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

He was not just my teacher and my friend. He was my hero, a man who was quietly but passionately committed to truth, to clarity, to understanding everything under the sun–and to making himself understood. More than anybody else he has made me proud to be a philosopher, so I would like to dedicate my Presidential Address to his memory.

At the heart of evolutionary theory lays the notion of replication. Unfortunately, this notion is far less exact than the weight of its importance. In this paper, it is argued that replication always involves coding. Furthermore, when a theory of evolution is built on replication based on coding, a unifying and coherent picture arises that sheds new light on some of the controversies and open questions in contemporary biology, such as what are the roles of phylogeny and ontogeny in evolution, (...) and how to characterize and explain the increase of biological complexity. (shrink)

This article reviews the seven “visions” of evolution proposed by Depew and Weber , concluding that each posited relationship between natural selection and self-organization has suited different aims and approaches. In the second section of the article, we show that these seven viewpoints may be collapsed into three fundamentally different ones: natural selection drives evolution; self-organization drives evolution; and natural selection and self-organization are complementary aspects of the evolutionary process. We then argue that these three approaches are not mutually exclusive, (...) since each may apply to different stages of development of different systems. What emerges from our discussion is a more encompassing view: that self-organization proposes what natural selection disposes. (shrink)

Bayesian learning theory and evolutionary theory both formalize adaptive competition dynamics in possibly high‐dimensional, varying, and noisy environments. What do they have in common and how do they differ? In this paper, we discuss structural and dynamical analogies and their limits, both at a computational and an algorithmic‐mechanical level. We point out mathematical equivalences between their basic dynamical equations, generalizing the isomorphism between Bayesian update and replicator dynamics. We discuss how these mechanisms provide analogous answers to the challenge of adapting (...) to stochastically changing environments at multiple timescales. We elucidate an algorithmic equivalence between a sampling approximation, particle filters, and the Wright‐Fisher model of population genetics. These equivalences suggest that the frequency distribution of types in replicator populations optimally encodes regularities of a stochastic environment to predict future environments, without invoking the known mechanisms of multilevel selection and evolvability. A unified view of the theories of learning and evolution comes in sight. (shrink)

Both paleobiology and investigations of ‘major evolutionary transitions’ are intimately concerned with the macroevolutionary shape of life. It is surprising, then, how little studies of major transitions are informed by paleontological perspectives and. I argue that this disconnect is partially justified because paleobiological investigation is typically ‘phenomena-led’, while investigations of major transitions are ‘theory-led’. The distinction turns on evidential relevance: in the former case, evidence is relevant in virtue of its relationship to some phenomena or hypotheses concerning those phenomena; in (...) the latter, evidence is relevant in virtue of providing insights into, or tests of, an abstract body of theory. Because paleobiological data is by-and-large irrelevant to the theory which underwrites the traditional conception of major transitions, it is of limited use to that research program. I suggest that although the traditional conception of major transitions is neither ad-hoc or problematically incomplete, its promise of providing unificatory explanations of the transitions is unlikely to be kept. Further, examining paleobiological investigations of mass extinctions and organogenesis, I further argue that whether or not transitions in paleobiology count as ‘major’ turns on how we conceive of major transitions ; although major transitions potentially have a unified theoretical basis, recent developments suggest that investigations are becoming increasingly phenomena-led; adopting phenomena-led investigations maximizes the evidence available to paleobiologists. (shrink)

Despite its current popularity, “emergence” is a concept with a venerable history and an elusive, ambiguous standing in contemporary evolutionary theory. This paper briefly recounts the history of the term and details some of its current usages. Not only are there radically varying interpretations about how to define emergence but “reductionist” and “holistic” theorists hold very different views about the issue of causation. However, these two seemingly polar positions are not irreconcilable. Reductionism, or detailed analysis of the parts and their (...) interactions, is essential for answering the “how” question in evolution—how does a complex living system work? But holism is equally necessary for answering the “why” question—why did a particular arrangement of parts evolve? In order to answer the “why” question, a broader, multi-leveled paradigm is required. The reductionist approach to explaining emergent complexity has entailed a search for underlying “laws of emergence.” In contrast, the “Synergism Hypothesis” focuses on the “economics”—the functional effects produced by emergent wholes and their selective consequences in evolutionary change. This paper also argues that emergent phenomena represent, in effect, a subset of a larger universe of cooperative, synergistic effects in the natural world. (shrink)

