We review Evolution and the Levels of Selection by Samir Okasha. This important book provides a cohesive philosophical framework for understanding levels-of-selections problems in biology. Concerning evolutionary transitions, Okasha proposes that three stages characterize the shift from a lower level of selection to a higher one. We discuss the application of Okasha’s three-stage concept to the evolutionary transition from unicellularity to multicellularity in the volvocine green algae. Okasha’s concepts are a provocative step towards a more general understanding of the major (...) evolutionary transitions; however, the application of certain ideas to the volvocine model system is not straightforward. (shrink)

Understanding good design requires addressing the question of what units undergo natural selection, thereby becoming adapted. There is, therefore, a natural connection between the formal Darwinism project (which aims to connect population genetics with the evolution of design and fitness maximization) and levels of selection issues. We argue that the formal Darwinism project offers contradictory and confusing lines of thinking concerning level(s) of selection. The project favors multicellular organisms over both the lower (cell) and higher (social group) levels as the (...) level of adaptation. Grafen offers four reasons for giving such special status to multicellular organisms: (1) they lack appreciable within-organism cell selection, (2) they have multiple features that appear contrived for the same purpose, (3) they possess a set of phenotypes, and (4) they leave offspring according to their phenotypes. We discuss why these rationales are not compelling and suggest that a more even-handed approach, in which multicellular organisms are not assumed to have special status, would be desirable for a project that aims to make progress on the foundations of evolutionary theory. (shrink)

Recent theoretical work has identified a tightly-constrained sense in which genes carry representational content. Representational properties of the genome are founded in the transmission of DNA over phylogenetic time and its role in natural selection. However, genetic representation is not just relevant to questions of selection and evolution. This paper goes beyond existing treatments and argues for the heterodox view that information generated by a process of selection over phylogenetic time can be read in ontogenetic time, in the course of (...) individual development. Recent results in evolutionary biology, drawn both from modelling work, and from experimental and observational data, support a role for genetic representation in explaining individual ontogeny: both genetic representations and environmental information are read by the mechanisms of development, in an individual, so as to lead to adaptive phenotypes. Furthermore, in some cases there appears to have been selection between individuals that rely to different degrees on the two sources of information. Thus, the theory of representation in inheritance systems like the genome is much more than just a coherent reconstruction of information talk in biology. Genetic representation is a property with considerable explanatory utility. (shrink)

The concept of innateness is used to make inferences between various better-understood properties, like developmental canalization, evolutionary adaptation, heritability, species-typicality, and so on (‘innateness-related properties’). This article uses a recently-developed account of the representational content carried by inheritance systems like the genome to explain why innateness-related properties cluster together, especially in non-human organisms. Although inferences between innateness-related properties are deductively invalid, and lead to false conclusions in many actual cases, where some aspect of a phenotypic trait develops in reliance on (...) a genetic representation it will tend, better than chance, to have many of the innateness-related properties. The account also shows why inferences between innateness-related properties sometimes fail and argues that such inferences are especially misleading when applied to human psychology and behaviour because human psychological development is especially reliant on non-genetic inherited representations. (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)

How all philosophical explanations of human consciousness and the fundamental structure of the cosmos are bizarre—and why that’s a good thing Do we live inside a simulated reality or a pocket universe embedded in a larger structure about which we know virtually nothing? Is consciousness a purely physical matter, or might it require something extra, something nonphysical? According to the philosopher Eric Schwitzgebel, it’s hard to say. In The Weirdness of the World, Schwitzgebel argues that the answers to these fundamental (...) questions lie beyond our powers of comprehension. We can be certain only that the truth—whatever it is—is weird. Philosophy, he proposes, can aim to open—to reveal possibilities we had not previously appreciated—or to close, to narrow down to the one correct theory of the phenomenon in question. Schwitzgebel argues for a philosophy that opens. According to Schwitzgebel’s “Universal Bizarreness” thesis, every possible theory of the relation of mind and cosmos defies common sense. According to his complementary “Universal Dubiety” thesis, no general theory of the relationship between mind and cosmos compels rational belief. Might the United States be a conscious organism—a conscious group mind with approximately the intelligence of a rabbit? Might virtually every action we perform cause virtually every possible type of future event, echoing down through the infinite future of an infinite universe? What, if anything, is it like to be a garden snail? Schwitzgebel makes a persuasive case for the thrill of considering the most bizarre philosophical possibilities. (shrink)

