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 Biology3.1 The explanatory targets of evolutionary biology3.2 Selection-based explanations3.3 Lineage explanations3.4 Evolvability-based explanations3.5 What properties must evolvability have?4 What Evolvability Really Is4.1 Making sense of ft4.2 Making sense of x and b5 What of the Limbs? The Power of E6 Conclusion. (shrink)

Godfrey-Smith ( 2001 ) has distinguished three types of adaptationism. This article builds on his analysis, and revises it in places, by distinguishing seven varieties of adaptationism. This taxonomy allows us to clarify what is at stake in debates over adaptationism, and it also helps to cement the importance of Gould and Lewontin’s ‘Spandrels’ essay. Some adaptationists have suggested that their essay does not offer any coherent alternative to the adaptationist programme: it consists only in an exhortation to test adaptationist (...) hypotheses more thoroughly than was usual in the 1970s. Here it is argued that the ‘Spandrels’ paper points towards a genuinely non-adaptationist methodology implicit in much evolutionary developmental biology. This conclusion helps to expose the links between older debates over adaptationism and more recent questions about the property of evolvability. (shrink)

Universal Basic Income (UBI) has been proposed as a potential way in which welfare states could be made more responsive to the ever-shifting evolutionary challenges of institutional adaptation in a dynamic environment. It has been proposed as a tool of “real freedom” (Van Parijs) and as a tool of making the welfare state more efficient. (Friedman) From the point of view of complexity theory and evolutionary economics, I argue that only a welfare state model that is “polycentrically” (Polanyi, Hayek) organized (...) as an evolving network of distributed decision-making can respond to the challenges of complexity in an adaptively efficient manner. UBI has the theoretical possibility to facilitate flexible, bottom-up innovation, since a) it embodies the “rule of law” principles of generality, nondiscrimination, simplicity, and transparency, and b) it grants people widespread freedom to experiment, innovate, and deviate from established practices and norms; but whether it can be made to work as intended depends on a number of variables in its design. I situate my research in the contemporary UBI debate around automation and technological development, which I interpret creatively through the lens of contemporary evolutionary political economy. I advance a syncretic model of institutional design based upon the shared insights of the Santa Fe, Neo-Hayekian, and Neo-Schumpeterian schools of evolutionary economics that, following Adam Thierer and Michael Munger, I call the framework of Permissionless Innovation (PI). Under the PI framework, which is a development of the tradition of evolutionary liberalism (Hume, Smith, Mandeville, Mill, Hayek, Polanyi, Schumpeter, Hodgson), individuals are granted the default right to innovate without having to ask for permission, and the right to basic income, which they can use to support their right to innovate, mutate, and experiment. This framework is a liberal framework to the extent it relies on widespread human freedom as a means of fostering innovation and social learning. And it is an evolutionary framework to the extent that it seeks to facilitate evolutionary socioeconomic processes for the sake of social progress and welfare advancement. I proceed to show, through an analysis of how innovation operates in both the economy and the cultural domain, that UBI can be used as an important cornerstone of an evolutionary liberal model of “ecostructural” governance that treats the socioeconomic order as an ecological garden of spontaneous growths, adaptations, and innovations. I call this the Permissionless Innovation Universal Basic Income (PIUBI) framework. In it, the right to innovate, deviate, and mutate is upheld as the “gold standard” of institutional design whereby individuals are granted widespread freedoms combined with the provision of UBI and other innovation-fostering (carefully bound) services and regulations. In the coming decades, UBI may even enable poor people to take better advantage of Human Enhancement Technologies (HETs) and other experimental innovations. Such a framework is compatible with a fast pace of evolutionary development, but the regulatory model accompanying such a framework must be capable of responding to evolutionary lock-ins, maladaptive innovations, self-destructive behaviour on the part of UBI recipients, and the management of catastrophic and existential risks. In the end, I will have shown that the PIUBI framework can be justified as a plausible default institutional mechanism for responding to radical uncertainty, but it needs to be experimentally adapted to changing circumstances with the help of various situation-specific, polycentric, multi-level, approaches. (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)

