Pierrick Bourrat

This paper investigates the concept of reproduction in an evolutionary context. It draws a distinction between objects that are reproduced (reproducees), objects that reproduce thanks to some reproductive autonomy (reproducers), and Darwinian individuals that are reproducers with a high degree of reproductive causal control. This threefold distinction is then applied to different biological objects classically invoked in reproduction processes (e.g., genes, viruses, cells) to explain why they do not have the same status with respect to reproduction. The distinction also provides some fuel for the view proposed by Griesemer: that material overlap during reproduction is a condition for reproduction.

In this critical notice to Robert Wright’s The Evolution of God, we focus on the question of whether Wright’s God is one which can be said to be an adaptation in a well defined sense. Thus we evaluate the likelihood of different models of adaptive evolution of cultural ideas in their different levels of selection. Our result is an emphasis on the plurality of mechanisms that may lead to adaptation. By way of conclusion we assess epistemologically some of Wright’s more controversial claims concerning the directionality of evolution and moral progress.

I argue that Calcott (in Biol Philos 32(4):481–505, Calcott 2017) mischaracterizes in an important way the notion of causal specificity proposed by Woodward (in Biol Philos 25(3):287–318, Woodward 2010). This leads him to (1) rely too heavily on one single aspect of Woodward’s analysis on causal specificity; (2) propose an information-theoretic measure he calls ‘precision’ which is partly redundant with, but less general than one of the dimensions in Woodward’s analysis of specificity, without acknowledging Woodward’s analysis; and (3) claim that comparing the specificities of two or more causes under what he calls a competitive analysis of causes, does not permit to capture the distinction between permissive and instructive causes. After having restated Woodward’s analysis of causal specificity, I present an information-theoretic measure (variation of causal information) which, although related to Calcott’s measure, is more general than his and corresponds to the notion of specificity he missed in Woodward's analysis. I then show how this measure can be used, together with mutual causal information (which captures another dimension of specificity in Woodward’s analysis), to distinguish permissive from instructive causes in a competitive analysis of causes.

Contemporary evolutionary medicine has unified the idea of ‘evolutionary mismatch’, derived from the older idea of ‘adaptive lag’ in evolution, with ideas about the mismatch in development and physiology derived from the Developmental Origins of Health and Disease (DOHaD) paradigm. A number of publications in evolutionary medicine have tried to make this theoretical framework explicit. The integrative theory of mismatch captures how organisms track environments across space and time on multiple scales in order to maintain an adaptive match to the environment, and how failures of adaptive tracking lead to disease. In this review, we try to present this complex body of theory as clearly and simply as possible with the aim of facilitating its application in new domains. We introduce terminology, which is as far as possible consistent with earlier usage, to distinguish the different forms of mismatch. Mismatch in its modern form is a productive organizing concept that can help researchers articulate how physiology, development and evolution interact with one another and with environmental change to explain health outcomes.

The idea that selection can go in opposite directions or, more generally, be independent at different levels is well entrenched in both the biological and philosophical literatures. However, this idea is difficult to render precise. On the face of it, it seems unclear how two levels of selection could conflict with one another – and thus be independent if they ultimately refer to the same Darwinian substrate. In this paper, I present an analysis of this problem. I argue that it is impossible for selection at one level to be independent from selection at a different level if independence is to be understood in a strong (metaphysical) sense. However, I propose that independence can be understood in a weaker sense, so long as our conception of independence does not violate the metaphysical dependence of the higher levels on the lower ones. From there, I argue that none of the notions of particle-level or collective-level selection used in the classical formal approaches to multilevel selection capture this weaker form of independence. Finally, I propose a different approach that is compatible with both metaphysical dependence and the weaker form of independence outlined in this paper.

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.

