This chapter combines formal models of how the fitness of a collective can become decoupled from the fitness with more empirical work on the volvocine algae. It uses the Volvox clade as a model system. It describes the evolution of altruism in the volvocine green algae. This chapter suggests that altruism may evolve from genes involved in life-history trade-offs. It shows the several cooperation, conflict, and conflict mediation cycles in the volvocine green algae. This cycle of cooperation, conflict, and conflict (...) mediation drives evolutionary transitions in individuality. (shrink)

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. (shrink)

Social evolution theory conventionally takes an externalist explanatory stance, treating observed cooperation as explanandum and the positive assortment of cooperative behaviour as explanans. We ask how the circumstances bringing about this positive assortment arose in the first place. Rather than merely push the explanatory problem back a step, we move from an externalist to an interactionist explanatory stance, in the spirit of Lewontin and the Niche Construction theorists. We develop a theory of ‘social niche construction’ in which we consider biological (...) entities to be both the subject and object of their own social evolution. Some important cases of the evolution of cooperation have the side-effect of causing changes in the hierarchical level at which the evolutionary process acts. This is because the traits that act to align the fitness interests of particles in a collective can also act to diminish the extent to which those particles are bearers of heritable fitness variance, while augmenting the extent to which collectives of such particles are bearers of heritable fitness variance. In this way, we can explain upward transitions in the hierarchical level at which the Darwinian machine operates in terms of particle-level selection, even though the outcome of the process is a collective-level selection regime. Our theory avoids the logical and metaphysical paradoxes faced by other attempts to explain evolutionary transitions. (shrink)

Evolutionary Transitions in Individuality have been responsible for the major transitions in levels of selection and individuality in natural history, such as the origins of prokaryotic and eukaryotic cells, multicellular organisms, and eusocial insects. The integrated hierarchical organization of life thereby emerged as groups of individuals repeatedly evolved into new and more complex kinds of individuals. The Social Protocell Hypothesis proposes that the integrated hierarchical organization of human culture can also be understood as the outcome (...) of an ETI—one that produced a “cultural organism” from a substrate of socially learned traditions that were contained in growing and dividing social communities. The SPH predicts that a threshold degree of evolutionary individuality would have been achieved by 2.0–2.5 Mya, followed by an increasing degree of evolutionary individuality as the ETI unfolded. We here assess the SPH by applying a battery of criteria—developed to assess evolutionary individuality in biological units—to cultural units across the evolutionary history of Homo. We find an increasing agreement with these criteria, which buttresses the claim that an ETI occurred in the cultural realm. (shrink)

This chapter argues that the multilevel selection -1 to MLS-2 model of a major transition is incomplete because it overlooks a crucial component of fitness. It addresses that the evolution of individuality literature has failed to account for expansive fitness and that expansive fitness differences play an important role in the transition to regimes sensitive to the fitness of the corporate agent. It discusses multilevel evolution during the three phases of transitions in individuality: the aggregate phase, the (...) group phase, and the individual phase. This chapter shows that the path a lineage takes through the phases of transitions is not fixed, but determined by ecology. (shrink)

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. (shrink)

Yes, fitness is the central concept of evolutionary biology, but it is an elusive concept. Almost everyone who looks at it seriously comes out in a different place.

The model of major transitions in evolution devised by Maynard Smith and Szathmáry has exerted tremendous influence over evolutionary theorists. Although MTE has been criticized for inconsistently combining different types of event, its ongoing appeal lies in depicting hierarchical increases in complexity by means of evolutionary transitions in individuality. In this paper, we consider the implications of major evolutionary events overlooked by MTE and its ETI-oriented successors, specifically the biological oxygenation of Earth, and the (...) acquisitions of mitochondria and plastids. By reflecting on these missed events, we reveal a central philosophical disagreement over the explanatory goals of major transitions theory that has yet to be made explicit in the literature. We go on to argue that this philosophical disagreement is only reinforced by Szathmáry’s recent revisions of MTE in the form of MTE 2.0. This finding motivates us to propose an alternative explanatory strategy: specifically, an interactionist metabolic perspective on major transitions. A metabolic framework not only avoids many of the criticisms that beset classic and revised MTE models, but also accommodates missing events and provides crucial explanatory components for standard major transitions. Although we do not provide a full-blown alternative theory and do not claim to achieve unity, we explain why foregrounding metabolism is crucial for any attempt to capture the major turning points in evolution, and why it does not lead to unmanageable pluralism. (shrink)

