a previous elaboration of the concept of weak individuality and some examples of its instances in ecology and biology, the article focuses on general features of the concept, arguing that in any ontological field individuals are understood on the basis of our knowledge of interactions, through the application of these general formulas for extracting individuals from interactions. Then, the specificities of the individuality in the sense of this weak concept are examined in ecology; I conclude by addressing the differences (...) between ecosystems and organisms as they appear in the viewpoint of such concept. (shrink)

In this article we focus on the emergence of biological individuality by association, trying to formulate some theoretical conditions to think about the process of collective individualization. The starting point of our analysis is the notion of “major evolutionary transition.” A major evolutionary transition is the result of the integration of a multiplicity of initially independent biological entities that, by managing to organize their interactions, become a collective of components having an identity oriented towards a common goal. When biological organisms (...) are concerned, a major transition corresponds to a phenomenon of fusion between them. We shall argue that the emergence of a new biological level of individuality implies the establishment of constitutive relationships between individuals that change their status as autonomous entities. As a result, the emergence of a new type of entity in the living world implies that individuals enter into relationships that intrinsically transform them, a transformation sufficient for a “whole” to become a “part” that forms another “whole”, that is, a new level of organismality. (shrink)

In this article, I propose a new way of thinking about natural necessity and a new way of thinking about biological laws. I suggest that much of the lack of progress in making a positive case for distinctively biological laws is that we’ve been looking for necessity in the wrong place. The trend has been to look for exceptionlessness at the level of the outcomes of biological processes and to build one’s claims about necessity off of that. However, as Beatty (...) (1995) observed, even when we are lucky enough to find a biological ‘rule’ of some sort, that rule is apt to be a victim of ‘the rule-breaking capabilities of evolutionary change’. If indeed no distinctively biological generalization—even an exceptionless one—is safe, we need to locate necessity elsewhere. A good place to start is, I think, precisely the point at which Beatty sees the possibility of lawhood as breaking down—namely, at the level of chances. 1 The ‘Necessity’ Objection to Biological Laws2 Necessary Chances2.1 Necessary chances: random drift2.2 Necessary chances: fitness3 But is it Biological?4 Conclusion. (shrink)

The concept of fitness has generated a lot of discussion in philosophy of biology. There is, however, relative agreement about the need to distinguish at least two uses of the term: ecological fitness on the one hand, and population genetics fitness on the other. The goal of this paper is to give an explication of the concept of ecological fitness by providing a reconstruction of the theory of natural selection in which this concept was framed, that is, based on the (...) way the theory was put to use in Darwin’s main texts. I will contend that this reconstruction enables us to account for the current use of the theory of natural selection. The framework presupposed in the analysis will be that of metatheoretical structuralism. This framework will provide both a better understanding of the nature of ecological fitness and a more complete reconstruction of the theory. In particular, it will provide what I think is a better way of understanding how the concept of fitness is applied through heterogeneous cases. One of the major advantages of my way of thinking about natural selection theory is that it would not have the peculiar metatheoretical status that it has in other available views. I will argue that in order to achieve these goals it is necessary to make several concepts explicit, concepts that are frequently omitted in usual reconstructions. (shrink)

Darwin begins On the Origin of Species by asking the reader to “reflect on the vast diversity of the plants and animals which have been cultivated” (1859, p. 7); almost five-hundred pages later, he closes by having the reader consider the “endless forms most beautiful and wonderful” that have evolved (1859, p. 490). Darwin contemplates diversity throughout the Origin and presents the principle of divergence as a way to explain it. Darwin formulated the principle of divergence around 1857 (Browne 1980), (...) at which point he began replacing his previous view of “limited perfect adaptedness” with a view of relative adaptedness (Ospovat 1981; Burian 1983). Proponents of perfect adaptedness, such as Paley and .. (shrink)

In this paper, I discuss the concept of complexity. I show that the principle of natural selection as acting on complexity gives a solution to the problem of reconciling the seemingly contradictory notion of generally increasing complexity and the observation that most species don’t follow such a trend. I suggest the process of evolution to be illustrated by means of a schematic diagram of complexity versus time, interpreted as a form of the Tree of Life. The suggested model implies that (...) complexity is cumulatively increasing, giving evolution a direction, an arrow of time, thus also implying that the latest emerging species will be the one with the highest level of complexity. Since the human species is the last species evolved in the evolutionary process seen at large, this means that we are the species with the highest complexity. The model implies that the human species constitutes an integral part of organic evolution, yet rendering us the exclusive status as the species of the highest complexity. (shrink)

The Hardy–Weinberg equilibrium has been argued by Sober, Stephens and others to represent the zero-force state for evolutionary biology understood as a theory of forces. I investigate what it means for a model to involve forces, developing an explicit account by defining what the zero-force state is in a general theoretical context. I use this account to show that Hardy–Weinberg equilibrium is not the zero-force state in biology even in the contexts in which it applies, and argue based on this (...) that drift should not be understood as an evolutionary force. (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)