Darwin’s classic ‘Origin of Species’ (Darwin 1859) described forces of selection acting upon individuals, but there remains a great deal of controversy about what exactly the status and definition of a biological individual is. Recently some authors have argued that the individual is dispensable – that an inability to pin it down is not problematic because little rests on it anyway. The aim of this paper is to show that there is a real problem of biological individuality, and an urgent (...) need to arbitrate over the current plethora of solutions to it. (shrink)

The problem of plant individuality is something which has vexed botanists throughout the ages, with fashion swinging back and forth from treating plants as communities of individuals (Darwin 1800 ; Braun and Stone 1853 ; Münch 1938 ) to treating them as organisms in their own right, and although the latter view has dominated mainstream thought most recently (Harper 1977 ; Cook 1985 ; Ariew and Lewontin 2004 ), a lively debate conducted mostly in Scandinavian journals proves that the issues (...) are far from being resolved (Tuomi and Vuorisalo 1989b ; Fagerström 1992 ; Pan and Price 2001 ). In this paper I settle the matter once and for all, by showing which elements of each side are correct. (shrink)

Do multi-level selection explanations of the evolution of social traits deepen the understanding provided by single-level explanations? Central to the former is a mathematical theorem, the multi-level Price decomposition. I build a framework through which to understand the explanatory role of such non-empirical decompositions in scientific practice. Applying this general framework to the present case places two tasks on the agenda. The first task is to distinguish the various ways of suppressing within-collective variation in fitness, and moreover to evaluate their (...) biological interest. I distinguish four such ways: increasing retaliatory capacity, homogenising assortment, and collapsing either fitness structure or character distribution to a mean value. The second task is to discover whether the third term of the Price decomposition measures the effect of any of these hypothetical interventions. On this basis I argue that the multi-level Price decomposition has explanatory value primarily when the sharing-out of collective resources is `subtractable'. Thus its value is more circumscribed than its champions Sober and Wilson (1998) suppose. (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)

Many scholars assume a connection between the evolution of language and that of distinctively human group-level morality. Unfortunately, such thinkers frequently downplay a central implication of modern Darwinian theory, which precludes the possibility of innate psychological mechanisms evolving to benefit the group at the expense of the individual. Group level moral regulation is indeed central to public life in all known human communities. The production of speech acts would be impossible without this. The challenge, therefore, is to explain on a (...) Darwinian basis how life could have become subject to the rule of law. Only then will we have an appropriate social framework in which to contextualize our models of how language may have evolved. (shrink)

Understanding how cooperation evolves is central to explaining some core features of our biological world. Many important evolutionary events, such as the arrival of multicellularity or the origins of eusociality, are cooperative ventures between formerly solitary individuals. Explanations of the evolution of cooperation have primarily involved showing how cooperation can be maintained in the face of free-riding individuals whose success gradually undermines cooperation. In this paper I argue that there is a second, distinct, and less well explored, problem of cooperation (...) that I call the generation of benefit. Focusing on how benefit is generated within a group poses a different problem: how is it that individuals in a group can (at least in principle) do better than those who remain solitary? I present several different ways that benefit may be generated, each with different implications for how cooperation might be initiated, how it might further evolve, and how it might interact with different ways of maintaining cooperation. I argue that in some cases of cooperation, the most important underlying “problem” of cooperation may be how to generate benefit, rather than how to reduce conflict or prevent free-riding. (shrink)