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)

Evolutionary biology is a field currently animated by much discussion concerning its conceptual foundations. On the one hand, we have supporters of a classical view of evolutionary theory, whose backbone is provided by population genetics and the so-called Modern Synthesis (MS). On the other hand, a number of researchers are calling for an Extended Synthe- sis (ES) that takes seriously both the limitations of the MS (such as its inability to incorporate developmental biology) and recent empirical and theoretical research on (...) issues such as evolvability, modularity, and self-organization. In this article, I engage in an in-depth commentary of an influential paper by population geneticist Michael Lynch, which I take to be the best defense of the MS-population genetics position published so far. I show why I think that Lynch’s arguments are wanting and propose a modification of evolutionary theory that retains but greatly expands on population genetics. (shrink)

This volume explores questions about conceptual change from both scientific and philosophical viewpoints by analyzing the recent history of evolutionary developmental biology. It features revised papers that originated from the workshop "Conceptual Change in Biological Science: Evolutionary Developmental Biology, 1981-2011" held at the Max Planck Institute for the History of Science in Berlin in July 2010. The Preface has been written by Ron Amundson. In these papers, philosophers and biologists compare and contrast key concepts in evolutionary developmental biology and their (...) development since the original, seminal Dahlem conference on evolution and development held in Berlin in 1981. Many of the original scientific participants from the 1981 conference are also contributors to this new volume and, in conjunction with other expert biologists and philosophers specializing on these topics, provide an authoritative, comprehensive view on the subject. Taken together, the papers supply novel perspectives on how and why the conceptual landscape has shifted and stabilized in particular ways, yielding insights into the dynamic epistemic changes that have occurred over the past three decades. This volume will appeal to philosophers of biology studying conceptual change, evolutionary developmental biologists focused on comprehending the genesis of their field and evaluating its future directions, and historians of biology examining this period when the intersection of evolution and development rose again to prominence in biological science. (shrink)

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

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)

Evo-Devo exhibits a plurality of scientific “cultures” of practice and theory. When are the cultures acting—individually or collectively—in ways that actually move research forward, empirically, theoretically, and ethically? When do they become imperialistic, in the sense of excluding and subordinating other cultures? This chapter identifies six cultures – three /styles/ (mathematical modeling, mechanism, and history) and three /paradigms/ (adaptationism, structuralism, and cladism). The key assumptions standing behind, under, or within each of these cultures are explored. Characterizing the internal structure of (...) the cultures is necessary for understanding how they collaborate or compete, and how they are fragmented or integrated, in the rich interdisciplinary /trading zone/ (Galison 1997) of Evo-Devo. Evo-Devo is an important example of how science can progress through a radical plurality of perspectives and cultures. (shrink)

‘‘Theoretical biology’’ is a surprisingly heter- ogeneous field, partly because it encompasses ‘‘doing the- ory’’ across disciplines as diverse as molecular biology, systematics, ecology, and evolutionary biology. Moreover, it is done in a stunning variety of different ways, using anything from formal analytical models to computer sim- ulations, from graphic representations to verbal arguments. In this essay I survey a number of aspects of what it means to do theoretical biology, and how they compare with the allegedly much more restricted (...) sense of theory in the physical sciences. I also tackle a recent trend toward the presentation of all-encompassing theories in the biological sciences, from general theories of ecology to a recent attempt to provide a conceptual framework for the entire set of biological disciplines. Finally, I discuss the roles played by philosophers of science in criticizing and shap- ing biological theorizing. (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)

Recent debates between proponents of the modern evolutionary synthesis (the standard model in evolutionary biology) and those of a possible extended synthesis are a good example of the fascinating tangle among empirical, theoretical, and conceptual or philosophical matters that is the practice of evolutionary biology. In this essay, we briefly discuss two case studies from this debate, highlighting the relevance of philosophical thinking to evolutionary biologists in the hope of spurring further constructive cross-pollination between the two fields.