The new foundation for the propensity interpretation of fitness (PIF), developed by Pence and Ramsey (Br J Philos Sci 64:851–881, 2013), describes fitness as a probability distribution that encompasses all possible daughter populations to which the organism may give rise, including daughter populations in which traits might change and the possible environments that members of the daughter populations might encounter. This long-term definition of fitness is general enough to avoid counterexamples faced by previous mathematical conceptions of PIF. However, there seem to be downsides to its generality: the ecological role of fitness involves describing the degree of adaptedness between an organism and the specific environment it inhabits. When all possible changes in traits and all possible environments that a daughter population may encounter are included in the concept, it becomes difficult to see how fitness can fulfill this role. In this paper, we argue that this is a feature of Pence and Ramsey’s view rather than a bug: long-term fitness accommodates evolvability considerations, which concern the role that variation plays in evolutionary processes. Building on the foundations, we show that Pence and Ramsey’s fitness—_F_—can be partitioned into fourths: adaptedness, robustness of adaptedness, and two facets of evolvability. Conceptualizing these last three components forces us to consider the role played by grains of description of both organisms and the environment when thinking about long-term fitness. They track the possibility that there could be a change in type in a daughter population as a way of responding to environmental challenges, or that the type persists in the face of novel environments. We argue that these components are just as salient as adaptedness for long-term fitness. Together, this decomposition of _F_ provides a more accurate picture of the factors involved in long-term evolutionary success.

Philosophers have proposed many accounts of biological function. A coarse-grained distinction can be made between backward-looking views, which emphasise historical contributions to fitness, and forward-looking views, which emphasise the current contribution to fitness or role of a biological component within some larger system. These two views are often framed as being incompatible and conflicting with one another. The emerging field of synthetic biology, which involves applying engineering principles to the design and construction of biological systems, complicates things further by adding intentional design as a source of function. In the current study we explored how biology experts and novices think about function in the context of single-celled, multi-celled, and synthetic organisms. We also explored the extent to which each group were function pluralists, and if they were function pluralists, which accounts of function tended to be endorsed together. The results showed a surprising degree of similarity between experts and novices in most contexts, although certain differences were apparent. Most surprisingly, we found evidence not only of function pluralism in both groups, but pluralism between backward-looking and forward-looking accounts. We discuss these findings in the context of the philosophical debate on function and consider the practical implications for public acceptance of synthetic biology. First, we argue that philosophers of biology should re-examine the purported incompatibility between accounts of function. Second, we argue that due to the introduction of an intentional aetiology in synthetic biology, there may be an inherent conflict between the views of experts and novices when thinking about synthetic biology.

This thesis examines the theoretical and philosophical underpinnings of the concept of natural selection which is pervasively invoked in biology and other ‘evolutionary’ domains. Although what constitutes the process of natural selection appears to be very intuitive (natural selection results from entities exhibiting differences in fitness in a population), this conceals a number of theoretical ambiguities and difficulties. Some of these have been pointed out numerous times; others have hardly been noticed. One aim of this work is to unpack these difficulties and ambiguities; another is to provide new solutions and clarifications to them using a range of philosophical and conceptual tools. The result is a concept of natural selection stripped down from its biological specificities. I start by revisiting the entangled debates over whether natural selection is a cause of evolutionary change as opposed to a mere statistical effect of other causes, at what level this putative cause operates and whether it can be distinguished from drift. Borrowing tools from the causal modelling literature, I argue that natural selection is best conceived as a causal process resulting from individual level differences in a population. I then move to the question of whether the process of natural selection requires perfect transmission of types. I show that this question is ambiguous and can find different answers. From there, I distinguish the process of natural selection from some of its possible products, namely, evolution by natural selection and complex adaptation. I argue that reproduction and inheritance are conceptually distinct from natural selection, and using individual-based models, I demonstrate that they can be conceived as evolutionary products of it. This ultimately leads me to generalise the concepts of heritability and fitness used in the formal equations of evolutionary change. Finally, I argue that concepts of fitness and natural selection crucially depend on the grains of description at and temporal scales over which evolutionary explanations are given. These considerations reveal that the metaphysical status of the process of natural selection is problematic and why neglecting them can lead to flawed arguments in the levels of selection debate.

