The concept of monophyly is central to much of modern biology. Despite many efforts over many years, important questions remain unanswered that relate both to the concept itself and to its various applications. This essay focuses primarily on four of these: i) Is it possible to define monophyly operationally, specifically with respect to both the structures of genomes and at the levels of the highest phylogenetic categories (kingdoms, phyla, classes)? ii) May the mosaic and chimeric structures of genomes be sufficiently (...) important factors in phylogeny that situations exist in which the concept may not be applicable? iii) In the history of life on earth were there important groups of organisms that probably had polyphyletic, rather than monophyletic, origins? iv) Does the near universal search for monophyletic origins of clades lead, on occasion, to both undesirable narrowing of acceptable options for development of evolutionary scenarios and sometimes actual omission from consideration of less conventional types of both data and modes of thought, possibly at the expense of biological understanding? Three sections in the essay consider possible answers to these questions: i) A reassessment is made of major features of both the concept and some of its applications. Recent research results make it seem improbable that there could have been single basal forms for many of the highest categories of evolutionary differentiation (kingdoms, phyla, classes). The universal tree of life probably had many roots. Facts contributing to this perception include the phylogenetically widespread occurrences of: horizontal transfers of plasmids, viral genomes, and transposons; multiple genomic duplications; the existence and properties of large numbers of gene families and protein families; multiple symbioses; broad-scale hybridizations; and multiple homoplasys. Next, justifications are reassessed for the application of monophyletic frameworks to two major evolutionary developments usually interpreted as having been monophyletic: ii) the origins of life; and iii) the origins of the vertebrate tetrapods. For both cases polyphyletic hypotheses are suggested as more probable than monophyletic hypotheses. Major conclusions are, as answers to the four questions posed above: probably not, yes, yes, and yes. (shrink)

Many aspects of biology, such as population genetics and senescence, are predicated on identifying individuals and generations. Conventional demarcations of individuals and generations, such as physiological autonomy, unicellular bottlenecks, and alternation of generation, are rife with problems. Do physically separated cuttings or plant ramets constitute separate individuals or generations? Are chimaeras one or more individuals? To resolve these problems, Clarke : 321–361, 2012) proposed that individuals are circumscribed by mechanisms that constrain heritable variance in fitness. Simultaneously, Gorelick and Heng : (...) 1088–1098 2011) showed that sex constrains heritable variance Therefore, for eukaryotes, meiosis and karyogamy provide a consistent way to demarcate individuals and generations. Epigenetic reset associated with meiosis and karyogamy rejuvenates the next generation, but not the parent that engaged in the sex act. Wholesale epigenetic resets that probably only occur with meiosis and karyogamy imply that monozygotic twins are two different individuals, but apomictic progeny are diffuse parts of one disaggregated individual. Mitotic heritability circumscribes an individual, whereas meiotic heritability demarcates new individuals and generations. (shrink)

Universal Darwinism provides a methodology to study the evolution of anatomical form and sociocultural behavior that centers on defining the units and levels of selection, and it identifies the conditions whereby natural selection operates. In previous work, I have examined how this selection-focused evolutionary epistemology may be universalized to include theories that associate with an extended synthesis. Applied evolutionary epistemology is a metatheoretical framework that understands any and all kinds of evolution as phenomena where units evolve by mechanisms at levels (...) of an ontological hierarchy; and it provides three heuristics to search for these units, levels and mechanisms. The heuristics are applicable to language and sociocultural evolution, and here, we give an in-depth analysis of how the unit-heuristic can be implemented into language origin and evolution studies. The importance of developing hierarchy theories is also more fully explained. (shrink)