This is the second of two articles in which I reflect on “generalized Darwinism” as currently discussed in evolutionary economics. In the companion article (Callebaut, Biol Theory 6. doi: 10.1007/s13752-013-0086-2, 2011, this issue) I approached evolutionary economics from the naturalistic perspectives of evolutionary epistemology and the philosophy of biology, contrasted evolutionary economists’ cautious generalizations of Darwinism with “imperialistic” proposals to unify the behavioral sciences, and discussed the continued resistance to biological ideas in the social sciences. Here I assess Generalized Darwinism (...) as propounded by Geoffrey Hodgson, Thorbjørn Knudsen, and others, concentrating on the roles of theory and model building in science (and the roles of analogy and metaphor therein), generative replication, and the relation between selection and self-organization. I then point to advances in current biology that promise to be more fruitful as sources of inspiration for evolutionary economics than the project to generalize Darwinism in its current, “hardened Modern Synthesis” form; and I draw some conclusions. (shrink)

This is the first of two articles in which I reflect on “generalized Darwinism” as currently discussed in evolutionary economics. I approach evolutionary economics by the roundabouts of evolutionary epistemology and the philosophy of biology, and contrast evolutionary economists’ cautious generalizations of Darwinism with “imperialistic” proposals to unify the behavioral sciences. I then discuss the continued resistance to biological ideas in the social sciences, focusing on the issues of naturalism and teleology. In the companion article (Callebaut, Biol Theory 6. doi: (...) 10.1007/s13752-013-0087-1, 2011, this issue) I assess generalized Darwinism, concentrating on the roles of theory and model building, generative replication, and the relation between selection and self-organization; and I point to advances in biology that promise to be more fruitful as sources of inspiration for evolutionary economics than the project to generalize Darwinism in its current, “hardened Modern Synthesis” form. (shrink)

Kevin Laland and colleagues have put forward a number of arguments motivating an extended evolutionary synthesis. Here I examine Laland et al.'s central concept of reciprocal causation. Reciprocal causation features in many arguments supporting an expanded evolutionary framework, yet few of these arguments are clearly delineated. Here I clarify the concept and make explicit three arguments in which it features. I identify where skeptics can—and are—pushing back against these arguments, and highlight what I see as the empirical, explanatory, and methodological (...) issues at stake. (shrink)

In the past 150 years there have been many attempts to draw parallels between cultural and biological evolution. Most of these attempts were flawed due to lack of knowledge and false ideas about evolution. In recent decades these shortcomings have been cleared away, thus triggering a renewed interest in the subject. This paper offers a critical survey of the main issues and arguments in that discussion. The paper starts with an explication of the Darwinian algorithm of evolution. It is argued (...) that this ‘formula’ is substrate-neutral, which means that biological evolution might not be the only Darwinian process. Other dynamic systems could evolve as well provided that certain conditions are met. In the case of human culture this seems to be the case. The paper then focuses on the notion of niche construction. It is argued that niche construction plays a crucial role in human evolution because it has altered the sources of natural selection and thus the path of evolution. Next two approaches to cultural evolution are discussed: sociobiology and memetics. I will argue that both approaches have flaws because they either underestimate the influence of culture or they stretch analogies too far. Finally two common objections against the idea of cultural evolution are addressed: Lamarckian inheritance and the issue of guided variation. I will argue that although cultural evolution differs from biological evolution in several respects, these discrepancies do not jeopardize the claim that cultural evolution is essentially Darwinian. (shrink)

Schizophrenia is a worldwide, prevalent disorder with a multifactorial but highly genetic aetiology. A constant prevalence rate in the face of reduced fecundity has caused some to argue that an evolutionary advantage exists in unaffected relatives. Here, I critique this adaptationist approach, and review – and find wanting – Crow's “speciation” hypothesis. In keeping with available biological and psychological evidence, I propose an alternative theory of the origins of this disorder. Schizophrenia is a disorder of the social brain, and it (...) exists as a costly trade-off in the evolution of complex social cognition. Paleoanthropological and comparative primate research suggests that hominids evolved complex cortical interconnectivity (in particular, frontotemporal and frontoparietal circuits) to regulate social cognition and the intellectual demands of group living. I suggest that the ontogenetic mechanism underlying this cerebral adaptation was sequential hypermorphosis and that it rendered the hominid brain vulnerable to genetic and environmental insults. I argue that changes in genes regulating the timing of neurodevelopment occurred prior to the migration of Homo sapiens out of Africa 100,000–150,000 years ago, giving rise to the schizotypal spectrum. While some individuals within this spectrum may have exhibited unusual creativity and iconoclasm, this phenotype was not necessarily adaptive in reproductive terms. However, because the disorder shared a common genetic basis with the evolving circuitry of the social brain, it persisted. Thus schizophrenia emerged as a costly trade-off in the evolution of complex social cognition. Key Words: cortical connectivity; evolution; heterochrony; metarepresentation; primates; psychiatry; schizophrenia; social brain; social cognition. (shrink)