Two major works in recent evolutionary biology have in different ways touched upon the issue of cultural replicators in language, namely Dawkins’ Selfish Gene and Maynard Smith and Szathmáry’s Major Transitions in Evolution. In the latter, the emergence of language is referred to as the last major transition in evolution, a claim we argue to be derived from a crucial property of language, called Duality of Patterning. Prima facie, this property makes natural language look like a structural equivalent to DNA, (...) and its peer in terms of expressive power. We will argue that, if one takes seriously Maynard Smith and Szathmáry’s outlook and examines what has been proposed as linguistic replicators, amongst others phonemes and words, the analogy meme-gene becomes problematic. A key issue is the fact that genes and memes are assumed to carry and transmit information, while what has been described as the best candidate for replicatorhood in language, i.e. the phoneme, does by definition not carry meaning. We will argue that semiotic systems with Duality of Pattering force us to reconsider either the analogy between replicators in the biological and the cultural domain, or what it is to be a replicator in linguistics. (shrink)

If you’re a materialist, you probably think that rabbits are conscious. And you ought to think that. After all, rabbits are a lot like us, biologically and neurophysiologically. If you’re a materialist, you probably also think that conscious experience would be present in a wide range of naturally-evolved alien beings behaviorally very similar to us even if they are physiologically very different. And you ought to think that. After all, to deny it seems insupportable Earthly chauvinism. But a materialist who (...) accepts consciousness in weirdly formed aliens ought also to accept consciousness in spatially distributed group entities. If she then also accepts rabbit consciousness, she ought to accept the possibility of consciousness even in rather dumb group entities. Finally, the United States would seem to be a rather dumb group entity of the relevant sort. If we set aside our morphological prejudices against spatially distributed group entities, we can see that the United States has all the types of properties that materialists tend to regard as characteristic of conscious beings. (shrink)

In contrast to vertical gene transfer from parent to offspring, horizontal (or lateral) gene transfer moves genetic information between different species. Bacteria and archaea often adapt through horizontal gene transfer. Recent analyses indicate that eukaryotic genomes, too, have acquired numerous genes via horizontal transfer from prokaryotes and other lineages. Based on this we raise the hypothesis that horizontally acquired genes may have contributed more to adaptive evolution of eukaryotes than previously assumed. Current candidate sets of horizontally acquired eukaryotic genes may (...) just be the tip of an iceberg. We have recently shown that adaptation of the thermoacidophilic red algaGaldieria sulphurariato its hot, acid, toxic‐metal laden, volcanic environment was facilitated by the acquisition of numerous genes from extremophile bacteria and archaea. Other recently published examples of horizontal acquisitions involved in adaptation include ice‐binding proteins in marine algae, enzymes for carotenoid biosynthesis in aphids, and genes involved in fungal metabolism.Editor's suggested further reading in BioEssaysJumping the fine LINE between species: Horizontal transfer of transposable elements in animals catalyses genome evolution Abstract. (shrink)

A recent trend in economics has been to appeal to evolutionary theory when addressing various open questions in the subject. I here further investigate one particular such appeal to evolutionary biology: the argument that, since markets select firms as coherent units, firms should be seen to be genuine economic agents. To assess this argument, I present a model of firm/office selection in a competitive market, and show that there are cases where markets can select for firms/offices as collective units – (...) and thus, as agents of their own – but also that there are cases where they do not. In this way, I try to make the evolutionary argument for the agency-based view of the firm more precise. (shrink)

