If we are not to conclude that most planets like Earth have evolved life as intelligent as we are, we must presume Earth is not random. This selection effect, however, also implies that the origin of life need not be as easy as the early appearance of life on Earth suggests. If a series of major evolutionary transitions were required to produce intelligent life, selection implies that a subset of these were “critical steps,” with durations that are (...) similarly distributed. The time remaining from now until simple life is no longer possible on Earth must also be similarly distributed. I show how these results provide timing tests to constrain models of critical evolutionary transitions. (shrink)

Adaptations can occur at different hierarchical levels, but it can be difficult to identify the level 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 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 to be maladapted if they were to leave the group and enter a global cell-level population. Thus, as group-specific adaptations evolve, they create selective pressure for increased cohesiveness and individuality of groups. This article suggests that group-specific adaptations could be present in the simplest, earliest branching colonial volvocine species, which do not have distinct specialized cells. The article also makes predictions about the kind of empirical evidence needed to support or refute the hypothesis that a particular trait is a group-level adaptation. (shrink)

The 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)

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

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.

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

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

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

Drawing on recent advances in evolutionary biology, prominent scholars return to the question posed in a pathbreaking book: how evolution itself evolved.

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. (shrink)

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

This paper takes a close look at the role of behavior in the “major transitions” in evolution—events during which inheritance and development, and therefore the process of adaptation by natural selection, are reorganized at a new level of compositional hierarchy—and at the requirements for sufficiently explaining these important events in the history of life. I argue that behavior played a crucial role in driving at least some of the major transitions. Because behavioral interactions can become stably (...) organized in novel ways on timescales faster than the lifetime of an organism, behavior can lead the way into a transition—becoming organized at the new level prior to inheritance and development. It is widely acknowledged that behavioral plasticity can play an important role in evolution; environmental novelty can elicit novel behavior that may feed back on the evolutionary process through niche selection or cultural inheritance, for example (Jablonka and Lamb 2005). I argue here that not just novel behaviors but novel forms of behavioral organization (distributed or hierarchical control that produces functional coherence) can emerge, binding the evolutionary fates of a group of organisms which were previously independent in terms of behavior as well as reproduction, and leading the way into a transition to an aggregative or “higher-level” mode of reproduction. (shrink)

This paper offers a descriptive account of the transition in evolutionary ethics with reference to some major works from ethics, sociobiology, moral psychology, and primatology. The causes and nature of the transition are discussed by making a distinction between traditional and recent trends in evolutionary ethics enabling us to understand the significance of contemporary evolutionary ethics. The study is gradually directed toward a crucial question of ethics that is the place of reason in morality and what (...) evolutionary ethics implies for Kantian ethics in the question of reason. Many advocates of evolutionary ethics and evolutionary moral psychology claim that reason plays no role or plays a very insignificant role in morality. Such assertion often leads to a rejection of Kantian ethics in particular and cognitivism in general. In this backdrop, the scope of reconciling evolutionary ethics with Kantian ethics as proposed by Frederick Rauscher is analyzed. This paper is thus an attempt to expose the significance of evolutionary ethics with reference to the relation of reason and morality. (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)

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

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

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

We extend the comparatively simple processes of group symmetry-breaking in physical systems to groupoid/equivalence class phase transitions characterizing adiabatically, piecewise stationary, information transmission in prebiotic, biological, and social phenomena: High vs. Low probability paths $$\rightarrow$$ Interior and Exterior Interact $$\rightarrow$$ Multiple Interacting Tunable Workspaces Application to nonstationary processes seems possible via generalizations of the symmetry algebra, for example, to semigroupoids. The dynamic probability models explored here can be transformed into statistical tools for the analysis of real-time and other data (...) across a spectrum of important disciplines confronted by biological and other forms of cognition and their dysfunctions. (shrink)