In recent years, the concept of evolvability has been gaining in prominence both within evolutionary developmental biology (evo-devo) and the broader field of evolutionary biology. Despite this, there remains considerable disagreement about what evolvability is. This article offers a solution to this problem. I argue that, in focusing too closely on the role played by evolvability as an explanandum in evo-devo, existing philosophical attempts to clarify the evolvability concept have been overly narrow. Within evolutionary biology more broadly, evolvability offers a (...) robust explanation for the evolutionary trajectories of populations. Evolvability is an abstract, robust, dispositional property of populations, which captures the joint causal influence of their internal features on the outcomes of evolution (as opposed to the causal influence of selection, which is often characterized as external). When considering the nature of the physical basis of this disposition, it becomes clear that the many existing definitions of evolvability at play within evo-devo should be understood as capturing only aspects of a much broader phenomenon. 1 Introduction2 The Problem of Evolvability3 The Theoretical Role of Evolvability in Evolutionary Biology 3.1 The explanatory targets of evolutionary biology 3.2 Selection-based explanations 3.3 Lineage explanations 3.4 Evolvability-based explanations 3.5 What properties must evolvability have?4 What Evolvability Really Is 4.1 Making sense of ft 4.2 Making sense of x and b5 What of the Limbs? The Power of E6 Conclusion. (shrink)

Does biology have general laws that apply to all levels of biological organisation, across all evolutionary time? In their book “Biology’s first law: the tendency for diversity and complexity to increase in evolutionary systems” (2010), Daniel McShea and Robert Brandon propose that the most fundamental law of biology is that all levels of biological organisation have an underlying tendency to become more complex and diverse over time. A range of processes, most notably selection, can prevent the expression of this tendency, (...) but they predict that, on average, we should see that lineages tend toward greater diversity and complexity, driven by fundamentally neutral processes. Their hypothesis can be summarised as “diversity is easy, stasis is hard”. Here, I consider evidence for this “zero force evolutionary law”. It provides a fair description of evolutionary change at the genomic level, but the predictions of the proposed law are not met for broad scale patterns in the evolution of the animal kingdom. (shrink)

One of the major benefits of interdisciplinary research is the chance to swap tools between fields, to save having to reinvent the wheel. The fields of language evolution and evolutionary biology have been swapping tools for centuries to the enrichment of both. Here I will discuss three categories of tool swapping: conceptual tools, where analogies are drawn between hypotheses, patterns or processes, so that one field can take advantage of the path cut through the intellectual jungle by the other; theoretical (...) tools, where the machinery developed to process the data in one field is adapted to be applied to the data of the other; and analytical tools, where common problems encountered in both fields can be solved using useful tricks developed by one or the other. I will argue that conceptual tools borrowed from linguistics contributed to the Darwinian revolution in biology; that theoretical tools of evolutionary change can in some cases be applied to both genetic and linguistic data without having to assume the underlying evolutionary processes are exactly the same; and that there are practical problems that have long been recognised in historical linguistics that may be solved by borrowing some useful analytical tools from evolutionary biology. (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)

I argue that Grafen’s formal darwinism project could profitably incorporate a gene’s-eye view, as informed by the major transitions framework. In this, instead of the individual being assumed to maximise its inclusive fitness, genes are assumed to maximise their inclusive fitness. Maximisation of fitness at the individual level is not a straightforward concept because the major transitions framework shows that there are several kinds of biological individual. In addition, individuals have a definable fitness, exhibit individual-level adaptations and arise in a (...) major transition, only to the extent that the inclusive-fitness interests of genes within them coincide. Therefore, as others have suggested, the fundamental level at which fitness is maximised is the gene level. Previous reconciliations of the concepts of gene-level fitness and individual-level fitness implicitly recognise this point. Adaptations always maximise the fitness of their causative genes, but may be simple or complex. Simple adaptations may be controlled by single genes and be maladaptive at higher levels, whereas complex adaptations are controlled by multiple genes and rely on those genes having coinciding fitness interests at a higher level, for a given trait. (shrink)

