This article reviews the edited volume “Evolvability: A Unifying Concept in Evolutionary Biology?” through biological and philosophical lenses. The book provides diverse angles on evolvability, which is affected by various hierarchical levels, timescales, and types of variation, thus moving beyond a purely genomics perspective. Evolvability is important to biosemiotics because understanding the dynamics of topological genotype spaces could help one better comprehend the phenotypic spaces of meaning, as developmental codes and interrelations can influence the emergence of biological novelty over time. (...) This book is successful in provoking thought on evolvability's role as both a product and a driver of evolutionary innovations and explores the philosophical implications of how evolution has the capacity to generate novel forms of meaning. (shrink)

This dissertation concerns the active role of the organism in evolutionary theory. In particular, it concerns how our conception of the relationship between organism and environment, and the nature of natural selection, influences the causal and explanatory role of organismic activity and behavior in evolutionary explanations. The overarching aim is to argue that the behaviors and activities of organisms can serve both as the explananda (that which is explained) and the explanantia (that which explains) in evolutionary explanations. I attempt to (...) achieve this aim by offering three central arguments. First, that the organism-environment relation is the ontologically basic unit of biology. A common way of conceiving the relationship between organism and environment is as a duality—as two causally independent systems that overlap through interaction. I think this is a mistake. There cannot be organisms without environments and vice versa. They are codependent and codetermined. Second, that natural selection is an ecological process. By this I mean that natural selection acts primarily on organism-environment interactions. It is only in virtue of organisms interacting with their environments that there can be fitness differences amongst individual organisms in a population. The ecological approach to natural selection also entails that the nature of the system(s) of inheritance involved in the reoccurrence of favorable organism-environment interactions is, in principle, immaterial to the process of selection. It only matters for selection that organism-environment interactions actually do reoccur in subsequent generations. Third, that niche construction theory—the most well-developed theoretical and conceptual framework for studying how the activities of organisms influences evolutionary dynamics—can be seen as fully a compatible and integrated part of evolutionary theory. Together, these three arguments constitute an overarching argument. Namely that we both can, and should, take organismic activity and behavior to be crucial explanatory elements in evolutionary theory. Throughout the dissertation I also show how these three arguments have consequences for other debates in the philosophy of evolutionary theory, such as the nature of evolutionary processes, the structure of selection-based explanations, the validity and utility of the proximate-ultimate distinction, and the nature of teleology in evolutionary systems. (shrink)

Resource rationality may explain suboptimal patterns of reasoning; but what of “anti-Bayesian” effects where the mind updates in a direction opposite the one it should? We present two phenomena — belief polarization and the size-weight illusion — that are not obviously explained by performance- or resource-based constraints, nor by the authors’ brief discussion of reference repulsion. Can resource rationality accommodate them?

Philosophical analyses of scientific concepts are legion. However, this literature is replete with methodological errors that have largely gone unnoticed. Five distinct projects are conflated which has led to faulty inferences, ambiguities, and mischaracterizations. There has also been some recent enthusiasm for approaches that attempt to rectify problematic scientific concepts but the motivations for these approaches are questionable. I am hopeful that by bringing these various issues to light that it will lead practitioners to be more explicit about their aims (...) and subsequently result in better analyses. (shrink)

In identifying intrinsic molecular chance and extrinsic adaptive pressures as the only causally relevant factors in the process of evolution, the theoretical perspective of the Modern Synthesis had a major impact on the perceived tenability of an ontology of dispositional properties. However, since the late 1970s, an increasing number of evolutionary biologists have challenged the descriptive and explanatory adequacy of this “chance alone, extrinsic only” understanding of evolutionary change. Because morphological studies of homology, convergence, and teratology have revealed a space (...) of possible forms and phylogenetic trajectories that is considerably more restricted than expected, evo-devo has focused on the causal contribution of intrinsic developmental processes to the course of evolution. Evo-devo’s investigation into the developmental structure of the modality of morphology – including both the possibility and impossibility of organismal form – has led to the utilisation of a number of dispositional concepts that emphasise the tendency of the evolutionary process to change along certain routes. In this sense, and in contrast to the perspective of the Modern Synthesis, evo-devo can be described as a “science of dispositions.” This chapter discusses the recent philosophical literature on dispositional properties in evo-devo, exploring debates about both the metaphysical and epistemological aspects of the central dispositional concepts utilised in contemporary evo-devo and addressing the epistemological question of how dispositional properties challenge existing explanatory models in evolutionary biology. (shrink)