In this paper, I consider the recent resurgence of “evolutionary economics”—the idea that evolutionary theory can be very useful to push forward key debates in economics—and assess the extent to which it rests on a plausible foundation. To do this, I first distinguish two ways in which evolutionary theory can, in principle, be brought to bear on an economic problem—namely, evidentially and heuristically—and then apply this distinction to the three major hypotheses that evolutionary economists have come to defend: the implausibility (...) of rational choice theory as an account of economic rationality, the idea that firms are autonomous economic agents, and the need for a more dynamic, less equilibrium-focused economic methodology. In each of these cases, I conclude negatively: the relevant evolutionary considerations neither suggest interesting and novel hypotheses to investigate further nor are backed up by the needed data to constitute genuine evidence. I end by distinguishing this criticism of evolutionary economics from others that have been put forward in the literature: in particular, I make clear that, unlike those of other critics, the arguments of this paper are based on epistemic—not structural—considerations and therefore leave more room for a plausible form of evolutionary economics to come about in the future. (shrink)

Carrie Figdor's Pieces of mind lays the groundwork for critiquing the mind package view of minds. According to the mind package view, psychological properties travel in groups, such that an entity either has the whole mind package or lacks mentality altogether. Implicit commitment to the mind package view makes it seem absurd to attribute some psychological properties (e.g., preferences) to entities that lack other psychological properties (e.g., feelings). Contra the mind package view, we are psychologically continuous with plants, worms, and (...) bacteria: Our patterns of mindedness resemble theirs, even if such entities do not have the whole mind package. (shrink)

The question of whether “genetic information” is a merely causal factor in development or can be made sense of semantically, in a way analogous to a language or other type of representation, has generated a long debate in the philosophy of biology. It is intimately connected with another intense debate, concerning the limits of genetic determinism. In this paper I argue that widespread attempts to draw analogies between genetic information and information contained in books, blueprints or computer programs, are fundamentally (...) inadequate. In development, gene exons are the central part of an intricate and densely ramified semantic Genetic Informational Network. DNA in the entire genome is in a state of continuous positive and negative feedback with itself and with its ‘environment’, and is ‘read’ and acted upon by the cell in various alternative and complementary ways. The linear combinatorial coding relation between codons and amino acids is but one aspect of semantic genetic information, which is, when considered in its entirety, a far wider and richer concept. (shrink)

Maynard Smith’s defenses of adaptationism and of the value of optimization theory in evolutionary biology are both criticized. His defense does not adequately respond to the criticism of adaptationism by Gould and Lewontin. It is also argued here that natural selection cannot be interpreted as an optimization process if the objective function to be optimized is either (i) interpretable as a fitness, or (ii) correlated with the mean population fitness. This result holds even if fitnesses are frequency-independent; the problem is (...) further exacerbated in the frequency-dependent context modeled by evolutionary game theory. However, Eshel and Feldman’s new results on “long-term” evolution may provide some hope for the continuing relevance of the game-theoretic framework. These arguments also demonstrate the irrelevance of attempts by Intelligent Design creationists to use computational limits on optimization algorithms as evidence against evolutionary theory. It is pointed out that adaptation, natural selection, and optimization are not equivalent processes in the context of biological evolution. (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)

In this paper we review and argue for the relevance of the concept of open-ended evolution in biological theory. Defining it as a process in which a set of chemical systems bring about an unlimited variety of equivalent systems that are not subject to any pre-determined upper bound of organizational complexity, we explain why only a special type of self-constructing, autonomous systems can actually implement it. We further argue that this capacity derives from the ‘dynamic decoupling’ (in its minimal or (...) most basic sense: the phenotype–genotype decoupling) by means of which a radically new way of material organization (minimal living organization) is achieved, allowing for the long-term sustenance of systems whose individual-metabolic and collective-historical pathways become thereafter deeply intertwined. (shrink)

Simon Conway Morris, noted Cambridge University paleontologist, argues that in evolutionary natural history humans (or beings rather like humans) are an inevitable outcome of the developing speciating processes over millennia; humans are “inherent” in the system. This claim, in marked contrast to claims about contingency made by other prominent paleontologists, is based on numerous remarkable convergences—similar trends found repeatedly in evolutionary history. Conway Morris concludes approaching a natural theology. His argument is powerful and informed. But does it face adequately the (...) surprising events in such history, particularly notable in unexpected co‐options that redirect the course of life? The challenge to understand how humans are both on a continuum with other species and also utterly different remains a central puzzle in paleontology. (shrink)