Evolutionary transitions in individuality (ETIs) are events during which individuals at a given level of organization (particles) interact to form higher-level entities (collectives) which are then recognized as new individuals at that level. ETIs are intimately related to levels of selection, which, following Okasha, can be approached from two different perspectives. One, referred to as ‘synchronic’, asks whether selection occurs at the collective level while the partitioning of particles into collectives is taken for granted. The other, referred (...) to as ‘diachronic’, asks about the origins of the partitioning of particles into collectives. After having presented the two perspectives and a classical formalism used to deal with the levels-of-selection question in the literature, namely the multilevel version of the Price equation, I show that because this formalism treats the levels-of-selection question from a synchronic perspective, it is inadequate to explain ETIs. This is because a fundamental aspect of ETIs is the origin of collectives. From there, I develop a framework for levels of selection compatible with the diachronic perspective. This framework relies on the distinction between what I call, on the one hand, a ‘functional aggregative collective trait’, and on the other hand, a ‘functional non-aggregative collective trait’. After having presented this distinction, I implement it in the Price equation, leading to a new statistical partitioning of this equation which, I argue, represents a causal decomposition more relevant for ETIs. Finally, I exploit this partitioning to explain the critical stages of an ETI. In addition to its explanatory power, a measure of the degree of functional non-aggregativity could be used as a proxy for the degree of individuality of a collective. (shrink)

There is a worry that the ‘major transitions in evolution’ represent an arbitrary group of events. This worry is warranted, and we show why. We argue that the transition to a new level of hierarchy necessarily involves a nonselectionist chance process. Thus any unified theory of evolutionary transitions must be more like a general theory of fortuitous luck, rather than a rigid formulation of expected events. We provide a systematic account of evolutionary transitions based on (...) a second-order regularity of chance events, as stipulated by the ZFEL (Zero Force Evolutionary Law). And in doing so, we make evolutionary transitions explainable and predictable, and so not entirely contingent after all. (shrink)

Adaptations can occur at different hierarchical levels (e.g., cells and multicellular organisms), but it can be difficult to identify the level(s) of adaptation in specific cases. A major problem is that selection at a lower level can filter up, creating the illusion of selection at a higher level. We use optimality modeling of the volvocine algae to explore the emergence of genuine group (i.e., colony-level) adaptations. We find that it is helpful to develop an explicit model for what group fitness (...) would be in the absence of group-level relationships between traits and group fitness. We call this “counterfactual fitness,” because in many actual cases of interest there are group-level relationships. Once counterfactual fitness is modeled, the difference between effects that filter up and genuine group selection is explicit and so, therefore, is the distinction between apparent and genuine group adaptations. We call the latter group-specific adaptations. Recognizing group-specific adaptations is important because only group-specific adaptations would cause the lower-level units (cells in this case) to be maladapted if they were to leave the group and enter a global cell-level population. Thus, as group-specific adaptations evolve, they create selective pressure for increased cohesiveness and individuality of groups. This article suggests that group-specific adaptations could be present in the simplest, earliest branching colonial volvocine species, which do not have distinct specialized cells. The article also makes predictions about the kind of empirical evidence needed to support or refute the hypothesis that a particular trait is a group-level adaptation. (shrink)

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. (shrink)

What changes when an evolutionary transition in individuality takes place? Many different answers have been given, in respect of different cases of actual transition, but some have suggested a general answer: that a major transition is a change in the extent to which selection acts at one hierarchical level rather than another. The current paper evaluates some different ways to develop this general answer as a way to characterise the property ‘evolutionary individuality’; and offers a justification (...) of the option taken in Clarke :413–435, 2013)—to define evolutionary individuality in terms of an object’s capacity to undergo selection at its own level. In addition, I suggest a method by which the property can be measured and argue that a problem which is often considered to be fatal to that method—the problem of ‘cross-level by-products’—can be avoided. (shrink)

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. (shrink)

In this dissertation, I explore the reach of the Evolutionary Contingency Thesis—a view that emphasizes the role of dependency relations and chance in evolution. Contingency produces diverse biological entities, processes, and mechanisms. I analyze the implications of evolution’s contingency in three areas. First, I address the problem of evolutionary individuality, which concerns the nature of entities that selection acts on. If we accept Lewontin’s 1970 view that individuals are selected, then what exactly are these individuals? I argue (...) that evolutionary contingency and its diversity of outcomes causes a plurality of individuality types. These types of individuals emerge, evolve, and disappear through evolution. Second, I defend a disunified model of major transitions in evolution. Major transitions mark pivotal turning points in life’s history, such as the transition from unicellularity to multicellularity. Philosophers and biologists aim to unify evolution’s transitions by establishing a property all major events share. I argue that evolutionary contingency’s emphasis on diversity supports a disunified model of transitions. That is, we should expect numerous kinds of major transitions given the diverse sets of outcomes caused by evolutionary contingency. Finally, I extend the contingency analysis to a topic in philosophy of medicine: The nature of cancer. It is common to view cancer as a by-product of our evolution as multicelled creatures. However, Lean and Plutynski argue that cancer is also a direct product of evolution by selection. In both cases, cancer is characterized in terms of how it functions as an object of selection. I argue that the diversity among cancers suggests philosophers should shift their focus to cancers’ structural differences rather than only their functional similarities. (shrink)