In a recent reply to Takacs and Bourrat’s article (Biol Philos 37:12, 2022), Autzen and Okasha (Biol Philos 37:37, 2022) question our characterization of the relationship between the geometric mean and arithmetic mean measures of fitness. We here take issue with the claim that our characterization falls prey to the mistakes they highlight. Briefly revisiting what Takacs and Bourrat (Biol Philos 37:12, 2022) accomplished reveals that the key issue of difference concerns cases of deterministic but nonconstant growth. Restricting focus to such cases shows that there is in fact no reason for disagreement.

I extend work from Krakauer et al. (2020), who propose a conception of individuality as the capacity to propagate information through time. From this conception, they develop information-theoretic measures. I identify several shortcomings with these measures—in particular, that they are associative rather than causal. I rectify this shortcoming by deriving a causal information-theoretic measure of individuality. I then illustrate how this measure can be implemented and extended in the context of evolutionary transitions in individuality.

The formalism used to describe evolutionary change in a multilevel setting can be used equally to re-describe the situation as one where all the selection occurs at the individual level. Thus, whether multilevel or individual-level selection occurs seems to be a matter of convention rather than fact. Yet, group selection is regarded by some as an important concept with factual rather than conventional elements. I flesh out an alternative position that regards groups as a target of selection in a way that is not merely definitional fiat and provide a theoretical basis for this position.

The evolution of complex life forms, such as multicellular organisms, is the result of a number of evolutionary transitions in individuality (ETIs). Several attempts have been made to explain their origins, many of which have been internalist (i.e., based largely on internal properties of these life form's ancestors). Here, we show how an externalist perspective, via the ecological scaffolding model in which properties of complex life forms arise from an external scaffold, can shed new light on the question of ETIs. Ultimately, we anticipate progress in the field will occur by recognizing the importance of both the internalist and externalist modes of explanation for ETIs. We illustrate this by considering an extension of the ecological scaffolding model by niche construction in which particles modify the environment which later becomes the scaffold giving rise to collective-level individuality.

The distinction between multilevel selection 1 (MLS1) and multilevel selection 2 (MLS2) is classically regarded as a distinction between two multilevel selection processes involving two different kinds of higher-level fitness. It has been invoked to explain evolutionary transitions in individuality as a shift from an MLS1 to an MLS2 process. In this paper, I argue against the view that the distinction involves two different kinds of processes. I show, starting from the MLS2 version of the Price equation, that it contains the MLS1 version if, following the assumption that a collective constitutively depends (i.e., mereologically supervenes) on its particles, one considers that a necessary map between fitness at two levels exists. I defend the necessity of such a map, making the distinction between MLS1 and MLS2 a matter of perspective and limited knowledge (i.e., epistemic limitations) rather than objective facts. I then provide some reasons why the MLS1/MLS2 distinction nonetheless has some pragmatic value and might be invoked usefully in some contexts, particularly within the context of explaining evolutionary transitions in individuality.

Explaining the emergence of individuality in the process of evolution remains a challenge; it faces the difficulty of characterizing adequately what ‘emergence’ amounts to. Here, I present a pragmatic account of individuality in which I take up this challenge. Following this account, individuals that emerge from an evolutionary transition in individuality are coarse-grained entities: entities that are summaries of lower-level evolutionary processes. Although this account may _prima facie_ appear to ultimately rely on epistemic considerations, I show that it can be used to vindicate the emergence of individuals in a quasi-ontological sense. To this end, I discuss a recent account of evolutionary transitions in individuality proposed by Godfrey-Smith and Kerr (Brit J Philos Sci 64(1):205–222, 2013) where a transition occurs through several stages, each with an accompanying model. I focus on the final stage where higher-level entities are ascribed a separate fitness parameter, while they were not in the previous stages. In light of my account, I provide some justification for why such a change in parameters is necessary and cannot be dismissed as merely epistemic.