. Despite the classical prohibition of moving from fact to value, encounter with the biodiversity and plenitude of being in evolutionary natural history moves us to respect life, even to reverence it. Darwinian accounts are value-laden and necessary for understanding life at the same time that Darwinian theory fails to provide sufficient cause for the historically developing diversity and increasing complexity on Earth. Earth is a providing ground; matter and energy on Earth support life, but distinctive to life is information (...) coded in the genetic molecules that superintends this matter-energy. Life is generated and regenerated in struggle, persists in its perishing. Such life is also a gift; nature is grace. Biologists and theologians join in celebrating and conserving the genesis on Earth, awed in their encounter with this creativity that characterizes our home planet. (shrink)

More than one researcher is currently proposing that the notion of information become an important element for defining living systems as well as for explaining conditions that make their origins possible. During the pre-biotic era, the type of compounds encountered would mainly have been very simple in nature and might have been immersed in the natural dynamic of the physical world and in processes of self-organization. It is furthermore quite possible that they formed a relationship between and among certain types (...) of processes that here we are specifically proposing as central for the emergence of cell organization. Consequently, an important initial step towards constructing a theory of biological information is to ask ourselves the question: how do biological systems process information? In this way, we will be contributing to the proposals of this paper where we seek to identify general principles that govern biological computing and that deal with biosemiotic approaches as they are defended in naturalistic normative terms. (shrink)

Biological understanding of human cognitive functions is incomplete because of failure to understand the evolution of complex functions and organisms in general. Here, that failure is attributed to an aspect of the standard neo-Darwinian synthesis, namely commitment to evolution by natural selection of genetic programs in stable environments, a position that cannot easily explain the evolution of complexity. When we turn to consider more realistic, highly changeable environments, however, another possibility becomes clearer. An alternative to genetic programs—dubbed “biogrammars”—is proposed here (...) to deal with complex, changing environments and explain evolving complexity from pre-genetic life to human socio-cognitive functions. (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)

Any description of the emergence and evolution of different types of meaning processes (semiosis, sensu C.S.Peirce) in living systems must be supported by a theoretical framework which makes it possible to understand the nature and dynamics of such processes. Here we propose that the emergence of semiosis of different kinds can be understood as resulting from fundamental interactions in a triadically-organized hierarchical process. To grasp these interactions, we develop a model grounded on Stanley Salthe's hierarchical structuralism. This model can be (...) applied to establish, in a general sense, a set of theoretical constraints for explaining the instantiation of different kinds of meaning processes (iconic, indexical, symbolic) in semiotic systems. We use it to model a semiotic process in the immune system, namely, B-cell activation, in order to offer insights into the heuristic role it can play in the development of explanations for specific semiotic processes. (shrink)

The organism is one of the fundamental concepts of biology and has been at the center of many discussions about biological individuality, yet what exactly it is can be confusing. The definition that we find generally useful is that an organism is a unit in which all the subunits have evolved to be highly cooperative, with very little conflict. We focus on how often organisms evolve from two or more formerly independent organisms. Two canonical transitions of this type—replicators clustered in (...) cells and endosymbiotic organelles within host cells—demonstrate the reality of this kind of evolutionary transition and suggest conditions that can favor it. These conditions include co-transmission of the partners across generations and rules that strongly regulate and limit conflict, such as a fair meiosis. Recently, much attention has been given to associations of animals with microbes involved in their nutrition. These range from tight endosymbiotic associations like those between aphids and Buchnera bacteria, to the complex communities in animal intestines. Here, starting with a reflection about identity through time, we consider the distinctions between these kinds of animal–bacteria interactions and describe the criteria by which a few can be considered jointly organismal but most cannot. (shrink)