Distributed adaptations are cases in which adaptation is dependent on the population as a whole: the adaptation is conferred by a structural or compositional aspect of the population; the adaptively relevant information cannot be reduced to information possessed by a single individual. Possible examples of human-distributed adaptations are song lines, traditions, trail systems, game drive lanes and systems of water collection and irrigation. Here we discuss the possible role of distributed adaptations in human cultural macro-evolution. Several kinds of human-distributed adaptations (...) are presented, and their evolutionary implications are highlighted. In particular, we discuss the implications of population size, density and bottlenecks on the distributed adaptations that a population may possess and how they in turn would affect the population's resilience to ecological change. We discuss the implications that distributed adaptations may have for human collective action and the possibility that they played a role in colonization of new areas and niches, in seasonal migration, and in setting constraints for minimal inter-population connectivity. (shrink)

ABSTRACTThis article argues that Anglophone works on Chinese democracy have tended to build their analyses on assumptions that tradition is either a premodern phenomenon unrelated to China’s democratization process, a hindrance that should be gotten rid of if China is to democratize, a static phenomenon that cannot but appear antiquated with regard to a dynamic, fast-paced modern China, or an object from which modern agents can freely draw. In order to challenge these assumptions, this article suggests that modernity and democracy (...) were translated into a Chinese milieu already ripe with Gadamerian prejudices: prejudices that not only modified the meaning of modernity and democracy, but also provided the very conditions without which modernity and democracy would not have been meaningful or understood at all. Max Ko-wu Huang’s work can contribute to our understanding of the role played by various traditions in the process of translating democracy during the transitional period of modern Chin... (shrink)

The study of culture and cultural selection from a biological perspective has been hampered by the lack of any firm theoretical basis for how the information for cultural traits is stored and transmitted. In addition, the study of any living system with a decentralized or multi-level information structure has been somewhat restricted due to the focus in genetics on the gene and the particular hereditary structure of multicellular organisms. Here a different perspective is used, one which regards living systems as (...) self-constructing energy users that utilize their genome as a library of information, making the genetic system just another component that adds fitness to the overall integrated unit. In this framework, basic fitness is measured as the ability to gather energy for growth and reproduction, and the fitness of the genetic system is broken down into two aspects: first, the effectiveness in searching for new somatic functional information, and second, the effectiveness in searching for better structures to store and process information. With this more generalized perspective, major evolutionary transitions to higher levels of organization become competitions between different information structures; furthermore the functioning and fitness of cultural systems can be more easily described and compared with other modes of information storage within biological systems. Modern technological societies are self-constructing systems that rely on written (symbolic) information storage and very complex algorithms that effectively search for variation with a high probability of successful selection. These systems are currently competing with traditional organic systems, and this competition constitutes the latest major evolutionary transition. Upon comparison of the energy-gathering potential of symbolic-based systems with DNA-based life, it appears that symbolic systems have a tremendous fitness potential and the current shift to a higher level of selection may be as significant and far-reaching as any of the previous major evolutionary transitions. (shrink)

This research describes and empirically evaluates the application of a business ethics pedagogy informed by neuroscience and evolutionary biology that suggest ethical decisions are made unconsciously and emotionally. Moral Foundation Theory (MFT) provides a framework that considers a range of values individuals rely on for decision-making. This relates to Virtue ethics (VE) that develops intellectual and character virtues, requires emotional development and is thus suitable for guiding business ethics pedagogy. This study focuses on a business ethics course integrating intellectual (...) virtue and character virtue pedagogies, including contemplative and emotional intelligence skills practices. The aim of the course is to develop character, providing future managers the means to enact ethical decision-making through self-awareness of emotions, intrinsic values and translate these into ethical behavior. Pre-Post Emotional Intelligence (EI) and Moral Foundation Questionnaires (MFQ) are used to evaluate the development of students’ character virtues, i.e., increased emotional awareness, emotional regulation and intrinsic values. This paper mainly seeks to understand how to develop ethical behavior, beyond learning didactic definitions of what is ethically right and what is wrong, i.e. focusing on the learning process that can lead to personal ethical judgment and conduct. The related pedagogy integrates transdisciplinary theories and practices, and as such suggests a holistic approach to ethical behavior transformation. The results indicate limited, significant changes in all measures except emotional awareness. This developing field requires more research to identify the contributions of MFT and related pedagogy to students’ ethical development. (shrink)

