Methodological problems in evolutionary biology VI. The force of evolutionary epistemology

Evolutionary epistemology takes various forms. As a philosophical discipline, it may use analogies by borrowing concepts from evolutionary biology to establish new foundations. This is not a very successful enterprise because the analogies involved are so weak that they hardly have explanatory force. It may also veil itself with the garbs of biology. Proponents of this strategy have only produced irrelevant theories by transforming epistemology's concepts beyond recognition. Sensible theories about knowledge and biology should presuppose that various long-standing problems concerning relations between the mental and the physical are solved. Such problems are wrongly disregarded by evolutionary epistemologists.     

The concept of developmental reprogramming and the quest for an inclusive theory of evolutionary mechanisms

SUMMARY Evolutionary developmental biology has already made a major contribution to our understanding of evolutionary patterns, notably homology. However, while it has the potential to make an equally important contribution to our understanding of evolutionary mechanisms, and indeed to the integration of mechanism and pattern, it has not yet done so. This paper explores how this potential may be realized. In particular, I focus on the limitations of present-day neo-Darwinian theory, and indicate how a combination of the neo-Darwinian and "evo-devo" approaches provides a more inclusive view of evolutionary mechanisms with greater explanatory power. There is a particular focus on developmental reprogramming, which lies logically between mutation and selection, yet has been neglected in mainstream evolutionary theory. The inclusion of developmental reprogramming in the list of evolutionary mechanisms leads to a view that the direction of evolutionary change is determined by a combination of internal and external factors, rather than being controlled entirely by the environment.     

Evo-Devo: evolutionary developmental mechanisms

Evolutionary developmental biology (Evo-Devo) as a discipline is concerned, among other things, with discovering and understanding the role of changes in developmental mechanisms in the evolutionary origin of aspects of the phenotype. In a very real sense, Evo-Devo opens the black box between genotype and phenotype, or more properly, phenotypes as multiple life history stages arise in many organisms from a single genotype. Changes in the timing or positioning of an aspect of development in a descendant relative to an ancestor (heterochrony and heterotopy) were two evolutionary developmental mechanisms identified by Ernst Haeckel in the 1870s. Many more have since been identified, in large part because of our enhanced understanding of development and because new mechanisms emerge as development proceeds: the transfer from maternal to zygotic genomic control; cell-to-cell interactions; cell differentiation and cell migration; embryonic inductions; functional interactions at the tissue and organ levels; growth. Within these emergent processes, gene networks and gene cascades (genetic modules) link the genotype with morphogenetic units (cellular modules, namely germ layers, embryonic fields or cellular condensations), while epigenetic processes such as embryonic inductions, tissue interactions and functional integration, link morphogenetic units to the phenotype. Evolutionary developmental mechanisms also include interactions between individuals of the same species, individuals of different species, and species and their biotic and/or abiotic environment. Such interactions link ecological communities. Importantly, there is little to distinguish the causality that underlies these interactions from that which underlies inductive interactions within embryos.     

Evolutionary biology: a basic science for medicine in the 21st century

Evolutionary biology was a poorly developed discipline at the time of the Flexner Report and was not included in Flexner's recommendations for premedical or medical education. Since that time, however, the value of an evolutionary approach to medicine has become increasingly recognized. There are several ways in which an evolutionary perspective can enrich medical education and improve medical practice. Evolutionary considerations rationalize our continued susceptibility or vulnerability to disease; they call attention to the idea that the signs and symptoms of disease may be adaptations that prevent or limit the severity of disease; they help us understand the ways in which our interventions may affect the evolution of microbial pathogens and of cancer cells; and they provide a framework for thinking about population variation and risk factors for disease. Evolutionary biology should become a foundational science for the medical education of the future.     