Biological individuality is a major topic of discussion in biology and philosophy of biology. Recently, several objections have been raised against traditional accounts of biological individuality, including the objections of monism, theory-centrism, ahistoricity, disciplinary isolationism, and the multiplication of conceptual uncertainties. In this introduction, I will examine the current philosophical landscape about biological individuality, and show how the contributions gathered in this special issue address these five objections. Overall, the aim of this issue is to offer a more diverse, unifying, (...) and scientifically informed conception of what a biological individual is. (shrink)

Natural selection is an extremely powerful process – so powerful, in fact, that it is often tempting to deploy it in explaining phenomena as wide-ranging as the persistence of blue eyes, the origins or persistence of religious belief, or, the history of science. One long-standing debate among both critics and advocates of Darwin’s concerns the scope of Darwinian explanations, and how we are to draw the line. Peter Godfrey-Smith’s Darwinian Populations and Natural Selection is a detailed examination of this question. (...) The book explores the criteria for what may count as a “Darwinian population,” by which Godfrey-Smith means, which collections of entities have the capacity to undergo evolution via natural selection (p. 6). Drawing upon his answer to this question, Godfrey-Smith examines and provides his own solution to the following long-standing debates in philosophy of biology: (a) the twin problems of reproduction and individuation of biological entities, (b) the persistent “gene’s eye view,” (c) the levels and units of selection problem, and (d) the evolution of cultural artifacts and behaviors. (shrink)

Talk of a “genetic program” has become almost as common in cell and evolutionary biology as talk of “genetic information”. But what is a genetic program? I understand the claim that an organism’s genome contains a program to mean that its genes not only carry information about which proteins to make, but also about the conditions in which to make them. I argue that the program description, while accurate in some respects, is ultimately misleading and should be abandoned. After that, (...) I sketch an alternative framework which is better suited to capturing the full informational nature of genes. This framework is centered on the notion of a signaling game, as originally developed by David Lewis, but expanded upon considerably by Brian Skyrms in more recent years. On the view I develop, genes turn out to be the producers and consumers of regulatory or developmental information, rather than entities encoding such information. This finding has consequences that link up with a broader debate in the philosophy of biology concerning inheritance systems. I take this to be one form of theoretical payoff that results from applying the signaling games framework to genes. (shrink)

The debate about the levels of selection has been one of the most controversial both in evolutionary biology and in philosophy of science. Okasha’s book makes the sort of contribution that simply will not be able to be ignored by anyone interested in this field for many years to come. However, my interest here is in highlighting some examples of how Okasha goes about discussing his material to suggest that his book is part of an increasingly interesting trend that sees (...) scientists and philosophers coming together to build a broadened concept of “theory” through a combination of standard mathematical treatments and conceptual analyses. Given the often contentious history of the relationship between philosophy and science, such trend cannot but be welcome. (shrink)

The debate about the levels of selection has been one of the most controversial both in evolutionary biology and in philosophy of science. Okasha’s book makes the sort of contribution that simply will not be able to be ignored by anyone interested in this field for many years to come. However, my interest here is in highlighting some examples of how Okasha goes about discussing his material to suggest that his book is part of an increasingly interesting trend that sees (...) scientists and philosophers coming together to build a broadened concept of “theory” through a combination of standard mathematical treatments and conceptual analyses. Given the often contentious history of the relationship between philosophy and science, such trend cannot but be welcome. (shrink)

Microbes often live in association with dense multicellular aggregates, especially biofilms, and the construction of these aggregates typically requires microbial cells to produce public goods, such as enzymes and signaling molecules. Public-goods producers are, in turn, vulnerable to exploitation by free-rider cells that consume the public goods without paying for their production costs. The cell population of a biofilm or other microbial aggregates are expected to pass through bottlenecks due to a wide range of factors, such as antibiotic treatments and (...) dispersal. The goal of this article is to make the case for the relevance of population bottlenecks at shaping the social interactions within microbial aggregates. The effect of bottlenecks on microbial aggregates is complex in that bottlenecks can favor producers under certain circumstances, but not in others. The concept of the Volunteer’s Dilemma from game theory will be used to motivate the hypothesis that this partly occurs because of how bottlenecks alter the risk of being a producer in a microbial aggregate. Finally, the role of bottlenecks in the microbial world impacts key issues in evolutionary biology, including the importance of ecology at shaping social evolution, and the evolution of multicellularity from unicellular ancestors. (shrink)