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

This is a lovely and very useful book. It deals with the emergence of higher and higher level units of evolution, especially regarding what Queller (1997) called “fraternal major transitions.” These are evolutionary transitions where the lower-level units that gang up are genetically alike and, therefore, the initial advantage is likely to come from the economy of scale rather than the complementation of function, as in the case of “egalitarian transitions.” Simple division of labor may (...) arise from simple conditions (such as some cells being inside, others outside, so that even with the same physiological reactivity the responses of the two cell types may differ), but the spectacular forms arise from evolved epigenetic mechanisms (Jablonka and Lamb 2005). (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)

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)

In this commentary we advance Jagiello et al.'s proposal by zooming in on the possible evolutionary origins of the “bifocal stance” that may have enabled a major transition in human cultural evolution, arguing that the evolution of the bifocal stance was driven by an explosion in cultural complexity arising from cooperative foraging, which led to a feedback loop between the ritual and instrumental stances.

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

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

This article tries to shed light on the mystery of human culture. Human beings are the only extant species with cumulative, evolving cultures. Many animal species do have cultural traditions in the form of socially transmitted practices but they typically lack cumulative culture. Why is that? This discrepancy between humans and animals is even more puzzling if one realizes that culture seems highly advantageous. Thanks to their accumulated knowledge and techniques our early ancestors were able to leave their cradle in (...) Africa and swarm out across the planet, thereby adjusting themselves to a whole range of new environments. Without culture this would have been impossible. So we may ask once again: if cumulative culture is so useful, why don’t other animals have it? In order to explain this mystery I won’t appeal to the major transitions in human evolution—like walking upright, crafting stone tools and controlling fire, etc.—because that would be question begging. Instead I try to unearth the mechanisms that caused those evolutionary turning points to occur in the first place. It seems that unlike other animals, humans are predisposed to efficiently acquire, store and transmit cultural information in such ways that our cultures can genuinely evolve. (shrink)

The concept of scaffold is widespread in science and increasingly common in evolutionary biology. While this concept figures in causal explanations, it is not clear what scaffolds are and what role they play in those explanations. Here we present evolutionary scaffolding explanation as a distinct type of explanatory strategy, distinguishing it from other types of evolutionary explanation. By doing so, we clarify the meaning of “scaffold” as a causal concept and its potential contribution to accounts of (...) class='Hi'>evolutionary novelty and major transitions. (shrink)