Path-dependence offers a promising way of understanding the role historicity plays in explanation, namely, how the past states of a process can matter in the explanation of a given outcome. The two main existing accounts of path-dependence have sought to present it either in terms of dynamic landscapes or branching trees. However, the notions of landscape and tree both have serious limitations and have been criticized. The framework of causal networks is both more fundamental and more general that that of (...) landscapes and trees. Within this framework, I propose that historicity in networks should be understood as symmetry breaking. History matters when an asymmetric bias towards an outcome emerges in a causal network. This permits a quantitative measure for how path-dependence can occur in degrees, and offers suggestive insights into how historicity is intertwined both with causal structure and complexity. (shrink)

Traditional accounts of the role of learning in evolution have concentrated upon its capacity as a source of fitness to individuals. In this paper I use a case study from invasive species biology—the role of conditioned taste aversion in mitigating the impact of cane toads on the native species of Northern Australia—to highlight a role for learning beyond this—as a source of evolvability to populations. This has two benefits. First, it highlights an otherwise under-appreciated role for learning in evolution that (...) does not rely on social learning as an inheritance channel nor “special” evolutionary processes such as genetic accommodation (both of which many are skeptical about). Second, and more significantly, it makes clear important and interesting parallels between learning and exploratory behaviour in development. These parallels motivate the applicability of results from existing research into learning and learning evolution to our understanding of the evolution of evolvability more generally. (shrink)

Against the neo-Darwinian assumption that genetic factors are the principal source of variation upon which natural selection operates, a phenotype-first hypothesis strikes us as revolutionary because development would seem to constitute an independent source of variability. Richard Watson and his co-authors have argued that developmental memory constitutes one such variety of phenotypic variability. While this version of the phenotype-first hypothesis is especially well-suited for the late metazoan context, where animals have a sufficient history of selection from which to draw, appeals (...) to developmental memory seem less plausible in the evolutionary context of the early metazoans. I provide an interpretation of Stuart Newman’s account of deep metazoan phylogenesis that suggests that spandrels are, in addition to developmental memory, an important reservoir of phenotypic variability. I conclude by arguing that Gerd Müller’s “side-effect hypothesis” is an illuminating generalization of the proposed non-Watsonian version of the phenotype-first hypothesis. (shrink)

Cancer is typically spoken of as a “complex” disease. But, in what sense are cancers “complex”? Is there one sense, or several? What implications does this complexity have – both for how we study, and how we intervene upon cancers? The aim of this paper is first, to clarify the variety of senses in which cancer is spoken of as "complex" in the scientific literature, and second, to discover what explanatory and predictive roles such features play.

Synthesising arguments motivate changes to the conceptual tools, theoretical structure, and evaluatory framework employed in a given scientific domain. Recently, a broad coalition of researchers has put forward a synthesising argument in favour of an Extended Evolutionary Synthesis (‘EES’). Often this synthesising argument is evaluated using a virtue-based approach, which construes the EES as a wholesale alternative to prevailing practice. Here I argue this virtue-based approach is not fit for purpose. Taking the central concept of niche construction as a case (...) study, I show that an agenda-based approach better captures the pragmatic and epistemological goals of the EES synthesising argument and diagnoses areas of empirical disagreement with prevailing practice. (shrink)

While the notion of chance has been central in discussions over the probabilistic nature of natural selection and genetic drift, its role in the production of variants on which populational sampling takes place has received much less philosophical attention. This article discusses the concept of chance in evolution in the light of contemporary work in evo-devo. We distinguish different levels at which randomness and chance can be defined in this context, and argue that recent research on variability and evolvability demands (...) a causal understanding of variational probabilities under which development acquires a creative, rather than a constraining role in evolution. We then provide a propensity interpretation of variational probabilities that solves a conceptual confusion between causal properties, variational probabilities and extant variation present in the literature, and explore some metaphysical consequences that follow from our interpretation, specifically with regards to the nature of developmental types. (shrink)

‘Gouldian arguments’ appeal to the contingency of a scientific domain to establish that domain’s autonomy from some body of theory. For instance, pointing to evolutionary contingency, Stephen Jay Gould suggested that natural selection alone is insufficient to explain life on the macroevolutionary scale. In analysing contingency, philosophers have provided source-independent accounts, understanding how events and processes structure history without attending to the nature of those events and processes. But Gouldian Arguments require source-dependent notions of contingency. An account of contingency is (...) source-dependent when it is indexed to some pattern and some process. Positions like Gould’s do not turn on the mere fact of life’s contingency—that life’s shape could have been different due to its sensitivity to initial conditions, path-dependence or stochasticity. Rather, Gouldian arguments require that the contingency is due to particular kinds of processes: in this case, those which microevolutionary theory cannot account for. This source-dependent perspective clarifies both debates about the nature and importance of contingency, and empirical routes for testing Gould’s thesis. (shrink)