TRIBUTE: In Goethe's Wake: Marvalee Wake's conceptual contributions to the development and evolution of a science of morphology

Decrying the typological approach in much of the teaching of morphology, from the outset of her career Marvalee Wake advocated a synthetic, mechanistic and pluralistic developmental and evolutionary morphology. In this short essay, I do not evaluate Wake's contributions to our knowledge of the morphology of caecilians, nor her contributions to viviparity, both of which are seminal and substantive, nor do I examine her role as mentor, supervisor and collaborator, but assess her broader conceptual contributions to the development and evolution of morphology as a science. One of the earliest morphologists to take on board the concept of constraint, she viewed constraint explicitly in relation to adaptation and diversity. Her approach to morphology as a science was hierarchical – measure form and function in a phylogenetic context; seek explanations at developmental, functional, ecological, evolutionary levels of the biological hierarchy; integrate those explanations to the other levels. The explanatory power of morphology thus practised allows morphology to inform evolutionary biology and evolutionary theory, and paves the way for the integrative biology Wake has long championed.     

Probabilistic causation and the explanatory role of natural selection

The explanatory role of natural selection is one of the long-term debates in evolutionary biology. Nevertheless, the consensus has been slippery because conceptual confusions and the absence of a unified, formal causal model that integrates different explanatory scopes of natural selection. In this study we attempt to examine two questions: (i) What can the theory of natural selection explain? and (ii) Is there a causal or explanatory model that integrates all natural selection explananda? For the first question, we argue that five explananda have been assigned to the theory of natural selection and that four of them may be actually considered explananda of natural selection. For the second question, we claim that a probabilistic conception of causality and the statistical relevance concept of explanation are both good models for understanding the explanatory role of natural selection. We review the biological and philosophical disputes about the explanatory role of natural selection and formalize some explananda in probabilistic terms using classical results from population genetics. Most of these explananda have been discussed in philosophical terms but some of them have been mixed up and confused. We analyze and set the limits of these problems.     

Physiological anthropology: past and future

Environmental studies in adaptive human biology by North American anthropologists have a history of strong investigative research. From both laboratory and field work, we have gained major insights into human response to physical and social challenges. While these results were considered by most professionals to belong within evolutionary biology, in fact the intellectual structure sprang almost entirely from physiological equilibrium models. Consequently, physiological process itself was the focus. Further, most of the physiological patterns were not linked directly to important outcomes such as work output, reproductive success or survival.About 1975, American physiological anthropologists, led by Paul Baker, turned to studies of health, change and stress response. These studies were strong, but were still neither genetic nor evolutionary in intellectual structure. Evolutionary human biology was taken over by a new body of theory now called "behavior ecology", positing that selfish genes control human behavior to promote their own reproduction. This was paralleled by strong use of evolutionary theory in some areas of molecular biology. However, although physiological anthropologists have not focused on evolution, we have been developing powerful causal models that incorporate elements of physiology, morphology, physical environment and cultural behavior. In these "proximate" biocultural models, it is of little importance whether outcomes such as work or energy management are genetically based.Our future offers two major challenges. First, we must confirm causal links between specific physiological patterns and outcomes of practical importance to individuals and societies. Second, if we are to take our place in evolutionary biology, the one overarching theory of life on earth, we must understand the heritability of physiological traits, and determine whether they play a role in survival and reproduction.     

Students’ Mental Models of Evolutionary Causation: Natural Selection and Genetic Drift

In an effort to understand how to improve student learning about evolution, a focus of science education research has been to document and address students?? naive ideas. Less research has investigated how students reason about alternative scientific models that attempt to explain the same phenomenon (e.g., which causal model best accounts for evolutionary change?). Within evolutionary biology, research has yet to explore how non-adaptive factors are situated within students?? conceptual ecologies of evolutionary causation. Do students construct evolutionary explanations that include non-adaptive and adaptive factors? If so, how are non-adaptive factors structured within students?? evolutionary explanations? We used clinical interviews and two paper and pencil instruments (one open-response and one multiple-choice) to investigate the use of non-adaptive and adaptive factors in undergraduate students?? patterns of evolutionary reasoning. After instruction that included non-adaptive causal factors (e.g., genetic drift), we found them to be remarkably uncommon in students?? explanatory models of evolutionary change in both written assessments and clinical interviews. However, consistent with many evolutionary biologists?? explanations, when students used non-adaptive factors they were conceptualized as causal alternatives to selection. Interestingly, use of non-adaptive factors was not associated with greater understanding of natural selection in interviews or written assessments, or with fewer naive ideas of natural selection. Thus, reasoning using non-adaptive factors appears to be a distinct facet of evolutionary thinking. We propose a theoretical framework for an expert?Cnovice continuum of evolutionary reasoning that incorporates both adaptive and non-adaptive factors, and can be used to inform instructional efficacy in evolutionary biology.     