This volume describes features of biological autonomy and integrates them into the recent discussion of factors in evolution. In recent years ideas about major transitions in evolution are undergoing a revolutionary change. They include questions about the origin of evolutionary innovation, their genetic and epigenetic background, the role of the phenotype, and of changes in ontogenetic pathways. In the present book, it is argued that it is likewise necessary to question the properties of these innovations and what was (...) qualitatively generated during the macroevolutionary transitions. The author states that a recurring central aspect of macroevolutionary innovations is an increase in individual organismal autonomy whereby it is emancipated from the environment with changes in its capacity for flexibility, self-regulation and self-control of behavior. The first chapters define the concept of autonomy and examine its history and its epistemological context. Later chapters demonstrate how changes in autonomy took place during the major evolutionary transitions and investigate the generation of organs and physiological systems. They synthesize material from various disciplines including zoology, comparative physiology, morphology, molecular biology, neurobiology and ethology. It is argued that the concept is also relevant for understanding the relation of the biological evolution of man to his cultural abilities. Finally the relation of autonomy to adaptation, niche construction, phenotypic plasticity and other factors and patterns in evolution is discussed. The text has a clear perspective from the context of systems biology, arguing that the generation of biological autonomy must be interpreted within an integrative systems approach. (shrink)

I will argue that MLS theory does not provide a complete, self- sufficient approach to theorizing about evolutionary transitions. As a formal, mathematical theory about evolution within a population, it presupposes but does not address the material structure of the population that realizes the model. An MLS model might tell us whether a cooperative trait could be- come fixed in a population, for example, but it won’t be able to explain how the cooperation actually works to produce an (...) adaptive effect on the group’s fitness. It also won’t be able to account for the sources of variety in the pos- sible modes of cooperation available to a population. MLS theory can tell us when fitness has transferred from one level of units to another, but it can give misleading answers unless we have some other, principled guide for picking out units (Clarke 2012; Clarke 2014). Furthermore, it is commonly acknowl- edged among biologists that actually measuring the fitness of an individual— sometimes even obtaining and interpreting proxy measurements—is diffi- cult in practice and prone to error (Hendry 2005; Orr 2009). Hence even if MLS theory were sufficient in principle, there would still be room for other approaches in practice that avoided the difficulties and limitations of depend- ing on fitness alone. (shrink)

The ‘Major Transitions in Evolution’ framework has emerged as the dominant paradigm for understanding the origins of life's hierarchical organization, but it has been criticized on the grounds that it lacks theoretical unity, that is, that the events included in the framework do not constitute a coherent category. I agree with this criticism, and I argue that the best response is to modify the framework so that the events it includes do comprise a coherent category, one whose members share (...) fundamental similarities. Specifically, I recommend defining major transitions as all those, and only those, events and processes that result in the emergence of a new population of evolutionary individuals. Two sorts of change will be required to achieve this. First, events and processes that do not meet this criterion, such as the origins of the genetic code and of human language, should be excluded. Second, events and processes that do meet the criterion, but which have generally been neglected, should be included. These changes would have the dual benefits of making MTEs a philosophically coherent category and of increasing the sample size on which we may infer trends and general principles that may apply to all MTEs. (shrink)

Our aim in the present paper is to approach the nature of life from the perspective of autonomy, showing that this perspective can be helpful for overcoming the traditional Cartesian gap between the physical and cognitive domains. We first argue that, although the phenomenon of life manifests itself as highly complex and multidimensional, requiring various levels of description, individual organisms constitute the core of this multifarious phenomenology. Thereafter, our discussion focuses on the nature of the organization of individual living entities, (...) proposing autonomy as the main concept to grasp it. In the second part of the article we show how autonomy is also fundamental to explaining major evolutionary transitions, in an attempt to rethink evolution from the point of view of the organizational structure of the entities/organisms involved. This gives further support to the idea of autonomy not only as a key to understanding life in general but also the complex expressions of it that we observe on our planet. Finally, we suggest a possible general principle that underlies those evolutionary transitions, which allow for the open-ended redefinition of autonomous systems: namely, the relative dynamic decoupling that must be articulated among distinct parts, modules or modes of operation in these systems. (shrink)

The evolutionary trajectory from non-human to human forms of social organization involves change from experiential- to relational-based systems of social interaction. Social organization derived from biologically and experientially grounded social interaction reached a hiatus with the great apes due to an expansion of individualization of behaviour. The hiatus ended with the introduction of relational-based social interaction, culminating in social organization based on cultural kinship. This evolutionary trajectory links biological origins to cultural outcomes and makes evident the centrality of (...) distributed forms of information for both the boundary and internal structure of human societies as these evolved from prior forms of social organization. (shrink)

Among mycologists, questions persist about what entities should be treated as the fundamental units of fungal populations. This article articulates a coherent view about populations of heterokaryotic fungi and the individuals that comprise them. Using Godfrey-Smith’s minimal concept of a Darwinian population, I argue that entities at two levels of the biological hierarchy satisfy the minimal concept in heterokaryotic fungi: mycelia and nuclei. I provide a preliminary answer to the question of how to understand the relation between these two populations. (...) This article contributes to discussions about the nature of biological individuality, organismality, and evolutionary transitions. (shrink)

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.