This first part of a two‐part article illustrates how research in evolutionary biology and psychology illuminates questions arising in philosophy—specifically questions about the origins of severe, systemic aggression that arise in the mimetic theory of René Girard. Part I looks at: (i) how old the systemic practice of severe aggression is, (ii) how much results from humanity's mimetic/social and competitive nature and how much from ecological, resource, and cultural conditions, and (iii) if ecological and cultural conditions are important, might we (...) adapt them toward greater cooperativity today? After briefly reviewing mimetic theory, the article looks at evolutionary psychology and biology, including fossil and archeological evidence. Findings suggest that severe, systemic aggression might be relatively recent and that its occurrence depends on ecological/resource/socioeconomic conditions additional to our mimetic/social nature. Thus, distinguishing the conditions that prod aggression from those that support pro‐social behavior might aid us in structuring society today. (shrink)

Philosophers of biology, along with everyone else, generally perceive life to fall into two broad categories, the microbes and macrobes, and then pay most of their attention to the latter. ‘Macrobe’ is the word we propose for larger life forms, and we use it as part of an argument for microbial equality. We suggest that taking more notice of microbes – the dominant life form on the planet, both now and throughout evolutionary history – will transform some of the philosophy (...) of biology’s standard ideas on ontology, evolution, taxonomy and biodiversity. We set out a number of recent developments in microbiology – including biofilm formation, chemotaxis, quorum sensing and gene transfer – that highlight microbial capacities for cooperation and communication and break down conventional thinking that microbes are solely or primarily single-celled organisms. These insights also bring new perspectives to the levels of selection debate, as well as to discussions of the evolution and nature of multicellularity, and to neo-Darwinian understandings of evolutionary mechanisms. We show how these revisions lead to further complications for microbial classification and the philosophies of systematics and biodiversity. Incorporating microbial insights into the philosophy of biology will challenge many of its assumptions, but also give greater scope and depth to its investigations. (shrink)

Protistology, and evolutionary protistology in particular, is experiencing a golden research era. It is an extended one that can be dated back to the 1970s, which is when the molecular rebirth of microbial phylogeny began in earnest. John Archibald, a professor of evolutionary microbiology at Dalhousie University, focuses on the beautiful story of endosymbiosis in his book, John Archibald, One Plus One Equals One: Symbiosis and the Origin of Complex Life. However, this historical narrative could be treated as synecdochal of (...) how the molecular revolution has changed evolutionary biology forever, and that is how Archibald has structured his book. I will address the encompassing theme of molecular methods in detail, but also pay careful attention to the endosymbiosis thread in its own right. (shrink)

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)

For a number of years, the debate in evolutionary biology over the ’levels of selection’ has attracted intense interest from philosophers of science. The main question concerns the level of the biological hierarchy at which natural selection occurs. Does selection act on organisms, genes, groups, colonies, demes, species, or some combination of these? According to traditional Darwinian theory the answer is the organism -- it is the differential survival and reproduction of individual organisms that drives the evolutionary process. But there (...) are alternative views too, including pluralist views which say that the choice between different levels of selection is often a matter of perspective, not empirical fact. (shrink)

Kin selection and multilevel selection are alternative approaches for studying the evolution of social behaviour, the relation between which has long been a source of controversy. Many recent theorists regard the two approaches as ultimately equivalent, on the grounds that gene frequency change can be correctly expressed using either. However, this shows only that the two are formally equivalent, not that they offer equally good causal representations of the evolutionary process. This article articulates the notion of an ‘adequate causal representation’ (...) using causal graphs, and then seeks to identify circumstances under which kin and multilevel selection do and do not satisfy the test of causal adequacy. 1 Introduction2 The KS and MLS Approaches2.1 The MLS decomposition2.2 The KS decomposition3 Equivalence and Causality4 Two Problem Cases4.1 The non-social trait case4.2 Genotypic selection with meiotic drive5 Casual Adequacy: A Graphical Approach5.1 The basic idea5.2 Graphs with individual and group variables5.3 Cases where KS is causally adequate5.4 Cases where MLS is causally adequate6 Discussion6.1 Relation to previous work. (shrink)