The purpose of this paper is to defend, contra Fodor and Piattelli-Palmarini (F&PP), that the theory of natural selection (NS) is a perfectly bona fide empirical unified explanatory theory. F&PP claim there is nothing non-truistic, counterfactual-supporting, of an “adaptive” character and common to different explanations of trait evolution. In his debate with Fodor, and in other works, Sober defends NS but claims that, compared with classical mechanics (CM) and other standard theories, NS is peculiar in that its explanatory models are (...) a priori (a trait shared with few other theories). We argue that NS provides perfectly bona fide adaptive explanations of phenotype evolution, unified by a common natural-selection guiding principle. First, we introduce the debate and reply to F&PP’s main argument against NS. Then, by reviewing different examples and analyzing Fisher’s model in detail, we show that NS explanations of phenotypic evolution share a General Natural Selection Principle. Third, by elaborating an analogy with CM, we argue against F&PP’s claim that such a principle would be a mere truism and thus explanatorily useless, and against Sober’s thesis that NS models/explanations have a priori components that are not present in CM and other common empirical theories. Irrespective of differences in other respects, the NS guiding principle has the same epistemic status as other guiding principles in other highly unified theories such as CM. We argue that only by pointing to the guiding principle-driven nature that it shares with CM and other highly unified theories, something no-one has done yet in this debate, one can definitively show that NS is not defective in F&PP’s sense: in the respects relevant to the debate, Natural Selection is as defective and as epistemically peculiar as Classical Mechanics and other never questioned theories. (shrink)

The epistemic status of Natural Selection has seemed intriguing 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 NS, 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. Sober’s argument focuses on some “would promote” sentences that (...) according to him, play a central role in NS explanations and are both causal and a priori. Lange and Rosenberg criticize Sober arguing that, though there may be some unspecific a priori causal claims, there are not a priori causal claims that specify particular causal factors. Although we basically agree with Lange and Rosenberg’s criticism, we think it remains silent about a second important element in Sober’s dialectics, namely his claim that, contrary to what happens in mechanics, in NS explanatory conditionals are a priori, and that this is so in quite specific explanatory models. In this paper we criticize this second element of Sober’s argument by analyzing what we take to be the four possible interpretations of Sober’s claim, and argue that, terminological preferences aside, the possible senses in which explanatory models in NS can qualify, or include elements that can qualify, as a priori, also apply to CM and other standard, highly unified theories. We conclude that this second claim is unsound, or at least that more needs to be said in order to sustain that NS explanatory models are a priori in a sense in which CM models are not. (shrink)

The dominant view today on evolutionary progress is that it has been thoroughly debunked. Even value-neutral progress concepts are seen to lack important theoretical underpinnings: natural selection provides no rationale for progress, and natural selection need not even be invoked to explain large-scale evolutionary trends. In this paper I challenge this view by analysing how natural selection acts in heterogeneous environments. This not only undermines key debunking arguments, but also provides a selectionist rationale for a pattern of “evolutionary unfolding”, where (...) life radiates across an increased range of exploitation of environmental heterogeneity. (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)

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)

This inaugural handbook documents the distinctive research field that utilizes history and philosophy in investigation of theoretical, curricular and pedagogical issues in the teaching of science and mathematics. It is contributed to by 130 researchers from 30 countries; it provides a logically structured, fully referenced guide to the ways in which science and mathematics education is, informed by the history and philosophy of these disciplines, as well as by the philosophy of education more generally. The first handbook to cover the (...) field, it lays down a much-needed marker of progress to date and provides a platform for informed and coherent future analysis and research of the subject. -/- The publication comes at a time of heightened worldwide concern over the standard of science and mathematics education, attended by fierce debate over how best to reform curricula and enliven student engagement in the subjects There is a growing recognition among educators and policy makers that the learning of science must dovetail with learning about science; this handbook is uniquely positioned as a locus for the discussion. -/- The handbook features sections on pedagogical, theoretical, national, and biographical research, setting the literature of each tradition in its historical context. Each chapter engages in an assessment of the strengths and weakness of the research addressed, and suggests potentially fruitful avenues of future research. A key element of the handbook’s broader analytical framework is its identification and examination of unnoticed philosophical assumptions in science and mathematics research. It reminds readers at a crucial juncture that there has been a long and rich tradition of historical and philosophical engagements with science and mathematics teaching, and that lessons can be learnt from these engagements for the resolution of current theoretical, curricular and pedagogical questions that face teachers and administrators. (shrink)

This article analyzes the view of evolutionary theory as a theory of forces. The analogy with Newtonian mechanics has been challenged due to the alleged mismatch between drift and the other evolutionary forces. Since genetic drift has no direction several authors tried to protect its status as a force: denying its lack of directionality, extending the notion of force and looking for a force in physics which also lacks of direction. I analyse these approaches, and although this strategy finally succeeds, (...) this discussion overlooks the crucial point on the debate between causalists and statisticalists: the causal status of evolutionary theory. (shrink)