Despite remarkable empirical and methodological advances, our theoretical understanding of the evolutionary processes that made us human remains fragmented and contentious. Here, we make the radical proposition that the cultural communities within which Homo emerged may be understood as a novel exotic form of organism. The argument begins from a deep congruence between robust features of Pan community life cycles and protocell models of the origins of life. We argue that if a cultural tradition, meeting certain requirements, arises in (...) the context of such a “social protocell,” the outcome will be an evolutionary transition in individuality whereby traditions and hominins coalesce into a macroscopic bio-socio-technical system, with an organismal organization that is culturally inherited through irreversible fission events on the community level. We refer to the resulting hypothetical evolutionary individual as a “sociont.” The social protocell provides a preadapted source of alignment of fitness interests that addresses a number of open questions about the origins of shared adaptive cultural organization, and the derived genetic adaptations that support them. Also, social cooperation between hominins is no longer in exclusive focus since cooperation among traditions becomes salient in this model. This provides novel avenues for explanation. We go on to hypothesize that the fate of the hominin in such a setting would be mutualistic coadaptation into a part-whole relation with the sociont, and we propose that the unusual suite of derived features in Homo is consistent with this hypothesis. (shrink)

Although natural philosophers have long been interested in individuality, it has been of interest to contemporary philosophers of biology because of its role in different aspects of evolutionary biology. These debates include whether species are individuals or classes, what counts as a unit of selection, and how transitions in individuality occur evolutionarily. Philosophical analyses are often conducted in terms of metaphysics (“what is an individual?”), rather than epistemology (“how can and do researchers conceptualize individuals so as (...) to address some of their scientific goals?”). We review several philosophical distinctions in order to shift attention from metaphysics to epistemology. Many controversies involve epistemological differences rather than metaphysical disagreement. This implies that a pluralist stance about individuality in biology is warranted and has metaphysical consequences because the pluralism emerges from the diversity of scientific interests that investigate the complexity of living phenomena. (shrink)

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. (shrink)

The emergence of individuality during the evolutionary transition from single cells to multicellularity poses a range of problems. A key issue is how variation in lower‐level individuals generates a corporate (collective) entity with Darwinian characteristics. Of central importance to this process is the evolution of a means of collective reproduction, however, the evolution of a means of collective reproduction is not a trivial issue, requiring careful consideration of mechanistic details. Calling upon observations from experiments, we draw attention to (...) proto‐life cycles that emerge via unconventional routes and that transition, in single steps, individuality to higher levels. One such life cycle arises from conflicts among levels of selection and invokes cheats as a primitive germ line: it lays the foundation for collective reproduction, the basis of a self‐policing system, the selective environment for the emergence of development, and hints at a plausible origin for a soma/germ line distinction. (shrink)

Eukaryogenesis is widely viewed as an improbable evolutionary transition uniquely affecting the evolution of life on this planet. However, scientific and popular rhetoric extolling this event as a singularity lacks rigorous evidential and statistical support. Here, we question several of the usual claims about the specialness of eukaryogenesis, focusing on both eukaryogenesis as a process and its outcome, the eukaryotic cell. We argue in favor of four ideas. First, the criteria by which we judge eukaryogenesis to have required a (...) genuinely unlikely series of events 2 billion years in the making are being eroded by discoveries that fill in the gaps of the prokaryote:eukaryote “discontinuity.” Second, eukaryogenesis confronts evolutionary theory in ways not different from other evolutionary transitions in individuality; parallel systems can be found at several hierarchical levels. Third, identifying which of several complex cellular features confer on eukaryotes a putative richer evolutionary potential remains an area of speculation: various keys to success have been proposed and rejected over the five-decade history of research in this area. Fourth, and perhaps most importantly, it is difficult and may be impossible to eliminate eukaryocentric bias from the measures by which eukaryotes as a whole are judged to have achieved greater success than prokaryotes as a whole. Overall, we question whether premises of existing theories about the uniqueness of eukaryogenesis and the greater evolutionary potential of eukaryotes have been objectively formulated and whether, despite widespread acceptance that eukaryogenesis was “special,” any such notion has more than rhetorical value. (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)

The levels of selection problem was central to Maynard Smith’s work throughout his career. This paper traces Maynard Smith’s views on the levels of selection, from his objections to group selection in the 1960s to his concern with the major evolutionary transitions in the 1990s. The relations between Maynard Smith’s position and those of Hamilton and G.C. Williams are explored, as is Maynard Smith’s dislike of the Price equation approach to multi-level selection. Maynard Smith’s account of the ‘core (...) Darwinian principles’ is discussed, as is his debate with Sober and Wilson (1998) over the status of trait-group models, and his attitude to the currently fashionable concept of pluralism about the levels of selection. (shrink)