Evolutionary biology is striking for its ability to explain a large and diverse range of empirical phenomena on the basis of a few general theoretical principles. This article offers a philosophical perspective on the way that evolutionary biology has come to achieve such impressive generality, by focusing on “the strategy of endogenization”. This strategy involves devising evolutionary explanations for biological features that were originally part of the background conditions, or scaffolding, against which such explanations take place. Where successful, the strategy (...) moves biology a step closer to the ideal of explaining as much as possible from evolutionary first principles. The strategy of endogenization is illustrated through a series of biological examples, historical and recent, and its philosophical implications are explored. (shrink)

The ‘levels of selection’ question is one of the most fundamental in evolutionary biology, for it arises directly from the logic of Darwinism. As is well-known, the principle of natural selection is entirely abstract; it says that any entities satisfying certain conditions will evolve by natural selection, whatever those entities are. (These conditions are: variability, associated fitness differences, and heritability (cf. Lewontin 1970).) This fact, when combined with the fact that the biological world is hierarchically structured, i.e. smaller biological units (...) are nested within larger ones, gives rise to the levels of selection question. For in principle, entities at many different hierarchical levels, above and below that of the ‘individual organism’, (e.g. gene, chromosome, cell, kin group, colony, lineage, species) can satisfy the requirements for Darwinian evolution. This possibility has long been recognised by biologists, from Darwin himself to contemporary proponents of ‘multi-level selection’; and there exist numerous biological phenomena which suggest that it has actually occurred. (shrink)

In models of multi-level selection, the property of Darwinian fitness is attributed to entities at more than one level of the biological hierarchy, e.g. individuals and groups. However, the relation between individual and group fitness is a controversial matter. Theorists disagree about whether group fitness should always, or ever, be defined as total (or average) individual fitness. This paper tries to shed light on the issue by drawing on work in social choice theory, and pursuing an analogy between fitness and (...) utility. Social choice theorists have long been interested in the relation between individual and social utility, and have identified conditions under which social utility equals total (or average) individual utility. These ideas are used to shed light on the biological problem. (shrink)

Two alternative statistical approaches to modelling multi-level selection in nature, both found in the contemporary biological literature, are contrasted. The simple covariance approach partitions the total selection differential on a phenotypic character into within-group and between-group components, and identifies the change due to group selection with the latter. The contextual approach partitions the total selection differential into different components, using multivariate regression analysis. The two approaches have different implications for the question of what constitutes group selection and what does not. (...) I argue that the contextual approach is theoretically preferable. This has important implications for a number of issues in the philosophical debate about the levels of selection. Introduction Group selection and the covariance formulation of selection The contextual approach A modification of the simple covariance approach Consequences: frameshifting and additivity 5.1 Frameshifting 5.2 Additivity Conclusion. (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)

Taking the lead from complexity theory and complex systems methodology, the article argues that we are engaged in a contradictory process when encountering, analysing and dealing with complexity. We face opposite tendencies that indicate an in-built dynamic between the increase of complexity and its reduction. The increase partly comes through evolution, defined as the transmission of information and partly from the desire for a human-built world that functions more efficiently. The reduction of complexity is due partly to the necessity of (...) minimizing unwanted and unintended consequences of its increase, and partly due to the continued re-alignment between social systems and their environments. The article examines the public debate about human stem cell research, the debates in the 18th century about the free circulation of commodities and opinions, and the attempts to provide answers to the question debated between Einstein and Freud: why war? The article concludes by arguing that the plea for a re-alignment of scientific disciplines will not suffice. Instead, the study of the co-evolution between science and society offers itself as a strategic research site by focusing on the dynamic interplay between the increase of complexity and its reduction. (shrink)