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. (shrink)

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

For evolution by natural selection to occur it is classically admitted that the three ingredients of variation, difference in fitness and heredity are necessary and sufficient. In this paper, I show using simple individual-based models, that evolution by natural selection can occur in populations of entities in which neither heredity nor reproduction are present. Furthermore, I demonstrate by complexifying these models that both reproduction and heredity are predictable Darwinian products (i.e. complex adaptations) of populations initially lacking these two properties but (...) in which new variation is introduced via mutations. Later on, I show that replicators are not necessary for evolution by natural selection, but rather the ultimate product of such processes of adaptation. Finally, I assess the value of these models in three relevant domains for Darwinian evolution. (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.

This article illustrates how axiomatic social choice theory can be used in the evaluation of measures of group fitness for a biological hierarchy, thereby contributing to the dialogue between the philosophy of biology and social choice theory. It provides an axiomatic characterization of the ordering underlying the MichodSolariNedelcu index of group fitness for a multicellular organism. The MVSHN index has been used to analyse the germ-soma specialization and the fitness decoupling between the cell and organism levels that takes place during (...) the evolutionary transition to multicellularity. It is argued that some of the axioms satisfied by the MVSHN group fitness ordering are not appropriate for all stages in this transition. (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)

Despite a revived interest in explaining the evolution of anisogamy in recent years (i.e. different—micro and macrogametes), there remain more questions than answers. The topic is important because it is thought to be the foundation of the theory of gender differences and relations. Twelve of these questions are briefly reviewed here—(1) the distinction between sex and sexual types; (2) the distinction between mating types and anisogamy; (3) the possible role of ecological as well as social evolution in proto-gender differences and (...) relations; (4) the life history strategies involved; (5) whether the relevant social relationships are based on conflict or cooperation; (6) the origin and properties of meiosis; (7) the conformity of theories with sex allocation theory; (8) the relevance of multiple levels of selection; (9) appropriate modelling strategies; (10) the relationship between the kind of gametes produced and secondary sexual characteristics; (11) the relevance to humans; and (12) how realistic the search for a single explanation is. It is concluded that polarized sexuality may be a form of trade. (shrink)

Implicit God-like and ghost-in-the-machine metaphors underlie much current thinking about genomes. Although many criticisms of such views exist, none have succeeded in substituting a different, widely accepted view. Viewing the genome with its protein packaging as a brain gets rid of Gods and ghosts while plausibly integrating machine and information-based views. While the ‘wetware’ of brains and genomes are very different, many fundamental principles of how they function are similar. Eukaryotic cells are compound entities in which case the nuclear genome (...) might best be thought of more as a government than simply as a brain. (shrink)

Interest in the most general theory of the evolutionary ecology of life histories ever proposed—that of density-independent versus density-dependent selection—peaked in the 1980s and 1990s respectively but declined somewhat thereafter because of arguments over its theoretical coherence and some empirical failures; although as is shown here, not so much as is commonly supposed. In this article, some conceptual analyses and clarification of terms in the conditions for its application, and in the explanantia of the theory are presented. These make intelligible (...) plausible mechanisms linking the explanantia and the explananda of the theory that are both theoretically coherent and clarify what predictions it does not and does make including about senescence. (shrink)

Over the past two decades, Ginsburg and Jablonka have developed a novel approach to studying the evolutionary origins of consciousness: the Unlimited Associative Learning framework. The central idea is that there is a distinctive type of learning that can serve as a transition marker for the evolutionary transition from non-conscious to conscious life. The goal of this paper is to stimulate discussion of the framework by providing a primer on its key claims and a clear statement of its main empirical (...) predictions. (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)