Criticisms of the “container” model of pregnancy picturing female and embryo as separate entities multiply in various philosophical and scientific contexts during the last decades. In this paper, we examine how this model underlies received views of pregnancy in evolutionary biology, in the characterization of the transition from oviparity to viviparity in mammals and in the selectionist explanations of pregnancy as an evolutionary strategy. In contrast, recent evo-devo studies on eutherian reproduction, including the role of inflammation and new (...) maternal cell types, gather evidence in favor of considering pregnancy as an evolved relational novelty. Our thesis is that from this perspective we can identify the emergence of a newhistoricalindividual in evolution. In evo-devo, historical units are conceptualized as evolved entities which fulfill two main criteria, their continuous persistence and their non-exchangeability. As pregnancy can be individuated in this way, we contend that pregnant females are historical individuals. We argue that historical individuality differs from, and coexists with, other views of biological individuality as applied to pregnancy (the physiological, the evolutionary and the ecological one), but brings forward an important new insight which might help dissolve misguided conceptions. (shrink)

Species’ adaptation to their environments occurs via a range of mechanisms of adaptation. These include genetic adaptations as well as non-traditional inheritance mechanisms such as learned behaviors, niche construction, epigenetics, horizontal gene transfer, and alteration of the composition of a host’s associated microbiome. We propose to supplement these with another modality of eco-evolutionary dynamics: cases in which adaptation to the environment occurs via what may be called a “distributed adaptation,” in which the adaptation is not conferred via something carried (...) by an individual of the adapted species (as with genes, behavior, or associated microbes), but by some structural or compositional aspect of the population. Put differently, the adaptively relevant information cannot be reduced to information possessed by a single individual, whether genetic or otherwise. Rather, the adaptively relevant information is distributed, and is found strictly at the population level. While human culture is presumably such a case, as may be cases found in social insects, we want to suggest that there are other cases that belong to this category and to explore its evolutionary implications. In particular, we discuss the factors that affect whether adaptive information is stored in a distributed way, to what degree, and what kinds of adaptive information are most likely to be found in this modality of adaptation. (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)

One of the major developments in cancer research in recent years has been the construction of models that treat cancer as a cellular population subject to natural selection. We expand on this idea, drawing upon multilevel selection theory. Cancer is best understood in our view from a multilevel perspective, as both a by-product of selection at other levels of organization, and as subject to selection at several levels of organization. Cancer is a by-product in two senses. First, cancer cells co-opt (...) signaling pathways that are otherwise adaptive at the organismic level. Second, cancer is also a by-product of features distinctive to the metazoan lineage: cellular plasticity and modularity. Applying the multilevel perspective in this way permits one to explain transitions in complexity and individuality in cancer progression. Our argument is a reply to Germain’s scepticism towards the explanatory relevance of natural selection for cancer. The extent to which cancer fulfills the conditions for being a paradigmatic Darwinian population depends on the scale of analysis, and the details of the purported selective scenario. Taking a multilevel perspective clarifies some of the complexities surrounding how to best understand the relevance of evolutionary thinking in cancer progression. (shrink)

Transitive inference (TI) refers to social cognition that facilitates the discernment of unknown relationships between individuals using known relationships. It is extensively reported that TI evolves in animals living in a large group because TI could assess relative rank without deducing all dyadic relationships, which averts costly fights. The relationships in a large group become so complex that social cognition may not be developed adequately to handle such complexity. If members apply TI to all possible members in the group, TI (...) requires extremely highly developed cognitive abilities especially in a large group. Instead of developing cognitive abilities significantly, animals may apply simplified TI we call reference TI in this study as heuristic approaches. The reference TI allows members to recognize and remember social interactions only among a set of reference members rather than all potential members. Our study assumes that information processes in the reference TI comprises (1) the number of reference members based on which individuals infer transitively, (2) the number of reference members shared by the same strategists, and (3) memory capacity. We examined how information processes evolve in a large group using evolutionary simulations in the hawk–dove game. Information processes with almost any numbers of reference members could evolve in a large group as long as the numbers of shared reference member are high because information from the others’ experiences is shared. TI dominates immediate inference, which assesses relative rank on direct interactions, because TI could establish social hierarchy more rapidly applying information from others’ experiences. (shrink)

