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 Philosophers have recommended transitioning away from outdated philosophies that were originally derived from physics within the philosophical zeitgeist of logical positivism (i.e., monism, reductionism, and monocausation)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 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)

This chapter offers a critique of intelligent design arguments against evolution and a philosophical discussion of the nature of science, drawing several lessons for the teaching of evolution and for science education in general. I discuss why Behe’s irreducible complexity argument fails, and why his portrayal of organismal systems as machines is detrimental to biology education and any under-standing of how organismal evolution is possible. The idea that the evolution of complex organismal features is too unlikely to have occurred by (...) random mutation and selection (as recently promoted by Dembski) is very widespread, but it is easy to show students why such small probability arguments are fallacious. While intelligent design proponents have claimed that the exclusion of supernatural causes mandated by scientific methods is dogmatically presupposed by science, scientists have an empirical justification for using such methods. This justification is instructive for my discussion of how to demarcate science from pseudoscience. I argue that there is no universal account of the nature of science, but that the criteria used to judge an intellectual approach vary across historical periods and have to be specific to the scientific domain. Moreover, intellectual approaches have to be construed as practices based on institutional factors and values, and to be evaluated in terms of the activities of their practitioners. Science educators should not just teach scientific facts, but present science as a practice and make students reflect on the nature of science, as this gives them a better appreciation of the ways in which intelligent design falls short of actual science. (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)

In a macroevolutionary timescale, evolvability itself evolves. Lineages are sorted based on their ability to generate adaptive novelties, which leads to the optimization of their genotype-phenotype map. The system of translation of genetic or epigenetic changes to the phenotype may reach significant horizontal and vertical complexity, and may even exhibit certain aspects of learning behaviour. This continuously evolving semiotic system probably enables the origin of complex yet functional and internally compatible adaptations. However, it also has a second, “darker”, side. As (...) was pointed out by several authors, the same process gradually reduces the probability of the origination of significant evolutionary novelties. In a similar way to the evolution of societies, teachings, or languages, in which the growing number of internal linkages gradually solidifies their overall structure and the structure or interpretation of their constitutive elements, the evolutionary potential of lineages decreases during biological evolution. Possible adaptations become limited to small “peripheral” modifications. According to the Frozen Evolution theory, some of the proximate causes of this “macroevolutionary freezing” are more pronounced or present exclusively in sexual lineages. Sorting based on the highest evolvability probably leads to the establishment of certain structural features of complex organisms, e.g. the modular character of their development and morphology. However, modules also “macroevolutionary freeze” whereas the hypothetical “thawing” of modules or their novel adaptive combinations becomes rarer and rarer. Some possible ways out of this dead end include the rearrangement of individual development, e.g. neoteny, radical simplification, i.e. sacculinization, and transition to a higher level of organization, e.g. symbiosis or symbiogenesis. The evolution of evolvability is essentially a biosemiotic process situated at the intersection of the genocentric modern synthesis and the evo-devo-centric extended synthesis. Therefore, evolvability may eventually connect these three not necessarily contradictory approaches. (shrink)

In the 1930s, two concepts excited the European biological community: the organizer phenomenon and organicism. This essay examines the history of and connection between these two phenomena in order to address the conventional ‘rise-and-fall’ narrative that historians have assigned to each. Scholars promoted the ‘rise-and-fall’ narrative in connection with a broader account of the devitalizing of biology through the twentieth century. I argue that while limited evidence exists for the ‘fall of the organizer concept’ by the 1950s, the organicism that (...) often motivated the organizer work had no concomitant fall – even during the mid-century heyday of molecular biology. My argument is based on an examination of shifting social networks of life scientists from the 1920s to the 1970s, many of whom attended or corresponded with members of the Cambridge Theoretical Biology Club . I conclude that the status and cohesion of these social networks at the micro scale was at least as important as macro-scale conceptual factors in determining the relative persuasiveness of organicist philosophy. (shrink)

What is the relationship between evolutionary contingency and diversity? The evolutionary contingency thesis emphasizes dependency relations and chance as the hallmarks of evolution. While contingency can be destructive of, for example, the fragile and complex dynamics in an ecosystem, I will mainly focus on the productive or causal aspect of contingency for a particular sort of diversity. There are many sorts of diversities: Gould is most famous for his diversity-to-decimation model, which includes disparate body plans distinguishing different phyla. However, structural (...) diversity construed more broadly spans scales, such as organization in and among cells, structural arrangements and biomechanics on various scales, and even the profile of ancestor-descendent relationships or community structure of interactions within ecosystems. By focusing on stochastic processes in contingent evolution, I argue that contingency causes structural diversity. Specifically, I focus on the plurality of structural types of cells, genetic codes, and phyla diversity as case studies. (shrink)

What gets integrated in integrative scientific practices has been a topic of much discussion. Traditional views focus on theories and explanations, with ideas of reduction and unification dominating the conversation. More recent ideas focus on disciplines, fields, or specialties; models, mechanisms, or methods; phenomena, problems. How integration works looks different on each of these views since the objects of integration are ontologically and epistemically various: statements, boundary conditions, practices, protocols, methods, variables, parameters, domains, laboratories, and questions all have their own (...) structures, functions and logics. I focus on one particular kind of scientific practice, integration of “approaches” in the context of a research system operating on a special kind of “platform.” Rather than trace a network of interactions among people, practices, and theoretical entities to be integrated, in this essay I focus on the work of a single investigator, David Wake. I describe Wake’s practice of integrative evolutionary biology and how his integration of approaches among biological specialties worked in tandem with his development of the salamanders as a model taxon, which he used as a platform to solve, re-work and update problems that would not have been solved so well by non-integrative approaches. The larger goal of the project to which this paper contributes is a counter-narrative to the story of 20th century life sciences as the rise and march of the model organisms and decline of natural history. (shrink)