The compatibility of the Gaia hypothesis with Darwinism is often challenged on the grounds that (1) to be potent, natural selection requires the existence of a population (whereas Gaia is a single entity), and (2) natural selection requires the entities forming a population to reproduce (whereas Gaia merely persists). However, using the Price equation, I argue, following others, that the Gaia hypothesis can fit squarely within a Darwinian framework because Gaia can exhibit adaptations if a process at a lower level (e.g., an ecosystem) can occur, and the notion of natural selection can be extended to accommodate evolution without reproduction.

This article proposes two conditions to assess whether an entity at a level of description is a unit of selection qua interactor. These two conditions make it possible to (1) distinguish biologically relevant entities from arbitrary ones and (2) distinguish units that can _potentially_ enter a selection process from those that have already done so. I show that the classical approaches used in the literature on units and levels of selection do not fare well with respect to either or both of these desiderata.

Reproductive division of labor has been proposed to play a key role for evolutionary transitions in individuality (ETIs). This chapter provides a guide to a theoretical model that addresses the role of a tradeoff between life-history traits in selecting for a reproductive division of labor during the transition from unicellular to multicellular organisms. In particular, it focuses on the five key assumptions of the model, namely (1) fitness is viability times fecundity; (2) collective traits are linear functions of their cellular counterparts; (3) there is a tradeoff between cell viability and fecundity; (4) cell contribution to the collective is optimal; and (5) there is an initial reproductive cost in large collectives. Thereafter the chapter contrasts two interpretations of the model in the context of ETIs. Originally, the model was interpreted as showing that during the transition to multicellularity the fitness of the lower-level (the cells) is “transferred” to the higher level (the collective). Despite its apparent intuitiveness, fitness transfer may obscure actual mechanisms in metaphorical language. Thus, an alternative and more conservative interpretation of the model that focuses on cell traits and the evolutionary constraints that link them is advocated. In addition, it allows for pursuing subsequent questions, such as the evolution of development.

Evolutionary transitions in individuality (ETIs) involve the formation of Darwinian collectives from Darwinian particles. The transition from cells to multicellular life is a prime example. During an ETI, collectives become units of selection in their own right. However, the underlying processes are poorly understood. One observation used to identify the completion of an ETI is an increase in collective-level performance accompanied by a decrease in particle-level performance, for example measured by growth rate. This seemingly counterintuitive dynamic has been referred to as fitness decoupling and has been used to interpret both models and experimental data. Extending and unifying results from the literature, we show that fitness of particles and collectives can never decouple because calculations of fitness performed over appropriate and equivalent time intervals are necessarily the same provided the population reaches a stable collective size distribution. By way of solution, we draw attention to the value of mechanistic approaches that emphasise traits, and tradeoffs among traits, as opposed to fitness. This trait-based approach is sufficient to capture dynamics that underpin evolutionary transitions. In addition, drawing upon both experimental and theoretical studies, we show that while early stages of transitions might often involve tradeoffs among particle traits, later—and critical—stages are likely to involve the rupture of such tradeoffs. Thus, when observed in the context of ETIs, tradeoff-breaking events stand as a useful marker of these transitions.

The question of whether cultural transmission is faithful has attracted significant debate over the last 30 years. The degree of fidelity with which an object is transmitted depends on 1) the features chosen to be relevant, and 2) the quantity of details given about those features. Once these choices have been made, an object is described at a particular grain. In the absence of conventions between different researchers and across different fields about which grain to use, transmission fidelity cannot be evaluated because it is relative to the choice of grain. In biology, because a genotype-to-phenotype mapping exists and transmission occurs from genotype to genotype, a privileged grain of description exists that circumvents this ‘grain problem.’ In contrast, in cultural evolution, the genotype–phenotype distinction cannot be drawn, rendering claims about fidelity dependent upon researchers’ choices. Thus, due to a lack of unified conventions, claims about fidelity transmission are difficult to evaluate.