Explanatory pluralism in evolutionary biology

The ontological dependence of one domain on another is compatible with the explanatory autonomy of the less basic domain. That autonomy results from the fact that the relationship between two domains can be very complex. In this paper I distinguish two different types of complexity, two ways the relationship between domains can fail to be transparent, both of which are relevant to evolutionary biology. Sometimes high level explanations preserve a certain type of causal or counterfactual information which would be lost at the lower level; I argue that this is central to the proper understanding of the adaptationist program. Sometimes high level kinds are multiply realised by lower level kinds: I argue that this is central to the understanding of macroevolution.     

Evolution and the Cumulative Nature of Science

One of the characteristics of science is its cumulative nature. As more discoveries are made and more is learned, we progressively come to a more and more complete understanding of the physical universe. Evolutionary biology serves as an excellent example of this progressive trend in knowledge, since we now understand significantly more about the mechanisms and details of the evolutionary process than we did in decades past. It is important to comprehend these progressive changes and communicate them effectively to students of evolutionary biology.     

The emerging structure of the Extended Evolutionary Synthesis: where does Evo-Devo fit in?

The Extended Evolutionary Synthesis (EES) debate is gaining ground in contemporary evolutionary biology. In parallel, a number of philosophical standpoints have emerged in an attempt to clarify what exactly is represented by the EES. For Massimo Pigliucci, we are in the wake of the newest instantiation of a persisting Kuhnian paradigm; in contrast, Telmo Pievani has contended that the transition to an EES could be best represented as a progressive reformation of a prior Lakatosian scientific research program, with the extension of its Neo-Darwinian core and the addition of a brand-new protective belt of assumptions and auxiliary hypotheses. Here, we argue that those philosophical vantage points are not the only ways to interpret what current proposals to ‘extend’ the Modern Synthesis-derived ‘standard evolutionary theory’ (SET) entail in terms of theoretical change in evolutionary biology. We specifically propose the image of the emergent EES as a vast network of models and interweaved representations that, instantiated in diverse practices, are connected and related in multiple ways. Under that assumption, the EES could be articulated around a paraconsistent network of evolutionary theories (including some elements of the SET), as well as models, practices and representation systems of contemporary evolutionary biology, with edges and nodes that change their position and centrality as a consequence of the co-construction and stabilization of facts and historical discussions revolving around the epistemic goals of this area of the life sciences. We then critically examine the purported structure of the EES—published by Laland and collaborators in 2015—in light of our own network-based proposal. Finally, we consider which epistemic units of Evo-Devo are present or still missing from the EES, in preparation for further analyses of the topic of explanatory integration in this conceptual framework.     

Deconstructing morphology

Scholtz, G. 2010. Deconstructing morphology. —Acta Zoologica (Stockholm) 91 : 44–63 Morphology as the science of form is, in particular, related to the overwhelming diversity of animal forms. Due to its long pre‐Darwinian tradition, organismic morphology is partly burdened by ahistorical typological views. On the other hand, the study of organismic form has always implied concepts of transformation, which helped to pave the way for evolutionary theories. This contradictory history and the fact that we need words to describe organismic form lead in many cases to morphological concepts implying a mixture of structural, functional, developmental, ecological, typological, and evolutionary aspects in current morphological approaches. Because these mixed views lead to contradictory and misleading interpretations of animal form, I stress the need to deconstruct morphological concepts at all levels. I propose a morphology that analyses transformation of animal forms strictly at the structural level in combination with genealogical thinking. Function and other biological aspects of form should be considered in an independent second analytical step. A comparative pattern approach, including developmental patterns, of animal structure in an evolutionary framework allows for the analysis of morphological change, in particular, phylogenetic reconstructions, homology assessment, and the recognition of evolutionary independent morphological units.     