Applied Evolutionary Epistemology is a scientific-philosophical theory that defines evolution as the set of phenomena whereby units evolve at levels of ontological hierarchies by mechanisms and processes. This theory also provides a methodology to study evolution, namely, studying evolution involves identifying the units that evolve, the levels at which they evolve, and the mechanisms and processes whereby they evolve. Identifying units and levels of evolution in turn requires the development of ontological hierarchy theories, and examining mechanisms and processes necessitates theorizing (...) about causality. Together, hierarchy and causality theories explain how biorealities form and diversify with time. This paper analyzes how Applied EE redefines both hierarchy and causality theories in the light of the recent explosion of network approaches to causal reasoning associated with studies on reticulate and macroevolution. Causality theories have often been framed from within a rigid, ladder-like hierarchy theory where the rungs of the ladder represent the different levels, and the elements on the rungs represent the evolving units. Causality then is either defined reductionistically as an upward movement along the strands of a singular hierarchy, or holistically as a downward movement along that same hierarchy. Upward causation theories thereby analyze causal processes in time, i.e. over the course of natural history or phylogenetically, as Darwin and the founders of the Modern Synthesis intended. Downward causation theories analyze causal processes in space, ontogenetically or ecologically, as the current eco-evo-devo schools are evidencing. This work demonstrates how macroevolution and reticulate evolution theories add to the complexity by examining reticulate causal processes in space–time, and the interactional hierarchies that such studies bring forth introduce a new form of causation that is here called reticulate causation. Reticulate causation occurs between units and levels belonging to different as well as to the same ontological hierarchies. This article concludes that beyond recognizing the existence of multiple units, levels, and mechanisms or processes of evolution, also the existence of multiple kinds of evolutionary causation as well as the existence of multiple evolutionary hierarchies needs to be acknowledged. This furthermore implies that evolution is a pluralistic process divisible into different kinds. (shrink)

This special issue for the Journal for General Philosophy of Science is devoted to exploring the impact and many ramifications of current research in evolutionary epistemology. Evolutionary epistemology is an inter- and multidisciplinary area of research that can be divided into two ever-inclusive research avenues. One research avenue expands on the EEM program and investigates the epistemology of evolution. The other research avenue builds on the EET program and researches the evolution of epistemology. Since its conception, EE has developed three (...) schools of thought: adaptationist, non-adaptationist, and applied EE. Although diverse in outlook and theoretical background, these research avenues and schools share the same agenda of understanding how knowledge evolves, and how it relates to the world. In this paper, we first explain wherefrom evolutionary epistemological schools of thought developed, and then we highlight current debates in EE by briefly reviewing the papers that form part of this special issue. (shrink)

The history and theoretical role of the concept of a ``replicator''is discussed, starting with Dawkins' and Hull's classic treatmentsand working forward. I argue that the replicator concept is still auseful one for evolutionary theory, but it should be revised insome ways. The most important revision is the recognition that notall processes of evolution by natural selection require thatsomething play the role of a replicator.

Birch, Ginsburg, and Jablonka suggest that Unlimited Associative Learning is a “transition marker” in the evolutionary process that produced consciousness, and organizes research by tying together a range of “hallmarks” of consciousness. I argue that the features they recognize as “hallmarks” are indeed important in the evolution of consciousness, but UAL may have a more limited role.

Group-structured and neighbor-structured populations are compared, especially in relation to multilevel selection theory and evolutionary transitions. I argue that purely neighborstructured populations, which can feature the evolution of altruism, are not properly described in multilevel terms. The ability to “gestalt switch” between individualist and multilevel frameworks is then linked to the investigation of “major transitions” in evolution. Some explanatory concepts are naturally linked to one framework or the other, but a full understanding is best achieved via the use of both.

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)

Formal methods developed for modeling levels of selection problems have recently been applied to the investigation of major evolutionary transitions. We discuss two new tools of this kind. First, the ‘near-variant test’ can be used to compare the causal adequacy of predictively equivalent representations. Second, ‘state-variable gestalt-switching’ can be used to gain a useful dual perspective on evolutionary processes that involve both higher and lower level populations.