Due to the variation, contingency and complexity of living systems, biology is often taken to be a science without fundamental theories, laws or general principles. I revisit this question in light of the quest for design principles in systems biology and show that different views can be reconciled if we distinguish between different types of generality. The philosophical literature has primarily focused on generality of specific models or explanations, or on the heuristic role of abstraction. This paper takes a different (...) approach in emphasizing a theory-constituting role of general principles. Design principles signify general dependency-relations between structures and functions, given a set of formally defined constraints. I contend that design principles increase our understanding of living systems by relating specific models to general types. The categorization of types is based on a delineation of the scope of biological possibilities, which serves to identify and define the generic features of classes of systems. To characterize the basis for general principles through generic abstraction and reasoning about possibility spaces, I coin the term constraint-based generality. I show that constraint-based generality is distinct from other types of generality in biology, and argue that general principles play a unifying role that does not entail theory reduction. (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. 2.1Mutational randomness 2.2Morphological randomness 2.3Adaptive randomness3Variability and Phenotypic Probabilities 3.1The RNA model as a model for evo-devo4Evolvability and the Probability of Evolving5Variational Propensities in Evo-Devo 5.1Individuating variational propensities 5.2Developmental types6Conclusion. (shrink)

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

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)

Despite its pervasiveness, the concept of ‘levels of organization’ has received relatively little attention in its own right. I propose here an emerging approach that posits ‘levels’ as a fragmentary concept situated within an interest-relative matrix of operational usage within scientific practice. To this end I propose one important component of meaning, namely the epistemic goal motivating the term’s usage, which recovers a remarkably conserved and sufficiently unifying significance attributable to ‘levels’ across different instances of usage. This epistemic goal, to (...) provide structure to scientific problems, delegates tasks whose execution generates the term’s expressed content in a given instance. This treatment of levels does not diminish the concept’s general importance to science, but rather allows for its use in, and usefulness for, scientific practice to be better contextualized to particular tasks encompassing varying breadths of activity. (shrink)

Understanding the role mechanistic constraints play in shaping evolution can relieve the tension between the generally accepted intuition that there are no strict laws in biology and empirical findings showing that evolutionary processes are biased toward preferred outcomes. Mechanistic constraints explain why some evolutionary outcomes are more probable than others and allow for predictions in specific lineages. At the same time, mechanistic constraints are neither necessary nor universal in the way laws are traditionally characterized: they remain contingent on the past (...) evolution of the biological mechanisms underpinning them and only constrain the future evolution of the organisms possessing them. (shrink)

Evolutionary developmental biology (evo-devo) is considered a ‘mechanistic science,’ in that it causally explains morphological evolution in terms of changes in developmental mechanisms. Evo-devo is also an interdisciplinary and integrative approach, as its explanations use contributions from many fields and pertain to different levels of organismal organization. Philosophical accounts of mechanistic explanation are currently highly prominent, and have been particularly able to capture the integrative nature of multifield and multilevel explanations. However, I argue that evo-devo demonstrates the need for a (...) broadened philosophical conception of mechanisms and mechanistic explanation. Mechanistic explanation (in terms of the qualitative interactions of the structural parts of a whole) has been developed as an alternative to the traditional idea of explanation as derivation from laws or quantitative principles. Against the picture promoted by Carl Craver, that mathematical models describe but usually do not explain, my discussion of cases from the strand of evo-devo which is concerned with developmental processes points to qualitative phenomena where quantitative mathematical models are an indispensable part of the explanation. While philosophical accounts have focused on the actual organization and operation of mechanisms, properties of developmental mechanisms that are about how a mechanism reacts to modifications are of major evolutionary significance, including robustness, phenotypic plasticity, and modularity. A philosophical conception of mechanisms is needed that takes into account quantitative changes, transient entities and the generation of novel types of entities, feedback loops and complex interaction networks, emergent properties, and, in particular, functional-dynamical aspects of mechanisms, including functional (as opposed to structural) organization and distributed, system-wide phenomena. I conclude with general remarks on philosophical accounts of explanation. (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)

Due to the variation, contingency and complexity of living systems, biology is often taken to be a science without fundamental theories, laws or general principles. I revisit this question in light of the quest for design principles in systems biology and show that different views can be reconciled if we distinguish between different types of generality. The philosophical literature has primarily focused on generality of specific models or explanations, or on the heuristic role of abstraction. This paper takes a different (...) approach in emphasizing a theory-constituting role of general principles. Design principles signify general dependencyrelations between structures and functions, given a set of formally defined constraints. I contend that design principles increase our understanding of living systems by relating specific models to general types. The categorization of types is based on a delineation of the scope of biological possibilities, which serves to identify and define the generic features of classes of systems. To characterize the basis for general principles through generic abstraction and reasoning about possibility spaces, I coin the term constraint-based generality. I show that constraintbased generality is distinct from other types of generality in biology, and argue that general principles play a unifying role that does not entail theory reduction. (shrink)