The 1981 Dahlem conference was a catalyst for contemporary evolutionary developmental biology (Evo-devo). This introductory chapter rehearses some of the details of the history surrounding the original conference and its associated edited volume, explicates the philosophical problem of conceptual change that provided the rationale for a workshop devoted to evaluating the epistemic revisions and transformations that occurred in the interim, explores conceptual change with respect to the concept of evolutionary novelty, and highlights some of the themes and patterns in the (...) different contributions to the present volume, Conceptual Change in Biology: Scientific and Philosophical Perspectives on Evolution and Development. (shrink)

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

Pere Alberch played a pivotal role in shaping the field of evolutionary developmental biology during the 1980s and 1990s. Whereas initially his contributions were sidelined by the empirical advancements of the molecular revolution in developmental and evolutionary biology, his theoretical insights have left a lasting impact on the discipline. This article provides a comprehensive review of the legacy and evolvability of Alberch's ideas in contemporary evo-devo, which included the study of morphogenesis as the proper level of developmental causation, the interplay (...) between developmental constraints and natural selection, the epistemic role of teratologies, the origin of evolutionary novelties and the concept of evolvability. (shrink)

The developmental properties of organisms play important roles in the generation of variation necessary for evolutionary change. But how can individual development steer the course of evolution? To answer this question, we introduce developmental channeling as a disposition of individual organisms that shapes their possible developmental trajectories and evolutionary dappling as an evolutionary outcome in which the space of possible organismic forms is dappled—it is only partially filled. We then trace out the implications of the channeling-dappling framework for contemporary debates (...) in the philosophy of evolution, including evolvability, reciprocal causation, and the extended evolutionary synthesis. (shrink)

The theory of evolution, which provides the conceptual framework for all modern research in organismal biology and informs research in molecular bi- ology, has gone through several stages of expansion and refinement. Darwin and Wallace (1858) of course proposed the original idea, centering on the twin concepts of natural selection and common descent. Shortly thereafter, Wallace and August Weismann worked toward the complete elimination of any Lamarckian vestiges from the theory, leaning in particular on Weismann’s (1893) concept of the separation (...) of soma and germ, resulting in what is some- times referred to as “neo-Darwinism”. (shrink)

A foundational idea of evo‐devo is that morphological variation is not isotropic, that is, it does not occur in all directions. Instead, some directions of morphological variation are more likely than others from DNA‐level variation and these largely depend on development. We argue that this evo‐devo perspective should apply not only to morphology but to evolution at all phenotypic levels. At other phenotypic levels there is no development, but there are processes that can be seen, in analogy to development, as (...) constructing the phenotype (e.g., protein folding, learning for behavior, etc.). We argue that to explain the direction of evolution two types of arguments need to be combined: generative arguments about which phenotypic variation arises in each generation and selective arguments about which of it passes to the next generation. We explain how a full consideration of the two types of arguments improves the explanatory power of evolutionary theory. Also see the video abstract here: https://youtu.be/Egbvma_uaKc. (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)

Sewall Wright introduced the metaphor of evolution on “adaptive landscapes” in a pair of papers published in 1931 and 1932. The metaphor has been one of the most influential in modern evolutionary biology, although recent theoretical advancements show that it is deeply flawed and may have actually created research questions that are not, in fact, fecund. In this paper I examine in detail what Wright actually said in the 1932 paper, as well as what he thought of the matter at (...) the very end of his career, in 1988. While the metaphor is flawed, some of the problems which Wright was attempting to address are still with us today, and are in the process of being reformulated as part of a forthcoming Extended Evolutionary Synthesis. (shrink)