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

Change is the fundamental idea of evolution. Explaining the extraordinary biological change we see written in the history of genomes and fossil beds is the primary occupation of the evolutionary biologist. Yet it is a surprising fact that for the majority of evolutionary research, we have rarely studied how evolution typically unfolds in nature, in changing ecological environments, over space and time. While ecology played a major role in the eventual acceptance of the population genetic viewpoint of (...) evolution in the synthetic era (circa 1918-1956), it held a lesser role in the development of evolutionary theory until the 1980s, when we began to systematically study the evolutionary dynamics of natural populations in space and time. As a result, early evolutionary theory was initially constructed in an abstract vacuum that was unrepresentative of evolution in nature. The subtle synthesis between ecology with evolutionary biology (eco-evo synthesis) over the past 40 years has progressed our knowledge of natural selection dynamics as they are found in nature, thus revealing how natural selection varies in strength, direction, form, and, more surprisingly, level of biological organization. Natural selection can no longer be reduced to lower levels of biological organization (i.e., individuals, selfish genes) over shorter timescales but should be expanded to include adaptation at higher levels and over longer timescales. Long-term and/or emergent evolutionary phenomena, such as multilevel selection or evolvability, have thus become tenable concepts within an evolutionary biology that embraces ecology and spatiotemporal change. Evolutionary biology is currently suspended at an intermediate stage of scientific progress that calls for the organization of all the recent knowledge revealed by the eco-evo synthesis into a coherent and unified theoretical framework. This is where philosophers of biology can be of particular use, acting as a bridge between the subdisciplines of biology and inventing new theoretical strategies to organize and accommodate the recent knowledge. Philosophers have recommended transitioning away from outdated philosophies that were originally derived from physics within the philosophical zeitgeist of logical positivism (i.e., monism, reductionism, and monocausation) and toward a distinct philosophy of biology that can capture the natural complexity of multifaceted biological systems within diverse ecosystems—one that embraces the emerging philosophies of pluralism, emergence, and multicausality. Therefore, I see recent advances in ecology, evolutionary biology, and the philosophy of biology as laying the groundwork for another major biological synthesis, what I refer to as the Second Synthesis because, in many respects, it is analogous to the aims and outcomes of the first major biological synthesis (but is notably distinct from the inorganic and contrived progressive movement known as the extended evolutionary synthesis). With the general development of a distinctive philosophy of science, biology has rightfully emerged as an autonomous science. Thus, while the first synthesis legitimized biology, the Second Synthesis autonomized biology and afforded biology its own philosophy, allowing biology to finally realize its full scientific potential. (shrink)

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)