Inclusive fitness theory provides conditions for the evolutionary success of a gene. These conditions ensure that the gene is selfish in the sense of Dawkins (The selfish gene, Oxford University Press, Oxford, 1976): genes do not and cannot sacrifice their own fitness on behalf of the reproductive population. Therefore, while natural selection explains the appearance of design in the living world (Dawkins in The blind watchmaker: why the evidence of evolution reveals a universe without design, W. W. Norton, New York, (...) 1996), inclusive fitness theory does not explain how. Indeed, Hamilton’s rule is equally compatible with the evolutionary success of prosocial altruistic genes and antisocial predatory genes, whereas only the former, which account for the appearance of design, predominate in successful organisms. Inclusive fitness theory, however, permits a formulation of the central problem of sociobiology in a particularly poignant form: how do interactions among loci induce utterly selfish genes to collaborate, or to predispose their carriers to collaborate, in promoting the fitness of their carriers? Inclusive fitness theory, because it abstracts from synergistic interactions among loci, does not answer this question. Fitness-enhancing collaboration among loci in the genome of a reproductive population requires suppressing alleles that decrease, and promoting alleles that increase the fitness of its carriers. Suppression and promotion are effected by regulatory networks of genes, each of which is itself utterly selfish. This implies that genes, and a fortiori individuals in a social species, do not maximize inclusive fitness but rather interact strategically in complex ways. It is the task of sociobiology to model these complex interactions. (shrink)

Kerr and Godfrey-Smith argue that two mathematically equivalent, alternative formal representations drawn from population genetics, the contextualist and collectivist formalisms, may be equally good for quantifying the dynamics of some natural systems, despite important differences between the formalisms. I draw on constraints on causal representation from Woodward (Making things happen, Oxford University Press, New York, 2003) and Eberhardt and Scheines (Philos Sci 74(5):981–995, 2006) to argue that one or the other formalism will be superior for arbitrary natural systems in which (...) individuals form different types of groups. (shrink)

The σ profile is presented as a tool to analyze the organization of systems at different scales, and how this organization changes in time. Describing structures at different scales as goal‐oriented agents, one can define σ ∈ [0,1] (satisfaction) as the degree to which the goals of each agent at each scale have been met. σ reflects the organization degree at that scale. The σ profile of a system shows the satisfaction at different scales, with the possibility to study their (...) dependencies and evolution. It can also be used to extend game theoretic models. The description of a general tendency on the evolution of complexity and cooperation naturally follows from the σ profile. Experiments on a virtual ecosystem are used as illustration. (shrink)

Darwin’s Conjecture is a bold attempt to bring evolutionary explanation to the social sciences, particularly economics. The book outlines the history of Darwinian explanation in social science then puts forward a generalized replicator account of social evolution by natural selection. The authors identify habits and routines as examples of the generative replicators necessary in order that social evolution is Darwinian. This reviewer notes that the replicator approach limits the generality of this account and suggests that habits and routines might better (...) be seen as members of a Darwinian population in the tradition of Godfrey-Smith’s (2009) generalized account of natural selection instead. Furthermore, the existence of levels of social evolution will constrain what evolution can occur low in the hierarchy. (shrink)

Somatic mutation plays a key role in transforming normal cells into cancerous cells. The analysis of cancer progression therefore requires the study of how point mutations and chromosomal mutations accumulate in cellular lineages. The spread of somatic mutations depends on the mutation rate, the number of cell divisions in the history of a cellular lineage, and the nature of competition between different cellular lineages. We consider how various aspects of tissue architecture and cellular competition affect the pace of mutation accumulation. (...) We also discuss the rise and fall of somatic mutation rates during cancer progression. BioEssays 26:291–299, 2004. © 2004 Wiley Periodicals, Inc. (shrink)

For many years the evolution of language has been seen as a disreputable topic, mired in fanciful “just so stories” about language origins. However, in the last decade a new synthesis of modern linguistics, cognitive neuroscience and neo-Darwinian evolutionary theory has begun to make important contributions to our understanding of the biology and evolution of language. I review some of this recent progress, focusing on the value of the comparative method, which uses data from animal species to draw inferences about (...) language evolution. Discussing speech first, I show how data concerning a wide variety of species, from monkeys to birds, can increase our understanding of the anatomical and neural mechanisms underlying human spoken language, and how bird and whale song provide insights into the ultimate evolutionary function of language. I discuss the “descended larynx” of humans, a peculiar adaptation for speech that has received much attention in the past, which despite earlier claims is not uniquely human. Then I will turn to the neural mechanisms underlying spoken language, pointing out the difficulties animals apparently experience in perceiving hierarchical structure in sounds, and stressing the importance of vocal imitation in the evolution of a spoken language. Turning to ultimate function, I suggest that communication among kin (especially between parents and offspring) played a crucial but neglected role in driving language evolution. Finally, I briefly discuss phylogeny, discussing hypotheses that offer plausible routes to human language from a non-linguistic chimp-like ancestor. I conclude that comparative data from living animals will be key to developing a richer, more interdisciplinary understanding of our most distinctively human trait: language. (shrink)