The origin and prevalence of transposable elements may best be understood as resulting from “selfish” evolutionary processes at the within-genome level, with relevant populations being all members of the same TE family or all potentially mobile DNAs in a species. But the maintenance of families of TEs as evolutionary drivers, if taken as a consequence of selection, might be better understood as a consequence of selection at the level of species or higher, with the relevant populations being species or ecosystems (...) varying in their possession of TEs. In 2015, Brunet and Doolittle made the case for legitimizing claims for an evolutionary role for TEs by recasting such claims as being about species selection. Here I further develop this “how possibly” argument. I note that with a forgivingly broad construal of evolution by natural selection we might come to appreciate many aspects of Life on earth as its products, and TEs as—possibly—contributors to the success of Life by selection at several levels of a biological hierarchy. Thinking broadly makes this proposition a testable Darwinian one. (shrink)

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

Darwinism is defined here as an evolving research tradition based upon the concepts of natural selection acting upon heritable variation articulated via background assumptions about systems dynamics. Darwin’s theory of evolution was developed within a context of the background assumptions of Newtonian systems dynamics. The Modern Evolutionary Synthesis, or neo-Darwinism, successfully joined Darwinian selection and Mendelian genetics by developing population genetics informed by background assumptions of Boltzmannian systems dynamics. Currently the Darwinian Research Tradition is changing as it incorporates new information (...) and ideas from molecular biology, paleontology, developmental biology, and systems ecology. This putative expanded and extended synthesis is most perspicuously deployed using background assumptions from complex systems dynamics. Such attempts seek to not only broaden the range of phenomena encompassed by the Darwinian Research Tradition, such as neutral molecular evolution, punctuated equilibrium, as well as developmental biology, and systems ecology more generally, but to also address issues of the emergence of evolutionary novelties as well as of life itself. (shrink)

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

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)

The assumption that natural selection alone is sufficient to explain not only which traits get fixed in a population/species, but also how they develop, has been questioned since Darwin’s times, and increasingly in the last decades. Alternative theories, linked to genetic and phenotypic processes, or to the theory of complex systems, have been proposed to explain the rise of the phenotypic variety upon which natural selection acts. In this article, we illustrate the current state of the issue and we propose (...) a logical space based on phenotypic robustness that allows a classification of evolutionary phenomena and can provide a framework for unifying all these accounts. (shrink)

There is an active research program currently underway, which treats cancer progression as an evolutionary process. This contribution investigates the ways that cancer progression is like and unlike evolution in other contexts. The aim is to take a multi-level perspective on cancer, investigating the levels at which selection may be acting, the unit or target of selection, the relative roles of selection and drift, and the idea that cancer progression may be a by-product of selection at other levels of organization.

Genetic variability is considered a key to the evolvability of species. The conversion of an adenosine (A) to inosine (I) in primary RNA transcripts can result in an amino acid change in the encoded protein, a change in secondary structure of the RNA, creation or destruction of a splice consensus site, or otherwise alter RNA fate. Substantial transcriptome and proteome variability is generated by A‐to‐I RNA editing through site‐selective post‐transcriptional recoding of single nucleotides. We posit that this epigenetic source of (...) phenotypic variation is an unrecognized mechanism of adaptive evolution. The genetic variation introduced through editing occurs at low evolutionary cost since predominant production of the wild‐type protein is retained. This property even allows exploration of sequence space that is inaccessible through mutation, leading to increased phenotypic plasticity and provides an evolutionary advantage for acclimatization as well as long‐term adaptation. Furthermore, continuous probing for novel RNA editing sites throughout the transcriptome is an intrinsic property of the editing machinery and represents the molecular basis for increased adaptability. We propose that higher organisms have therefore evolved to systems with increasing RNA editing activity and, as a result, to more complex systems. (shrink)

Ever since Darwin a great deal of the conceptual history of biology may be read as a struggle between two philosophical positions: reductionism and holism. On the one hand, we have the reductionist claim that evolution has to be understood in terms of changes at the fundamental causal level of the gene. As Richard Dawkins famously put it, organisms are just ‘lumbering robots’ in the service of their genetic masters. On the other hand, there is a long holistic tradition that (...) focuses on the complexity of developmental systems, on the non-linearity of gene– environment interactions, and on multi-level selective processes to argue that the full story of biology is a bit more complicated than that. Reductionism can marshal on its behalf the spectacular successes of genetics and molecular biology throughout the 20th and 21st centuries. Holism has built on the development of entirely new disciplines and conceptual frameworks over the past few decades, including evo-devo and phenotypic plasticity. Yet, a number of biologists are still actively looking for a way out of the reductionism–holism counterposition, often mentioning the word ‘emergence’ as a way to deal with the conundrum. This paper briefly examines the philosophical history of the concept of emergence, distinguishes between epistemic and ontological accounts of it, and comments on conceptions of emergence that can actually be useful for practising evolutionary biologists. (shrink)