Inclusive fitness theory was not originally designed to explain the major transitions in evolution, but there is a growing consensus that it has the resources to do so. My aim in this paper is to highlight, in a constructive spirit, the puzzles and challenges that remain. I first consider the distinctive aspects of the cooperative interactions we see within the most complex social groups in nature: multicellular organisms and eusocial insect colonies. I then focus on one aspect in particular: the (...) extreme redundancy these societies exhibit. I argue that extreme redundancy poses a distinctive explanatory puzzle for inclusive fitness theory, and I offer a potential solution which casts coercion as the key enabler. I suggest that the general moral to draw from the case is one of guarded optimism: while inclusive fitness is a powerful tool for understanding evolutionary transitions, it must be integrated within a broader framework that recognizes the distinctive problems such transitions present and the distinctive mechanisms by which these problems may be overcome. (shrink)

The variation and selection form of explanationcan be prescinded from the evolutionary biologyhome ground in which it was discovered and forwhich it has been most developed. When this isdone, variation and selection explanations arefound to have potential application to a widerange of phenomena, far beyond the classicalbiological ground and the contemporaryextensions into epistemological domains. Itappears as the form of explanation most suitedto phenomena of fit. It is also found toparticipate in multiple interestingrelationships with other forms of explanation. We proceed with (...) an examination of multiplekinds of phenomena, interrelationships withother members of the family of forms ofexplanation, and some novel applications evenwithin the home ground of evolutionary biology. (shrink)

Response to commentaries on “The Transmission Sense of Information” Content Type Journal Article Pages 195-200 DOI 10.1007/s10539-011-9257-3 Authors Carl T. Bergstrom, Department of Biology, University of Washington, Seattle, WA 98195-1800, USA Martin Rosvall, Integrated Science Lab, Department of Physics, Umeå University, 901 87 Umeå, Sweden Journal Biology and Philosophy Online ISSN 1572-8404 Print ISSN 0169-3867 Journal Volume Volume 26 Journal Issue Volume 26, Number 2.

In this philosophical paper, I discuss and illustrate the necessary three ingredients that together could allow a collective phenomenon to be labelled as “emergent.” First, the phenomenon, as usual, requires a group of natural objects entering in a non-linear relationship and potentially entailing the existence of various semantic descriptions depending on the human scale of observation. Second, this phenomenon has to be observed by a mechanical observer instead of a human one, which has the natural capacity for temporal or spatial (...) integration, or both. Finally, for this natural observer to detect and select the collective phenomenon, it needs to do so on account of the adaptive advantage this phenomenon is responsible for. The necessity for such a teleological characterization and the presence of natural selection drive us to defend, with many authors, the idea that emergent phenomena should belong only to biology. Following a brief philosophical plea, we present a simple and illustrative computer thought experiment in which a society of agents evolves a stigmergic collective behavior as an outcome of its greater adaptive value. The three ingredients are illustrated and discussed within this experimental context. Such an inclusion of the mechanical observer and the natural selection to which this phenomenon is submitted should underlie the necessary de-subjectivation that strengthens any scientific endeavor. I shall finally show why the short paths taken by ant colonies, the collective flying of birds and the maximum consumption of nutrients by a cellular metabolism are strongly emergent. (shrink)

Communication is an important feature of the living world that mainstream biology fails to adequately deal with. Applying two main disciplines can be contemplated to fill in this gap: semiotics and information theory. Semiotics is a philosophical discipline mainly concerned with meaning; applying it to life already originated in biosemiotics. Information theory is a mathematical discipline coming from engineering which has literal communication as purpose. Biosemiotics and information theory are thus concerned with distinct and complementary possible meanings of the word (...) ‘communication’. Since literal communication needs to be secured so as to enable semantics being communicated, information theory is a necessary prerequisite to biosemiotics. Moreover, heredity is a purely literal communication process of capital importance fully relevant to literal communication, hence to information theory. A short introduction to discrete information theory is proposed, which is centred on the concept of redundancy and its use in order to make sequences resilient to errors. Information theory has been an extremely active and fruitful domain of researches and the motor of the tremendous progress of communication engineering in the last decades. Its possible connections with semantics and linguistics are briefly considered. Its applications to biology are suggested especially as regards error-correcting codes which are mandatory for securing the conservation of genomes. Biology needs information theory so biologists and communication engineers should closely collaborate. (shrink)