This collection reports on the latest research on an increasingly pivotal issue for evolutionary biology: cooperation. The chapters are written from a variety of disciplinary perspectives and utilize research tools that range from empirical survey to conceptual modeling, reflecting the rich diversity of work in the field. They explore a wide taxonomic range, concentrating on bacteria, social insects, and, especially, humans. -/- Part I (“Agents and Environments”) investigates the connections of social cooperation in social organizations to the conditions that (...) make cooperation profitable and stable, focusing on the interactions of agent, population, and environment. Part II (“Agents and Mechanisms”) focuses on how proximate mechanisms emerge and operate in the evolutionary process and how they shape evolutionary trajectories. Throughout the book, certain themes emerge that demonstrate the ubiquity of questions regarding cooperation in evolutionary biology: the generation and division of the profits of cooperation; transitions in individuality; levels of selection, from gene to organism; and the “human cooperation explosion” that makes our own social behavior particularly puzzling from an evolutionary perspective. (shrink)

This article argues that the (epi)genetic, cultural, symbolic, and environmental transmission channels are insufficient to explain the structure of modern human societies. Economic exchange of knowledge embodied in goods and services constitutes an additional transmission channel that makes more efficient use of limited human cognitive capacity. Economic exchange results in a gradual shift in societies from task-based division of labor to cognitive specialization. This shifts scarce cognitive resources away from production and into learning. It accelerates learning and reinforces the drive (...) towards specialization. Cognitive specialization may constitute another “major transition” towards a higher level of aggregation in human societies, with properties that differ from symbolic transmission. Collective control of individual market-based exchange is ensured by means of economic institutions that put a constraint on individual behavior. (shrink)

The paper links discussions of two topics: biological individuality and the simplest forms of mentality. I discuss several attempts to locate the boundary between metabolic activity and ‘minimal cognition.’ I then look at differences between the kinds of individuality present in unicellular life, multicellular life in general, and animals of several kinds. Nervous systems, which are clearly relevant to cognition and subjectivity, also play an important role in the form of individuality seen in animals. The last part (...) of the paper links these biological transitions to the evolutionary history of subjective experience. (shrink)

Andrew Bourke’s Principles of Social Evolution identifies three stages that characterize an evolutionary transition in individuality and deploys inclusive fitness theory to explain each stage. The third stage, social group transformation, has hitherto received relatively little attention from inclusive fitness theorists. In this review, I first discuss Bourke’s “virtual dominance” hypothesis for the evolution of the germ line. I then contrast Bourke’s inclusive fitness approach to the major transitions with the multi-level approach developed by Richard Michod, Samir (...) Okasha and others. I suggest that, rather than choosing between these approaches, we should exploit the strengths of both. Finally, I stress the need for a firmer conceptual grasp of the nature of social group transformation. (shrink)

Evolutionary theory is formulated in terms of individuals that carry heritable information and are subject to selective pressures. However, individuality itself is a trait that had to evolve - an individual is not an indivisible entity, but a result of evolutionary processes that necessarily begin at the lower level of hierarchical organisation. Traditional approaches to biological individuality focus on cooperation and relatedness within a group, division of labour, policing mechanisms and strong selection at the higher level. (...) Nevertheless, despite considerable theoretical progress in these areas, a full dynamical first-principles account of how new types of individuals arise is missing. To the extent that individuality is an emergent trait, the problem can be approached by recognising the importance of individuating mechanisms that are present from the very beginning of the transition, when only lower-level selection is acting. Here we review some of the most influential theoretical work on the role of individuating mechanisms in these transitions, and demonstrate how a lower-level, bottom-up evolutionary framework can be used to understand biological complexity involved in the origin of cellular life, early eukaryotic evolution, sexual life cycles and multicellular development. Some of these mechanisms inevitably stem from environmental constraints, population structure and ancestral life cycles. Others are unique to specific transitions - features of the natural history and biochemistry that are co-opted into conflict mediation. Identifying mechanisms of individuation that provide a coarse-grained description of the system's evolutionary dynamics is an important step towards understanding how biological complexity and hierarchical organisation evolves. In this way, individuality can be reconceptualised as an approximate model that with varying degrees of precision applies to a wide range of biological systems. (shrink)