Darwin’s claim that Natural Selection, through optimization of fitness, explains complex biological design has not yet been properly formalized. Alan Grafen’s Formal Darwinism Project aims at providing such a formalization and at demonstrating that fitness maximization is coherent with results from Population Genetics, usually interpreted as denying it. We suggest that Grafen’s proposal suffers from some limitations linked to its concept of design as optimized fitness. In order to overcome these limitations, we propose a classification of evolutionary facts based on (...) a bi-dimensional complexity space, which adds robustness to fitness as measure of the quality of a design. In this space, each point represents an organismic architecture, and movement between two points would be an evolutionary fact. Natural Selection explains movement along the fitness axis, while non-selective mechanisms explain movement on the robustness axis. Moreover, we propose a thermodynamic metaphor to draw parallelisms between notions of fitness and entropy, robustness and energy, and movement in this space and reversible and irreversible cycles. We argue that this metaphor illustrates different evolutionary processes usually overlooked by proposals of formalization of Darwinian theory, such Grafen’s Formal Darwinism Project. (shrink)

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)

According to the modern synthesis, evolution is the gradual change of gene frequencies in a population. The MS is closely allied to adaptationist explanations of phenotypes, where organismic form and behavior is treated as previously selected for and owes its genesis to some remote past. However, some new theories of evolution broadly aligned with the extended evolutionary synthesis, in particular developmental plasticity theory and niche construction theory, foreground the fact that evolution is sometimes much more rapid than previously imagined, and (...) occurs through the active engagement of organisms accommodating and modifying their environments. This article describes how these contemporary theories reveal two interconnected sides of being an adaptive subject, a situated agent that modifies itself and its environment as it lives, and contributes to evolution in turn. MS and adaptationism have a generic logical structure that can be taught anywhere, but because developmental plasticity theory and niche construction theory point to an ontology that foregrounds the agency of the organism, they benefit from in situ exploration. I argue biology as a subject needs to adapt, and call for the renewed importance of field studies, outlining some elements of how such studies might be conceived. I close by considering how understanding organisms as adaptive subjects of evolution has important implications for sustainability education. (shrink)

El proyecto del ‘Darwinismo Formal’ de Alan Grafen propone una formulación matemática de la teoría de Darwin que pretende demostrar que la selección natural moldea los rasgos fenotípicos a través de la maximización de la eficacia. El proyecto de Grafen reposa sobre tres premisas: la selección natural es la única fuerza que moldea los fenotipos; la eficacia es la única medida de le evolución; y el diseño biológico surge como resultado de un proceso de optimización selectiva. En este trabajo argumentamos (...) que estas tres premisas limitan la aplicabilidad del modelo a los hechos evolutivos más simples. Esto se debe a que Grafen implícitamente presupone un concepto de eficacia que es a la vez causa y efecto de las novedades fenotípicas; lo que, por un lado, vacía la teoría de Darwin de poder explicativo y, por el otro, lleva a ignorar la potencial contribución de fuerzas evolutivas no selectivas en la configuración de la complejidad biológica. Para superar estos escollos del proyecto de del Darwinismo Formal - que son además comunes a todos los modelos formales adaptacionistas - proponemos sustituir el concepto causal de eficacia por el de robustez, definiendo así el diseño biológico como forma y función en lugar de como maximización de la eficacia. (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)

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)

Essay review of Michael Ruse (ed.), The Oxford Handbook of the Philosophy of Biology (2008).