What I suggest we can see in this brief overview of the literature is an extensive interpenetration on both sides of these debates between scientific, political, and social values. Important shifts in political and social values were of course occurring over the same period, some of them in parallel with, and perhaps even contributing to, these transitions I have been speaking of in evolutionary discourse. The developments that I think of as at least suggestive of possible parallels include (...) the progressive encroachment of public values into the private domain of post-World War II American life, the cold war, the rise of consumerism, and the flowering of what Christopher Lasch calls a “narcissistic individualism.”35 In popular language, the 1960s gave birth to the “me” generation. Perhaps the most tantalizing analogue is suggested by Barbara Ehrenreich's argument for the emergence of a new meaning of masculinity — an ideal of masculinity measured not by commitment, responsibility, or success as family provider, but precisely by the strength of a man's autonomy in the private sphere, his resistance to the demands of a hampering female.36 It is tempting to speculate about possible connections between changes in scientific discourse and developments in the social and political spheres, but such connections, however suggestive, would clearly have to be demonstrated.For now, however, I want to focus on another kind of change —a transformation not so much in the social or political sphere as in the scientific sphere. I make this turn, or return, in support of a more complex account of scientific change that incorporates reverberations within the scientific communty along with social and political changes.In the 1960s, all of biology was undergoing a major transformation in direct response to the dramatic successes of molecular biology. These successes seemed to completely vindicate the values on which the molecular revolution was premised — namely, simplicity and mechanism. Following the victory of Watson and Crick, and of others after them, the fever of that endeavor swept through biology leaving in its wake a new standard of science, and of scientific discourse — one predicated on clarity, simplicity, and analyzability; on the definition of legitimate questions as those capable of clear and unambiguous answers. Every biological discipline felt it — even evolutionary biology, which in some respects was at the furthest pole. Perhaps precisely because it seemed conceptually so remote, evolutionary biology may have felt it most of all. Lewontin inadvertently provides us with some direct support for this view. Indeed, he begins his introduction to Population Biology and Evolution with the following remarks: The twenty years since World War II have seen a vindication in biology of our faith in the Cartesian method as a way of doing science. Some of the most fundamental and interesting problems of biology have been solved or are very nearly solved by an analytic technique that is now loosely called “molecular biology.” But it is not specifically the “molecular” aspect of biology of the last twenty years that has led to its success. It is, rather, the analytic aspect, the belief that by breaking systems down into their component parts, by simplifying them or using simpler organisms, one can learn about more complex systems. As it happens, the problems that were attacked and are being attacked by this method lead to answers in terms of molecules and cell organelles.... There is a host of problems in biology, however, that has been much neglected in these twenty exciting years, because the answers to them cannot be meaningfully framed in molecular and cellular terms.37Lewontin is referring, of course, to problems in evolution. The remainder of his remarks is devoted to an argument for the applicability of the method, if not the content, of molecular biology to these problems. He writes, “It is not the case that molecular biology is Cartesian and analytic while population biology is holistic. Population biology is properly analytic and operates, within the framework of its own problems, by the process of simplification, analysis, and resynthesis.”38 With these remarks, he leads into the criticism of the “holists” who have “held up progress.”This new ethic of simplicity, clarity, and mechanism — embodying the very virtues lauded by Williams — was explicitly carried into evolutionary biology in the name of scientific progress. As it happened, the values implied also fit conveniently well with other values — each set of values providing crucial support for the other.However substantive the scientific gains may have been in some respects, the net effect of this ethic has also been a systematic “perceptual bias” — a bias with profound practical consequences for the entire program of methodological individualism in evolutionary biology, if not elsewhere as well. It may well be that the whole is equivalent to the reconstituted aggregate of its parts, if, in the process of aggregation or summation, all possible interactions among the parts are included. But if certain kinds of interactions are systematically excluded, our confidence in that program necessarily founders. My claim here is that such systematic exclusion does occur, and that it occurs on a number of different levels. To briefly review the interlocking kinds of “bias” that I see occurring in practice, I suggest the following schematic listing:On the most general level: The ethic of simplicity — the privileging of certain values, even certain methodologies, as having an a priori superior claim to scientific credibility.Only slightly less general, and crucially related, is the equation of “scientific” with “tractible”: Given the techniques of analysis available, the equation of science with what we can do inevitably leads to a systematic technical bias favoring simplicity. That is, because we don't know how to model complex dynamics, nonlinear interactions are systematically biased against because of the limitations of our technical know-how. The consequences of this equation of the scientific with the tractible are greatly compounded by the additional equation between what we can do and what is — that is, by our temptation to confuse tractibility with reality.Finally, and also closely related, a further kind of elision occurs even within the confines of tractibility. This kind of elision — taking the form almost of inferring tractibility from one's prior assumptions of what is real — is exemplified by the history of a mathematical ecology of mutualism. Even when mutualism can be introduced into the same technical machinery, it is still not pursued. The basic assumption is that competition is what is real, not because it is easier to model, but because it is what we expect. When the actual difficulties of modeling competition are then in turn suppressed, as in the Robert May story, what we have, given the temptation to equate the tractible with the real, is the possibility of a truly self-fulfilling prophecy. (shrink)

A large body of experience and expertise on the implementation of sustainable public school food procurement policies has developed in recent years. However, there has been little investigation of the values and motivations of the public officials implementing the policies. To address this gap, we examine how the city of Avignon took a step toward transition to local fresh food procurement for public schools, under French government calls for sustainable food products in public canteens. Our analysis combines the Multi-Level Perspective (...) on sustainable transitions with the Public Service Motivation construct. Unlike other studies addressing the MLP at a macro level, we focus on individual motivations behind public action. We demonstrate that staff motivations have a major impact on how policies supporting transition to local fresh food procurement for public schools are implemented. These public officials’ approach to food shows concern for the public interest regardless of financial return. It is not the rational choice that primarily drives the implementation of urban food policies, but normative conformity and affective bonding. Interestingly, we conclude that the success of Avignon’s politically-driven school food procurement policy is essentially due to the catering and procurement staff who overcame the barriers to its implementation. (shrink)