Evolutionary explanations are not only common in the biological sciences, but also widespread outside biology. But an account of how evolutionary explanations perform their explanatory work is still lacking. This paper develops such an account. I argue that available accounts of explanations in evolutionary science miss important parts of the role of history in evolutionary explanations. I argue that the historical part of evolutionary science should be taken as having genuine explanatory force, and that it provides how‐possibly explanations sensu Dray. (...) I propose an account of evolutionary explanations as comparative‐composite explanations consisting of two distinct kinds of explanations, one processual and one historical, that are connected via the explanandum's evolvability to show how the explanandum is the product of its evolutionary past. The account is both a reconstruction of how evolutionary explanations in biology work and a guideline specifying what kind of explanations evolutionary research programs should develop. (shrink)

A disposition or dispositional property is a capacity, ability, or potential to display or exhibit some outcome. Evolvability refers to a disposition to evolve. This chapter discusses why the dispositional nature of evolvability matters—why philosophical distinctions about dispositions can have scientific implications. To that end, we build a conceptual toolkit with vocabulary from prior philosophical analyses using a different disposition: protein foldability. We then apply this toolkit to address several methodological questions related to evolvability. What entities are the bearers of (...) evolvability? What features causally contribute to the disposition of evolvability? How does evolvability manifest? The various possible answers to these questions available from philosophical distinctions suggest key implications for why the concept of evolvability as a disposition is useful in evolutionary research. These include (1) securing scientific virtues (e.g., explanatory depth and generalization, prediction or retrodiction, and control or manipulation) and (2) fostering interdisciplinary collaboration through the coordination of definitional diversity and different types of inquiry. Together these facilitate concentration on a variety of research questions at different levels of organization and on distinct time scales, all of which should be expected for a complex dispositional property such as evolvability. (shrink)

A number of biologists and philosophers have noted the diversity of interpretations of evolvability in contemporary evolutionary research. Different clusters of research defined by co-citation patterns or shared methodological orientation sometimes concentrate on distinct conceptions of evolvability. We examine five different activities where the notion of evolvability plays conceptual roles in evolutionary biological investigation: setting a research agenda, characterization, explanation, prediction, and control. Our analysis of representative examples demonstrates how different conceptual roles of evolvability are quasi-independent and yet exhibit important (...) relationships across scientific activities. It also provides us with the resources to detail two distinct strategies for how evolvability can help to synthesize disparate areas of research and thereby potentially serve as a unifying concept in evolutionary biology. (shrink)

The origins of human social cooperation confound simple evolutionary explanation. But from Darwin and Durkheim onward, theorists (anthropologists and sociologists especially) have posited a potential link with another curious and distinctively human social trait that cries out for explanation: religion. This dissertation explores one contemporary theory of the co-evolution of religion and human social cooperation: the signalling theory of religion, or religious signalling theory (RST). According to the signalling theory, participation in social religion (and its associated rituals and sanctions) acts (...) as an honest signal of one’s commitment to a religiously demarcated community and its way of doing things. This signal would allow prosocial individuals to positively assort with one another for mutual advantage, to the exclusion of more exploitative individuals. In effect, the theory offers a way that religion and cooperation might explain one another, but which that stays within an individualist adaptive paradigm. My approach is not to assess the empirical adequacy of the religious signalling explanation or contrast it with other explanations, but rather to deal with the theory in its own terms – isolating and fleshing out its core commitments, explanatory potential, and limitations. The key to this is acknowledging the internal complexities of signalling theory, with respect to the available models of honest signalling and the extent of their fit (or otherwise) with religion as a target system. The method is to take seriously the findings of formal modelling in animal signalling and other disciplines, and to apply these (and methods from the philosophy of biology more generally) to progressively build up a comprehensive picture of the theory, its inherent strengths and weaknesses. The first two chapters outline the dual explanatory problems that cooperation and religion present for evolutionary human science, and surveys contemporary approaches toward explaining them. Chapter three articulates an evolutionary conception of the signalling theory, and chapters four to six make the case for a series of requirements, limitations, and principles of application. Chapters seven and eight argue for the value of formal modelling to further flesh out the theory’s commitments and potential and describe some simple simulation results which make progress in this regard. Though the inquiry often problematizes the signalling theory, it also shows that it should not be dismissed outright, and that it makes predictions which are apt for empirical testing. (shrink)

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

The traditional practice of establishing morphological types and investigating morphological organization has found new support from evolutionary developmental biology (evo-devo), especially with respect to the notion of body plans. Despite recurring claims that typology is at odds with evolutionary thinking, evo-devo offers mechanistic explanations of the evolutionary origin, transformation, and evolvability of morphological organization. In parallel, philosophers have developed non-essentialist conceptions of natural kinds that permit kinds to exhibit variation and undergo change. This not only facilitates a construal of species (...) and higher taxa as natural kinds, but also broadens our perspective on the diversity of kinds found in biology. There are many different natural kinds relevant to the investigative and explanatory aims of evo-devo, including homologues and developmental modules. (shrink)