Don’t Call it “Darwinism”

Evolutionary biology owes much to Charles Darwin, whose discussions of common descent and natural selection provide the foundations of the discipline. But evolutionary biology has expanded well beyond its foundations to encompass many theories and concepts unknown in the 19th century. The term “Darwinism” is, therefore, ambiguous and misleading. Compounding the problem of “Darwinism” is the hijacking of the term by creationists to portray evolution as a dangerous ideology—an “ism”—that has no place in the science classroom. When scientists and teachers use “Darwinism” as synonymous with evolutionary biology, it reinforces such a misleading portrayal and hinders efforts to present the scientific standing of evolution accurately. Accordingly, the term “Darwinism” should be abandoned as a synonym for evolutionary biology.     

Evolution of evolutionary physiology

In 19th century and at the beginning 20th century, reports appeared in the field of comparative and ontogenetic physiology and the value of these methods for understanding of evolution of functions. The term "evolutionary physiology" was suggested by A. N. Severtsov in 1914. In the beginning of 30s, in the USSR, laboratories for researches in problems of evolutionary physiology were created, the results of these researches having been published. In 1956 in Leningrad, the Institute of Evolutionary Physiology was founded by L. A. Orbeli. He formulates the goals and methods of evolutionary physiology. In the following half a century, the evolutionary physiology was actively developed. The evolutionary physiology solves problems of evolution of function of functions evolution, often involving methods of adjacent sciences, including biochemistry, morphology, molecular biology.     

Methodological Problems in Evolutionary Biology. XII. Against Evolutionary Ethics

Evolutionary ethics has recently become popular again. Some of its representatives elaborate new attempts to derive ethics from evolutionary biology. The attempts, like previous ones, fail because they commit the naturalistic fallacy. Premises from evolutionary biology together with normative premises also do not justify ethical principles. Other representatives argue that evolutionary considerations imply that ethics cannot be justified at all. Their arguments presuppose an unacceptable form of foundationalism. In principle, evolutionary biology might explain some aspects of morality, but in practice explanations are hard to come by. All this does not imply that evolutionary theory is irrelevant in normative settings. To the contrary, it may help us devise guidelines in environmental policy and health care policy. It is to be hoped that evolutionary ethicists will divert their research efforts to the elaboration of such guidelines.     

How to Identify and Interpret Evolutionary Tree Diagrams

Evolutionary trees are key tools for modern biology and are commonly portrayed in textbooks to promote learning about biological evolution. However, many people have difficulty in understanding what evolutionary trees are meant to portray. In fact, some ideas that current professional biologists depict with evolutionary trees are neither clearly defined nor conveyed to students. To help biology teachers and students learn how to more deeply interpret, understand and gain knowledge from diagrams that represent ancestor–descendant relationships and evolutionary lineages, we describe the different rooted and unrooted evolutionary tree visualisations and explain how they are best read. Examples from a study of tree-shaped diagrams in the journal Science are used to illustrate how to distinguish evolutionary trees from other tree-shaped representations that are easily misunderstood as visualising evolutionary relationships. We end by making recommendations for how our findings may be implemented in teaching practice in this important area of biology education.     

Proper names in twin worlds: Monophyly, paraphyly, and the world around us

This paper addresses the theoretical relevance of monophyletic, paraphyletic and polyphyletic groups under the paradigm of sophisticated scientific realism. The doctrine of metaphysical realism is introduced using the philosophy of Karl Popper as an example, which is then contrasted with scientific realism. A discussion of the nature of causal relations presents an account of counterfactual conditionals. The current state of art casts the theory of phylogenetic systematics in a stark contrast of classes (universals) and individuals (particulars). In practice, however, individuals piggyback on classes, or sets. Natural kinds are introduced in order to overcome this deep dichotomy. The theoretical relevance of natural kinds lies in their explanatory value, and that may change with changing context. It is for this reason that non-monophyletic groups can have explanatory value (their members can function as tokens of causally relevant kinds) within certain domains of evolutionary biology. Explanatory value is maximized by integration of the genealogical hierarchy of species and monophyletic taxa with other areas of evolutionary biology.