Here I discuss the basic elements, major stages, and completion of progressive evolution. The cosmic world of self-realization is based on extensive self-development within a closed contour: temporal counter-transitions of spatial counter-elements (energy bonds and media and, basically, substance structures) form of local worlds within it through evolution of informational structures. The organic world of reproduction develops through the open informational path: the initial substance, through energy exchange and metabolism, reproduces similar substance; the latter interacts with the environment (...) and, subsequently, reproduces its like, and so on. The animal world of self-regulation builds up a closed informational contour in the environment through the informational input and command output. The human world of self-cognition forms the intensive type of development within the internal closed informational contour of cognition. Counter-transitions of ideal images and signs relate to their real prototypes. In the course of cognition, abstractive thinking develops and brings man to the possibility of reflection of the initial world in its integrity (thus, elevates man to the infinite, by Hegel). (shrink)

This chapter highlights Carl Woese’s message about cellular complexity. It addresses the phenotypic distance between a putative, hypothetical ur-replicator or ur-chromosome and anything that can function as a cell that can effectively respond to its environment in ways that maintain its metabolic and physiological integrity. It describes what self-preserving and self-extending behavior in a chemical system minimally involves. This chapter shows that The Major Transitions in Evolution assumes that the emergence of a replicating molecular complex coincides with the (...) emergence of structures with self-preserving characteristics. It then speculates a reason why the “missing transition” has not been missed. (shrink)

Darwin provided us with a powerful theoretical framework to explain the evolution of living systems. Natural selection alone, however, has sometimes been seen as insufficient to explain the emergence of new levels of selection. The problem is one of “circularity” for evolutionary explanations: how to explain the origins of Darwinian properties without already invoking their presence at the level they emerge. That is, how does evolution by natural selection commence in the first place? Recent results in experimental evolution suggest (...) a way forward: Paul Rainey and his collaborators have shown that Darwinian properties could be exogenously imposed via what they call “ecological scaffolding.” This could solve the “black box” dilemma faced by Darwinian explanations of new levels of organization. Yet, despite “scaffolding” recently becoming a popular term in the study of cognition, culture, and evolution, the concept has suffered from vagueness and ambiguity. This article aims to show that scaffolding can be turned into a proper scientific concept able to do explanatory work within the context of the major evolutionary transitions. Doing so will allow us to once again extend the scope of the Darwinian model of evolution by natural selection. (shrink)

It is striking that the concept of fitness although fundamental in evolutionary theory, still remains ambiguous. I argue here that time, although usually neglected, is an important parameter in regards to the concept of fitness. I will show some of the benefits of taking it seriously using the example of recent debates over evolutionary transitions in individuality. I start from Okasha's assertion that once an evolutionary transition in individuality is completed an ontologically new level of selection (...) emerges from lower levels of organization. I argue that Okasha's claim to have identified two ontologically distinct levels of selection is an artifact created by an undeserved comparison between the fitness of the collective level and the fitness of its constituents. Once fitness is assessed over the same period of time at the two levels of organization it becomes clear that only one, unique process of selection is acting upon both levels. (shrink)

We appreciate the depth and breadth of comments we received. They reflect the interdisciplinary challenge of our inquiry and reassured us of its broad interest. We believe that our target article and the criticisms, elaborations, and extensions of the commentators can be an important contribution to establishing human ultrasociality as a new field of social science inquiry. A few of the commentators questioned our definition of ultrasociality, and we begin our response with an elaboration of that definition and a defense (...) of our argument that human ultrasociality began with agriculture. We then respond to the second major area of controversy, namely, our use of group selection to explain the economic drivers behind the agricultural transition. We then focus on the issue of human intentionality raised by the phenomenon of collective intelligence. The intriguing question is to what extent can an entire culture change its own destiny? We then address the issue of the division of labor raised by a number of commentators. The complex division of labor was both a driver and a defining characteristic of ultrasociality, even though it was present in simpler forms in earlier societies. The remaining issues addressed include energy and complexity, expansion and sustainability, and the accelerating evolution of human ultrasociality. These were raised by only a few commentators, but their importance warrants further elaboration. (shrink)

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

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

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)