When the history of life on earth is viewed as a history of cell division, all of life becomes a single cell lineage. The growth and differentiation of this lineage in reciprocal interaction with its environment can be viewed as a developmental process; hence the evolution of life on earth can also be seen as the development of life on earth. Here, in reviewing this field, some potentially fruitful research directions suggested by this change in perspective are highlighted. Variation and (...) selection become, for example, bidirectional information flows between scales, while the notions of “cooperation” and “competition” become scale relative. The language of communication, inference, and information processing becomes more useful than the language of causation to describe the interactions of both homogeneous and heterogeneous living systems at any scale. Emerging scale‐free theoretical frameworks such as predictive coding and active inference provide conceptual tools for reconceptualizing biology as the study of a unified, multiscale dynamical system. (shrink)

The course of biological evolution is regarded by many authors as an ascending path toward higher levels of variety, complexity and integration. There are similar but partly conflicting accounts of the nature and causes of this ascending course. With the aim of reaching a unified conception I start by summarily reviewing three notable examples. These are, in their latest presentations, those of Hoffmeyer and Stjernfelt 2015, Szathmáry 2015, and Lane 2015a. Comparison of their commonalities and divergences, combined with further reflections, (...) leads to the following views: 1) cooperation between preexistent traits and processes is the primary determinant in the emergence of evolutionary novelty; 2) cooperation is a triadic relation similar to semiosis ; 3) regulation, the key element of both metabolism and replication, results from the cooperation of energetic and semiotic causation; 4) conceptual generalization and concrete generalization are fruits of cooperation. Conceptual generalizations spring from the cooperation of ideas. I believe these realizations supply essential elements for a unified view of the ascent of biological evolution towards higher levels of organization. Some of these conceptions are also applicable to similar ascending trajectories through the stages of cultural and technological evolution. (shrink)

Expanding on the results of previous contributions I advance several hypotheses on the interaction of physical and semiotic processes, both in organisms and in human artifacts. I then proceed to employ these ideas to formulate a general account of evolutionary processes in terms of concrete generalization, where, in analogy with conceptual generalization, novel creations retain antecedent features as special or restricted cases. I argue the following theses: 1) the main point of intersection of physical and semiotic causation is the process (...) of regulation; 2) The broadest form of regulation is a course of actions known as modulation ; 3) Modulation is a universal means of conveyance by which a form is lifted from one supporting vehicle and re-embodied into another replica. These considerations suggest viewing biological evolution as a complex generalization of semiotic evolution. Biological reproduction is then regarded as a complex generalization of sign replication. The latter is a relatively simple affair: the embodied form is external and largely independent of its indifferent supporting medium. Biological reproduction, on the contrary, is an extremely complex dynamical process in which the embodied type is duplicated from within the medium, through the subsidiary internal replication of molecular genetic records. These ideas are developed and illustrated through comparisons between the evolution of organisms and that of human artifacts. (shrink)