The hierarchy of life is the result of a succession of evolutionary transitions in individuality (ETIs). During an ETI, individuals at a particular level of organization interact in such a way as to produce larger-level entities that become individuals in their own right. These new individuals are defined by their capacity to exhibit Darwinian properties of variation, differences in fitness, and heredity. One difficulty in accounting for ETIs is articulating how these properties are acquired at a higher level from the lower ones. Collaborators and I recently proposed the ‘ecological scaffolding’ model in which imposing an ecological scaffold (that is, a structure in the environment) on lower-level entities initiates an ETI. Here, I present a new model that extends this work. Within this new model, I propose a mechanism of scaffold endogenization, demonstrating that collectives can become resilient to the ecological scaffold being removed. This type of resilience is not observed in the ecological scaffolding model. However, classically, a biological individual would be regarded as an entity capable of withstanding environmental changes. Thus, the new model proposed here represents a step towards a more complete explanation for ETIs.

Showing that the arithmetic mean number of offspring for a trait type often fails to be a predictive measure of fitness was a welcome correction to the philosophical literature on fitness. While the higher mathematical moments of a probability-weighted offspring distribution can influence fitness measurement in distinct ways, the geometric mean number of offspring is commonly singled out as the most appropriate measure. For it is well-suited to a compounding process and is sensitive to variance in offspring number. The geometric mean thus proves to be a predictively efficacious measure of fitness in examples featuring discrete generations and within- or between-generation variance in offspring output. Unfortunately, this advance has subsequently led some to conclude that the arithmetic mean is never a good measure of fitness and that the geometric mean should accordingly be the default measure of fitness. We show not only that the arithmetic mean is a perfectly reasonable measure of fitness so long as one is clear about what it refers to, but also that it functions as a more general measure when properly interpreted. It must suffice as a measure of fitness in any case where the geometric mean has been effectively deployed as a measure. We conclude with a discussion about why the mathematical equivalence we highlight cannot be dismissed as merely of mathematical interest.

Natural selection is commonly seen not just as an explanation for adaptive evolution, but as the inevitable consequence of “heritable variation in fitness among individuals”. Although it remains embedded in biological concepts, such a formalisation makes it tempting to explore whether this precondition may be met not only in life as we know it, but also in other physical systems. This would imply that these systems are subject to natural selection and may perhaps be investigated in a biological framework, where properties are typically examined in light of their putative functions. Here we relate the major questions that were debated during a three-day workshop devoted to discussing whether natural selection may take place in non-living physical systems. We start this report with a brief overview of research fields dealing with “life-like” or “proto-biotic” systems, where mimicking evolution by natural selection in test tubes stands as a major objective. We contend the challenge may be as much conceptual as technical. Taking the problem from a physical angle, we then discuss the framework of dissipative structures. Although life is viewed in this context as a particular case within a larger ensemble of physical phenomena, this approach does not provide general principles from which natural selection can be derived. Turning back to evolutionary biology, we ask to what extent the most general formulations of the necessary conditions or signatures of natural selection may be applicable beyond biology. In our view, such a cross-disciplinary jump is impeded by reliance on individuality as a central yet implicit and loosely defined concept. Overall, these discussions thus lead us to conjecture that understanding, in physico-chemical terms, how individuality emerges and how it can be recognised, will be essential in the search for instances of evolution by natural selection outside of living systems.

The most consistent definition of fitness makes it a static property of organisms. However, this is not how fitness is used in many evolutionary models. In those models, fitness is permitted to vary with an organism’s circumstances. According to this second conception, fitness is dynamic. There is consequently tension between these two conceptions of fitness. One recently proposed solution suggests resorting to conditional properties. We argue, however, that this solution is unsatisfactory. Using a very simple model, we show that it can lead to incompatible fitness values and indecision about whether selection actually occurs.