Comparing engineering to evolution typically involves adaptationist thinking, where well-designed artifacts are likened to well-adapted organisms, and the process of evolution is likened to the process of design. A quite different comparison is made when biologists focus on evolvability instead of adaptationism. Here, the idea is that complex integrated systems, whether evolved or engineered, share universal principles that affect the way they change over time. This shift from adaptationism to evolvability is a significant move for, as I argue, we can (...) make sense of these universal principles without making any adaptationism claims. Furthermore, evolvability highlights important aspects of engineering that are ignored in the adaptationist debates. I introduce some novel engineering examples that incorporate these key neglected aspects, and use these examples to challenge some commonly cited contrasts between engineering and evolution, and to highlight some novel resemblances that have gone unnoticed. (shrink)

Against the background of “arms race” style competitive explanations for complex human cognition, such as the Social Intelligence Hypothesis Growing points in ethology, Cambridge University Press, pp 303–317, 1976; Jolly in Science, 10.1126/science.153.3735.501, 1966), and theories that tie complex cognition with environmental variability more broadly The evolution of intelligence, Lawrence Earlbaum and Associates, 2001), the idea that culturally inherited mechanisms for social cognition would be more capable of responding to the labile social environment is a compelling one. Whilst it is (...) tempting to think that the evolvability of culturally inherited cognitive mechanisms such as Cecilia Heyes’ cognitive gadgets would be akin to culturally inherited tools like axes or canoes, I draw on established theory in evolutionary developmental biology to show that this is a mistake. Their causal translucency, along with the degree to which they would be integrated within the organism, make cognitive gadgets far more like genetically inherited traits with respect to their evolvability. Consequently, their evolution is unlikely to be particularly fast or nimble. In making clear the constraints on the evolution of culturally inherited cognition and how they must influence our theorising the discussion also highlights the value of thinking about evolvability in this domain. (shrink)

Although the concept of modularity is pervasive across fields and disciplines, philosophers and scientists use the term in a variety of different ways. This paper identifies two distinct ways of thinking about modularity, and considers what makes them similar and different. For philosophers of mind and cognitive science, cognitive modularity helps explain the capacities of brains to process sundry and distinct kinds of informational input. For philosophy of biology and evolutionary science, biological modularity helps explain the capacity of random evolutionary (...) processes to give rise to highly complex and sophisticated biological systems. Although these different ways of thinking about modularity are largely distinct, this paper proposes a unifying feature common to both: isolability, or the capacity of subsystems to undergo changes without resulting in substantial changes to neighboring or interconnected subsystems. (shrink)

I develop an account of homology and homoplasy drawing on their use in biological inference and explanation. Biologists call on homology and homoplasy to infer character states, support adaptationist explanations, identify evolutionary novelties and hypothesize phylogenetic relationships. In these contexts, the concepts must be understood phylogenetically and kept separate: as they play divergent roles, overlap between the two ought to be avoided. I use these considerations to criticize an otherwise attractive view defended by Gould, Hall, and Ramsey & Peterson. By (...) this view, homology and homoplasy can only be delineated qua some level of description, and some homoplasies (parallelisms) are counted as homologous. I develop an account which retains the first, but rejects the second, aspect of that view. I then characterize parallelisms and convergences in terms of their causal role. By the Strict Continuity account, homology and homoplasy are defined phylogenetically and without overlaps, meeting my restriction. Convergence and parallelisms are defined as two types of homoplasy: convergent homoplasies are largely constrained by external factors, while parallelisms are due to internal constraints. (shrink)

I identify six types of phenotypic plasticity and categorize them with respect to their cognitive status. I look at differences and relations between some of these types of plasticity and then analyze how phenotypic outcomes are transmitted across generations. I engage with the relevant literature on developmental scaffolding and entrenchment in cultural evolution. I argue that that the typology I present here can be beneficial for such a debate and therefore instructive to better comprehend the evolution and development of human (...) cognition. (shrink)

In Denis Walsh’s Organisms, Agency, and Evolution, he argues that new developments in the science of biology motivate a radical change to our metaphysical picture of life: what he calls ‘Situated Darwinism’. The central claim is that we should take the biological world to be at base about organisms, and organisms in a fundamentally teleological sense. We critically examine Walsh’s arguments and suggest further developments.

A modularidade massiva pode ser descrita como um programa de pesquisa, nos termos da filosofia da ciência de Lakatos. Cumpre, para tanto, identificar o núcleo, mostrar evidências de que ele é assumido como infalsificável e examinar estratégias de articulação e sofisticação do cinto de proteção. No presente trabalho, faço isso com o intuito de responder à pergunta lançada no título. Em atenção a três desenvolvimentos recentes, cujas predições encontraram corroboração empírica, sou levado a inferir que esse programa é, de fato, (...) empiricamente progressivo. (shrink)