The ‘managed-metabolism’ hypothesis suggests that a ‘cooperation barrier’ must be overcome if self-producing chemical organizations are to undergo the transition from non-life to life. This dynamical barrier prevents un-managed autocatalytic networks of molecular species from individuating into complex, cooperative organizations. The barrier arises because molecular species that could otherwise make significant cooperative contributions to the success of an organization will often not be supported within the organization, and because side reactions and other ‘free-riding’ processes will undermine cooperation. As a result, (...) the barrier seriously impedes the emergence of individuality, complex functionality and the transition to life. This barrier is analogous to the cooperation barrier that also impedes the emergence of complex cooperation at all levels of living organization. As has been shown at other levels of organization, the barrier can be overcome comprehensively by appropriate ‘management’. Management implements a system of evolvable constraints that can overcome the cooperation barrier by ensuring that beneficial co-operators are supported within the organization and by suppressing free-riders. In this way, management can control and manipulate the chemical processes of a collectively autocatalytic organization, producing novel processes that serve the interests of the organization as a whole and that could not arise and persist in an un-managed chemical organization. Management self-organizes because it is able to capture some of the benefits that are produced when its interventions promote cooperation, thereby enhancing productivity. Selection will therefore favour the emergence of managers that take over and manage chemical organizations so as to overcome the cooperation barrier. The managed-metabolism hypothesis demonstrates that if management is to overcome the cooperation barrier comprehensively, its interventions must be digitally coded. In this way, the hypothesis accounts for the two-tiered structure of all living cells in which a digitally-coded genetic apparatus manages an analogically-informed metabolism.GraphicalOpen image in new window. (shrink)

The ‘managed-metabolism’ hypothesis suggests that a ‘cooperation barrier’ must be overcome if self-producing chemical organizations are to undergo the transition from non-life to life. This dynamical barrier prevents un-managed autocatalytic networks of molecular species from individuating into complex, cooperative organizations. The barrier arises because molecular species that could otherwise make significant cooperative contributions to the success of an organization will often not be supported within the organization, and because side reactions and other ‘free-riding’ processes will undermine cooperation. As a result, (...) the barrier seriously impedes the emergence of individuality, complex functionality and the transition to life. This barrier is analogous to the cooperation barrier that also impedes the emergence of complex cooperation at all levels of living organization. As has been shown at other levels of organization, the barrier can be overcome comprehensively by appropriate ‘management’. Management implements a system of evolvable constraints that can overcome the cooperation barrier by ensuring that beneficial co-operators are supported within the organization and by suppressing free-riders. In this way, management can control and manipulate the chemical processes of a collectively autocatalytic organization, producing novel processes that serve the interests of the organization as a whole and that could not arise and persist in an un-managed chemical organization. Management self-organizes because it is able to capture some of the benefits that are produced when its interventions promote cooperation, thereby enhancing productivity. Selection will therefore favour the emergence of managers that take over and manage chemical organizations so as to overcome the cooperation barrier. The managed-metabolism hypothesis demonstrates that if management is to overcome the cooperation barrier comprehensively, its interventions must be digitally coded. In this way, the hypothesis accounts for the two-tiered structure of all living cells in which a digitally-coded genetic apparatus manages an analogically-informed metabolism.GraphicalOpen image in new window. (shrink)

Can a human society be constrained in such a way that self-organization will thereafter tend to produce outcomes that advance the goals of the society? Such a society would be self-organizing in the sense that individuals who pursue only their own interests would none-the-less act in the interests of the society as a whole, irrespective of any intention to do so. The paper sketches an agent-based model that identifies the conditions that must be met if such a self-organizing society is (...) to emerge. The model draws heavily on an understanding of how self-organizing societies have emerged repeatedly during the evolution of life on Earth. The model demonstrates that the key enabling requirement for a self-organizing society is ‘consequence-capture’. Broadly this means that all agents in the society must capture sufficient of the beneficial consequences of their actions for the goals of the society. ‘Consequence-capture’ can be organized in a society by appropriate management that suppresses free riders and supports pro-social actions. In human societies these constraints include institutions such as systems of governance and social norms. If a self-organizing society is to emerge, consequence-capture must apply to all agents in the society, including those involved in the establishment and adaptation of institutions. If this is achieved, the result will be a fully self-organizing society in which the interests of all agents are aligned with the interests of the society as a whole. (shrink)