The paper investigates Maurice Merleau-Ponty’s discussion of body and space and Gilles Deleuze’s reading of Francis Bacon’s work, in order to derive a renegotiated interrelation between habitual body, phenomenal space, preferential plane and constructive line. The resulting system is ap- plied as a filter to understand the sartorial fashion of Rei Kawakubo and Hussein Chalayan and their potential as a spatial prosthesis: the operative third skin. If the evolutionary nature of culture demands a constant change, how does the surface (...) of a third skin, which embodies the generative of stable/ unstable, respond to changes of context? The fleeting, shifting conditions of contemporary culture/ lifestyle rely upon, result in, and reflect one constant, change: change of working conditions, family structures, modes of inhabitation, relation networks, of userprofile and identity, of social and territorial boundaries. We occupy the shift- ing spatial parameters of a transitional supermodern environment. Culture, as enacted or embodied through each of these fields, is regulated by a number of abstract and factual variables that interplay constantly: time, space, movement, surface, individual, and data. Elizabeth Grosz argues that culture is an evolutionary effect: it regenerates itself in order to ensure the survival of the species. Each “prosthetic” expression of culture – language, fashion, architecture, etc. – changes repeatedly. Here change is not an end in itself, but a means. And the most successful prosthesis may not be the one that is able to answer the largest number of challenges, but one which itself undergoes a process of learning, self-modification, and differentiation – in short, a process of evolution. Any prosthesis is by nature an extension of the body. In the case of architecture and fashion the prosthesis addressing change is most often external to the body – a “cultural fur” or a surface phenomenon, that is, a highly profiled supplementary skin. As with all prostheses, their respective life-span depends on their ability to reflect a change in context and value systems. They are adapted or updated, if not, they vanish. Any situation of change is processed as a differentiation between the actual and the virtual of a given context. Grosz identifies distinctions between the actual and virtual, the real and the possible: the possible is a preformed real that has not yet received its final materiality, and thus delineates a range of options of becoming. The real is the blueprint of the possible, negotiated by factual limitations, and it is conjoint with the actual through a process of differentiation and divergence. The virtual comprises alternate variations of the actual, it defines a realm of deviation from the blueprint. In order to be responsive to change, the balance between the actual and virtual thus must be rendered unstable: “The virtual requires the actual to diverge, to differentiate itself, to proceed by way of division and disruption, forging modes of actualisation that will transform this virtual into others unforeseen or uncontained within it.” 4 The integration of the virtual allows a re-ordering of the blueprint, a return to the crossroad of possibilities, unlimiting and processing an alternative real, and establishing a state of continual change. A repeated change – not as a choice between a number of options but as a gradual process – marks the moment of evolution, and requires a dynamic system. Such a dynamic can be rendered as an adaptable, flexible, modular, mobile, or morphing system of change. The key lies with the fluency and ability of adaptation for the proposed differentiation between actual and virtual – thus it is an elastic change that is required. The nature of this elasticity is encoded in a repeated repositioning of the variables: a constant fine-tuning of a maximum number of parameters that engineer, alter and define the blueprint. When looking for a dynamic system that incorporates a transformation of skin or space, we are in search of dynamics through an operative surface, controlled by means of the constructive line. Both operative surface and constructive line are generative methods for the formation and form finding of the second and third skin of sartorial fashion and architecture respectively, as they both produce inhabitable or wearable envelopes with a specific responsiveness. Both professions share communication, coding and signage, form information programs, pattern charts, volume outlines, texture fields, surface operations, and implement electronic or digital extensions. In both, the constructive line shapes the surface twice: before production and in operation. The surface demarcates space, spatial envelope, enclosing garment, field of action. In which way can an operative system of the surface with stable/ unstable conditioning generate a phenomenological or evolutionary change in the reconstruction of body and context? How do time, space, movement, surface, individual and data interact in this framework? What is the impact of surface strategies and constructive line on that system? (shrink)

The fitness of any evolutionary unit can be understood in terms of its two basic components: fecundity (reproduction) and viability (survival). Trade-offs between these fitness components drive the evolution of life-history traits in extant multicellular organisms. We argue that these trade-offs gain special significance during the transition from unicellular to multicellular life. In particular, the evolution of germ–soma specialization and the emergence of individuality at the cell group (or organism) level are also consequences of trade-offs between the two (...) basic fitness components, or so we argue using a multilevel selection approach. During the origin of multicellularity, we study how the group trade-offs between viability and fecundity are initially determined by the cell level trade-offs, but as the transition proceeds, the fitness trade-offs at the group level depart from those at the cell level. We predict that these trade-offs begin with concave curvature in single-celled organisms but become increasingly convex as group size increases in multicellular organisms. We argue that the increasingly convex curvature of the trade-off function is driven by the cost of reproduction which increases as group size increases. We consider aspects of the biology of the volvocine green algae – which contain both unicellular and multicellular members – to illustrate the principles and conclusions discussed. (shrink)

Extensive social choice theory is used to study the problem of measuring group fitness in a two-level biological hierarchy. Both fixed and variable group size are considered. Axioms are identified that imply that the group measure satisfies a form of consequentialism in which group fitness only depends on the viabilities and fecundities of the individuals at the lower level in the hierarchy. This kind of consequentialism can take account of the group fitness advantages of germ-soma specialization, which is not possible (...) with an alternative social choice framework proposed by Okasha, but which is an essential feature of the index of group fitness for a multicellular organism introduced by Michod, Viossat, Solari, Hurand, and Nedelcu to analyze the unicellular-multicellular evolutionary transition. The new framework is also used to analyze the fitness decoupling between levels that takes place during an evolutionary transition. (shrink)