Have the large-scale evolutionary patterns illustrated by the fossil record been driven by fluctuations in environmental opportunity, by biotic factors, or by changes in the types of phenotypic variants available for evolutionary change? Since the Modern Synthesis most evolutionary biologists have maintained that microevolutionary processes carrying on over sufficient time will generate macroevolutionary patterns, with no need for other pattern-generating mechanisms such as punctuated equilibrium or species selection. This view was challenged by paleontologists in the 1970s with proposals that the (...) differential sorting and selection of species and clades, and the effects of biotic crises such as mass extinctions, were important extensions to traditional evolutionary theory. More recently those interested in macroevolution have debated the relative importance of abiotic and biotic factors in driving macroevolutionary patterns and have introduced comparative phylogenetic methods to analyze the rates of change in taxonomic diversity. Applying Peter Godfrey-Smith’s distinction between distributional explanations and explanations focusing on the origin of variation, most macroevolutionary studies have provided distributional explanations of macroevolutionary patterns. Comparative studies of developmental evolution, however, have implicated the origin of variants as a driving macroevolution force. In particular, the repatterning of gene regulatory networks provides new insights into the origins of developmental novelties. This raises the question of whether macroevolution has been pulled by the generation of environmental opportunity, or pushed by the introduction of new morphologies. The contrast between distributional and origination scenarios has implications for understanding evolutionary novelty and innovation and how macroevolutionary process may have evolved over time. (shrink)

The causal closure of physics is usually discussed in a context free way. Here I discuss it in the context of engineering systems and biology, where strong emergence takes place due to a combination of upwards emergence and downwards causation. Firstly, I show that causal closure is strictly limited in terms of spatial interactions because these are cases that are of necessity strongly interacting with the environment. Effective Spatial Closure holds ceteris parabus, and can be violated by Black Swan Events. (...) Secondly, I show that causal closure in the hierarchy of emergence is a strictly interlevel affair, and in the cases of engineering and biology encompasses all levels from the social level to the particle physics level. However Effective Causal Closure can usefully be defined for a restricted set of levels, and one can experimentally determine Effective Theories that hold at each level. This does not however imply those effective theories are causally complete by themselves. In particular, the particle physics level is not causally complete by itself in the contexts of solid state physics, digital computers, or biology. Furthermore Inextricably Intertwined Levels occur in all these contexts. (shrink)

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

This article investigates the rate at different periods of the evolutionary process on Earth. The investigation is based on the author's previous investigations of the growth of evolutionary complexity and on other recent investigations of the rates of scientific evolution and information technology. The measures of the evolutionary rates are given in terms of their doubling times, being several million years for animal evolution, and ranging from a million years to some thousand years for human cultural evolution. After the Galilean (...) scientific revolution the doubling time shows a dramatic drop to about only twenty years and is found to be still shorter for modern information technology. The result shows that the totality of the evolutionary processes on Earth follows a superexponentially increasing rate, characterized by steadily decreasing doubling times. (shrink)

In the present article, a depiction of complexity versus time will be used for the construction of a novel form of a tree of life, called The Pattern of Life, comprising the biological, cultural, and scientific forms of the evolutionary process. This diagram accentuates the implication of the successive modifications of developmental programs, in the cultural and scientific realms coupled to a feedback mechanism that is decisive for the accelerating pace of complexity growth, also suggested to be of support of (...) a cautious prediction of future evolution. (shrink)

In the present paper I develop a model of the evolutionary process associated to the widespread although controversial notion of a prevailing trend of increasing complexity over time. The model builds on a coupling of evolution to individual developmental programs and introduces an integrated view of evolution implying that human culture and science form a continuous extension of organic evolution. It is formed as a mathematical model that has made possible a quantitative estimation in relative terms of the growth of (...) complexity. This estimation is accomplished by means of computer simulations the result of which indicates a strong acceleration of complexity all the way from the appearance of multicellular organisms up to modern man. (shrink)

Developmental systems theory (DST) is a general theoretical perspective on development, heredity and evolution. It is intended to facilitate the study of interactions between the many factors that influence development without reviving `dichotomous' debates over nature or nurture, gene or environment, biology or culture. Several recent papers have addressed the relationship between DST and the thriving new discipline of evolutionary developmental biology (EDB). The contributions to this literature by evolutionary developmental biologists contain three important misunderstandings of DST.

Clade selection is unpopular with philosophers who otherwise accept multilevel selection theory. Clades cannot reproduce, and reproduction is widely thought necessary for evolution by natural selection, especially of complex adaptations. Using microbial evolutionary processes as heuristics, I argue contrariwise, that (1) clade growth (proliferation of contained species) substitutes for clade reproduction in the evolution of complex adaptation, (2) clade-level properties favoring persistence – species richness, dispersal, divergence, and possibly intraclade cooperation – are not collapsible into species-level traits, (3) such properties (...) can be maintained by selection on clades, and (4) clade selection extends the explanatory power of the theory of evolution. (shrink)

Interdisciplinarity is one of the features of modern science, defined as blurring the boundaries of disciplines and overcoming their limitations or excessive specialization by borrowing methods from one discipline into another, integrating different theoretical assumptions, and using the same concepts and terms. Often, theoretical knowledge of one discipline and technological advances of another are combined within an interdisciplinary science, and new branches or disciplines may also emerge. Biosemiotics, a field that arose at the crossroads of biology, semiotics, linguistics, and philosophy, (...) enables scientists to borrow theoretical assumptions from semiotics and extend them to different biological theories. The latter applies especially to extended synthesis, wherein culture is viewed as one of the factors influencing evolution. In the present research, the semiotic system of Ukrainian folk ornament is analyzed through the theory of fractals, key features of which are recursion and self-similarity. As a result, an assumption is made about the fractal structure of culture and social life on a conceptual level. What follows is a discussion of how this assumption can contribute to the multilevel selection theory, one of the foundations of extended synthesis, which employs the concept of self-similarity at all levels of the biological hierarchy. (shrink)

This paper examines three exemplary theories of living organization with respect to their common feature of defining life in terms of metabolic closure: autopoiesis, systems, and chemoton theory. Metabolic closure is broadly understood to denote the property of organized chemical systems that each component necessary for the maintenance of the system is produced from within the system itself, except for an input of energy. It is argued that two of the theories considered—autopoiesis and systems—participate in a hylomorphist pattern of thinking (...) which separates the “form” of the living system from its “matter.” The analysis and critique of hylomorphism found in the work of the philosopher Gilbert Simondon is then applied to these two theories, and on the basis of this critique it is argued that the chemoton model offers a superior theory of minimal life which overcomes many of the problems associated with the other two. Throughout, the relationship between hylomorphism and the understanding of living things as machines is explored. The paper concludes by considering how hylomorphism as a background ontology for theories of life fundamentally influences the way life is defined. (shrink)

Although classical evolutionary theory, i.e., population genetics and the Modern Synthesis, was already implicitly ‘gene-centred’, the organism was, in practice, still generally regarded as the individual unit of which a population is composed. The gene-centred approach to evolution only reached a logical conclusion with the advent of the gene-selectionist or gene’s eye view in the 1960s and 1970s. Whereas classical evolutionary theory can only work with fitness differences between individual organisms, gene-selectionism is capable of working with fitness differences among genes (...) within the same organism and genome. Here, we explore the explanatory potential of ‘intra-organismic’ and ‘intra-genomic’ gene-selectionism, i.e., of a behavioural-ecological ‘gene’s eye view’ on genetic, genomic and organismal evolution. First, we give a general outline of the framework and how it complements the—to some extent—still ‘organism-centred’ approach of classical evolutionary theory. Secondly, we give a more in-depth assessment of its explanatory potential for biological evolution, i.e., for Darwin’s ‘common descent with modification’ or, more specifically, for ‘historical continuity or homology with modular evolutionary change’ as it has been studied by evolutionary developmental biology during the last few decades. In contrast with classical evolutionary theory, evo-devo focuses on ‘within-organism’ developmental processes. Given the capacity of gene-selectionism to adopt an intra-organismal gene’s eye view, we outline the relevance of the latter model for evo-devo. Overall, we aim for the conceptual integration between the gene’s eye view on the one hand, and more organism-centred evolutionary models on the other. (shrink)

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

This essay examines the claim “path dependence entails irreversibility” from the point of view of evolutionary biology. I argue that evolutionary irreversibility possesses many faces, sometimes conflicting with path dependence. I propose an account of path dependence that does not rely on irreversibility and explains why it more naturally coexists with the notion of (contingent) irreversibility developed by the Belgian paleontologist Louis Dollo. However, I argue that we should not conceive of this relationship as necessary.