Attempts to explain the origin of macroevolutionary innovations have been only partially successful. Here it is proposed that the patterns of major evolutionary transitions have to be understood first, before it is possible to further analyse the forces behind the process. The hypothesis is that major evolutionary innovations are characterized by an increase in organismal autonomy, in the sense of emancipation from the environment. After a brief overview of the literature on this subject, increasing autonomy is defined (...) as the evolutionary shift in the individual system–environment relationship, such that the direct influences of the environment are gradually reduced and a stabilization of self-referential, intrinsic functions within the system is generated. This is described as relative autonomy because numerous interconnections with the environment and dependencies upon it are retained. Features of increasing autonomy are spatial separations, an increase in homeostatic functions and in body size, internalizations and an increase in physiological and behavioral flexibility. It is described how these features are present in different combinations in the major evolutionary transitions of metazoans and, consequently, how they should be taken into consideration when evolutionary innovations are studied. The hypothesis contributes to a reconsideration of the relationship between organisms and their environment. (shrink)

Successful transitions to a sustainable bioeconomy require novel technologies, processes, and practices as well as a general agreement about the overarching normative direction of innovation. Both requirements necessarily involve collective action by those individuals who purchase, use, and co-produce novelties: the consumers. Based on theoretical considerations borrowed from evolutionary innovation economics and consumer social responsibility, we explore to what extent consumers’ scope of action is addressed in the scientific bioeconomy literature. We do so by systematically reviewing bioeconomy-related publications (...) according to the extent to which consumers are regarded as passive vs. active, and different domains of consumer responsibility. We find all aspects of active consumption considered to varying degrees but observe little interconnection between domains. In sum, our paper contributes to the bioeconomy literature by developing a novel coding scheme that allows us to pinpoint different aspects of consumer activity, which have been considered in a rather isolated and undifferentiated manner. Combined with our theoretical considerations, the results of our review reveal a central research gap which should be taken up in future empirical and conceptual bioeconomy research. The system-spanning nature of a sustainable bioeconomy demands an equally holistic exploration of the consumers’ prospective and shared responsibility for contributing to its coming of age, ranging from the procurement of information on bio-based products and services to their disposal. (shrink)

For the first time, the article analyses certain aspects of Russian poetry of the “Silver Age” in order to identify the rudiments or features which are characteristic of the postmodern creative paradigm. It is noted that a number of poets almost do not have any postmodernist tendencies. Despite the fact it is proved that postmodernism denies the personality-centric and aesthetically oriented concept of modernism, it nevertheless arose on the basis of modernism and has sharpened evolutionary features formulated in the (...) first half of the 20th century. The article aims to prove a hypothesis that arises in the authors during a preliminary perceptual reading of the poets` works of the “Silver Age”: in the early 20th century. Sporadically and consistently in individual authors can be observed irony, play, reconstruction and performance as precursor of postmodernist creative thinking. Specialties of the Russian poetry of the “Silver Age”, which directly correlate with postmodernist tendencies of the second half of the 20th century is not a description itself, but the realization of reality, ambivalence, as well as following the linguistic and figurative, conceptual, motive levels of gradual transitions between the paradigms of “symbolism – modernism” and “modernism – postmodernism”. The international significance of the article is that the material of one of the Eastern European literatures has proved the existence of postmodern features in the first half of the 20th century for the first time, which can serve as a deeper research in the field of literary typology, continuity; culturology and anthropology. (shrink)

What are the prospects for a monistic view of biological individuality given the multiple epistemic roles the concept must satisfy? In this paper, I examine the epistemic adequacy of two recent accounts based on the capacity to undergo natural selection. One is from Ellen Clarke, and the other is by Peter Godfrey-Smith. Clarke’s position reflects a strong monism, in that she aims to characterize individuality in purely functional terms and refrains from privileging any specific material properties as important (...) in their own right. I argue that Clarke’s functionalism impairs the epistemic adequacy of her account compared to a middle-ground position taken by Godfrey-Smith. In comparing Clarke and Godfrey-Smith’s account, two pathways emerge to pluralism about biological individuality. The first develops from the contrast between functionalist and materialist approaches, and the second from an underlying temporal structure involved in using evolutionary processes to define individuality. (shrink)

The Committee on Common Problems of Genetics, Paleontology, and Systematics (United States National Research Council) marks part of a critical transition in American evolutionary studies. Launched in 1942 to facilitate cross-training between genetics and paleontology, the Committee was also designed to amplify paleontologist voices in modern studies of evolutionary processes. During coincidental absences of founders George Gaylord Simpson and Theodosius Dobzhansky, an opportunistic Ernst Mayr moved into the project's leadership. Mayr used the opportunity for programmatic reforms he had (...) been pursuing elsewhere for more than a decade. These are evident in the Bulletins he distributed under Committee auspices. In his brief tenure as Committee leader, Mayr gained his first substantial foothold within the coalescing community infrastructure of evolutionary studies. Carrying this momentum forward led Mayr directly into the project to launch the journal Evolution. The sociology of interdisciplinary activity provides useful tools for understanding the Committee's value in the broad sweep of change in evolutionary studies during the synthesis period. (shrink)