This article introduces a special issue on mechanisms in language evolution research. It describes processes relevant for the emergence of protolanguage and the transition thereof to modern language. Protolanguage is one of the key terms in the field of language evolution, used to designate a hypothesised intermediate stage in the emergence of language present in extinct hominins: qualitatively different from non-human primate communication in possessing some, but not all, of the features that characterise modern language. Much debate in language evolution (...) focuses on the exact delineation of these features, as well as the means whereby the transitions occurred: first from non-human primate communication systems to protolanguage, and then from protolanguage to modern language. In what follows, we first propose a comprehensive typology of protolanguage debates, taking into account the postulated structural organisation of protolanguage, its functions, and its communicative modality. This makes it possible to show how a specific focus on mechanisms and processes deemed relevant for the emergence of these features allows us to assess the explanatory scope of the existing theories of protolanguage. (shrink)

Traditionally defined as the science of the living, or as the field that beyond anatomical structure and bodily form studies functional organization and behaviour, physiology has long been excluded from evolutionary research. The main reason for this exclusion is that physiology has a presential and futuristic outlook on life, while evolutionary theory is traditionally defined as the study of natural history. In this paper, I re-evaluate these classic science divisions and situate physiology within the history of the evolutionary sciences, as (...) well as within debates on the Extended Evolutionary Synthesis and the need for a Third Way of Evolution. I then briefly point out how evolutionary physiology in particular contributes to research on function, causation, teleonomy, agency and cognition. (shrink)

The theory of evolution by natural selection is, perhaps, the crowning intellectual achievement of the biological sciences. There is, however, considerable debate about which entity or entities are selected and what it is that fits them for that role. This article aims to clarify what is at issue in these debates by identifying four distinct, though often confused, concerns and then identifying how the debates on what constitute the units of selection depend to a significant degree on which of these (...) four questions a thinker regards as central. (shrink)

Levels of organization are structures in nature, usually defined by part-whole relationships, with things at higher levels being composed of things at the next lower level. Typical levels of organization that one finds in the literature include the atomic, molecular, cellular, tissue, organ, organismal, group, population, community, ecosystem, landscape, and biosphere levels. References to levels of organization and related hierarchical depictions of nature are prominent in the life sciences and their philosophical study, and appear not only in introductory textbooks and (...) lectures, but also in cutting-edge research articles and reviews. In philosophy, perennial debates such as reduction, emergence, mechanistic explanation, interdisciplinary relations, natural selection, and many other topics, also rely substantially on the notion. -/- Yet, in spite of the ubiquity of the notion, levels of organization have received little explicit attention in biology or its philosophy. Usually they appear in the background as an implicit conceptual framework that is associated with vague intuitions. Attempts at providing general and broadly applicable definitions of levels of organization have not met wide acceptance. In recent years, several authors have put forward localized and minimalistic accounts of levels, and others have raised doubts about the usefulness of the notion as a whole. -/- There are many kinds of ‘levels’ that one may find in philosophy, science, and everyday life—the term is notoriously ambiguous. Besides levels of organization, there are levels of abstraction, realization, being, analysis, processing, theory, science, complexity, and many others. In this article, the focus will be on levels of organization and debates associated with them, and other kinds of levels will only be discussed when they are relevant to this main topic. (shrink)

The Modern Synthesis (MS) is the current paradigm in evolutionary biology. It was actually built by expanding on the conceptual foundations laid out by its predecessors, Darwinism and neo-Darwinism. For sometime now there has been talk of a new Extended Evolutionary Synthesis (EES), and this article begins to outline why we may need such an extension, and how it may come about. As philosopher Karl Popper has noticed, the current evolutionary theory is a theory of genes, and we still lack (...) a theory of forms. The field began, in fact, as a theory of forms in Darwin’s days, and the major goal that an EES will aim for is a unification of our theories of genes and of forms. This may be achieved through an organic grafting of novel concepts onto the foundational structure of the MS, particularly evolvability, phenotypic plasticity, epigenetic inheritance, complexity theory, and the theory of evolution in highly dimensional adaptive landscapes. (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)