Ingo Brigandt

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.

The theory of concepts advanced in the dissertation aims at accounting for a) how a concept makes successful practice possible, and b) how a scientific concept can be subject to rational change in the course of history. Traditional accounts in the philosophy of science have usually studied concepts in terms only of their reference; their concern is to establish a stability of reference in order to address the incommensurability problem. My discussion, in contrast, suggests that each scientific concept consists of three components of content: 1) reference, 2) inferential role, and 3) the epistemic goal pursued with the concept's use. I argue that in the course of history a concept can change in any of these three components, and that change in one component—including change of reference—can be accounted for as being rational relative to other components, in particular a concept's epistemic goal.This semantic framework is applied to two cases from the history of biology: the homology concept as used in 19th and 20th century biology, and the gene concept as used in different parts of the 20th century. The homology case study argues that the advent of Darwinian evolutionary theory, despite introducing a new definition of homology, did not bring about a new homology concept (distinct from the pre-Darwinian concept) in the 19th century. Nowadays, however, distinct homology concepts are used in systematics/evolutionary biology, in evolutionary developmental biology, and in molecular biology. The emergence of these different homology concepts is explained as occurring in a rational fashion. The gene case study argues that conceptual progress occurred with the transition from the classical to the molecular gene concept, despite a change in reference. In the last two decades, change occurred internal to the molecular gene concept, so that nowadays this concept's usage and reference varies from context to context. I argue that this situation emerged rationally and that the current variation in usage and reference is conducive to biological practice.The dissertation uses ideas and methodological tools from the philosophy of mind and language, the philosophy of science, the history of science, and the psychology of concepts.

Recent rival attempts in the philosophy of science to put forward a general theory of the properties that all (and only) natural kinds across the sciences possess may have proven to be futile. Instead, I develop a general methodological framework for how to philosophically study kinds. Any kind has to be investigated and articulated together with the human aims that motivate referring to this kind, where different kinds in the same scientific domain can answer to different concrete aims. My core contention is that non-epistemic aims, including environmental, ethical, and political aims, matter as well. This is defended and illustrated based on several examples of kinds, with particular attention to the role of social-political aims: species, race, gender, as well as personality disorders and oppositional defiant disorder as psychiatric kinds. Such non-epistemic aims and values need not always be those personally favoured by scientists, but may have to reflect values that matter to relevant societal stakeholders. Despite the general agenda to study ‘kinds,’ I argue that philosophers should stop using the term ‘natural kinds,’ as this label obscures the relevance of humans interests and the way in which many kinds are based on contingent social processes subject to human responsibility.

Throughout the recent history of research at the intersection of evolution and development, notions such as developmental constraint, evolutionary novelty, and evolvability have been prominent, but the term ‘developmental bias’ has scarcely been used. And one may even doubt whether a unique and principled definition of bias is possible. I argue that the concept of developmental bias can still play a vital scientific role by means of setting an explanatory agenda that motivates investigation and guides the formulation of integrative explanatory frameworks. Less crucial is a definition that would classify patterns of phenotypic variation and unify variational patterns involving different traits and taxa as all being ‘bias.’ Instead, what we should want is a concept that generates intellectual identity across various researchers, and that unites the diverse fields and approaches relevant to the study of developmental bias, from paleontology to behavioral biology. I point to some advantages of conducting research specifically under the label of ‘developmental bias,’ compared to employing other, more common terms such as ‘evolvability.’

Multilevel research strategies characterize contemporary molecular inquiry into biological systems. We outline conceptual, methodological, and explanatory dimensions of these multilevel strategies in microbial ecology, systems biology, protein research, and developmental biology. This review of emerging lines of inquiry in these fields suggests that multilevel research in molecular life sciences has significant implications for philosophical understandings of explanation, modeling, and representation.

The present paper analyzes the use and understanding of the homology concept across different biological disciplines. It is argued that in its history, the homology concept underwent a sort of adaptive radiation. Once it migrated from comparative anatomy into new biological fields, the homology concept changed in accordance with the theoretical aims and interests of these disciplines. The paper gives a case study of the theoretical role that homology plays in comparative and evolutionary biology, in molecular biology, and in evolutionary developmental biology. It is shown that the concept or variant of homology preferred by a particular biological field is used to bring about items of biological knowledge that are characteristic for this field. A particular branch of biology uses its homology concept to pursue its specific theoretical goals.

Ongoing empirical discoveries in molecular biology have generated novel conceptual challenges and perspectives. Philosophers of biology have reacted to these trends when investigating the practice of molecular biology and contributed to scientific debates on methodological and conceptual matters. This article reviews some major philosophical issues in molecular biology. First, philosophical accounts of mechanistic explanation yield a notion of explanation in the context of molecular biology that does not have to rely on laws of nature and comports well with molecular discovery. Second, reductionism continues to be debated and increasingly be rejected by scientists. Philosophers have likewise moved away from reduction toward integration across fields or integrative explanations covering several levels of organization. Third, although the gene concept has undergone substantial transformation and even fragmentation, it still enjoys widespread use by molecular biologists, which has prompted philosophers to understand the empirical reasons for this. At the same time, it has been argued the notion of ‘genetic information’ is largely an empty metaphor, which generates the illusion of explanatory understanding without offering an adequate explanation of molecular and developmental mechanisms.

Examining previous discussions on how to construe the concepts of gender and race, we advocate what we call strategic conceptual engineering. This is the employment of a (possibly novel) concept for specific epistemic or social aims, concomitant with the openness to use a different concept (e.g., of race) for other purposes. We illustrate this approach by sketching three distinct concepts of gender and arguing that all of them are needed, as they answer to different social aims. The first concept serves the aim of identifying and explaining gender-based discrimination. It is similar to Haslanger’s well-known account, except that rather than offering a definition of ‘woman’ we focus on ‘gender’ as one among several axes of discrimination. The second concept of gender is to assign legal rights and social recognitions, and thus is to be trans-inclusive. We argue that this cannot be achieved by previously suggested concepts that include substantial gender-related psychological features, such as awareness of social expectations. Instead, our concept counts someone as being of a certain gender solely based on the person’s self-identification with this gender. The third concept of gender serves the aim of personal empowerment by means of one’s gender identity. In this context, substantial psychological features and awareness of one’s social situation are involved. While previous accounts of concepts have focused on their role in determining extensions, we point to contexts where a concept’s role in explanation and moral reasoning can be more important.

Philosophical discussions of systems biology have enriched the notion of mechanistic explanation by pointing to the role of mathematical modeling. However, such accounts still focus on explanation in terms of tracking a mechanism's operation across time (by means of mental or computational simulation). My contention is that there are explanations of molecular systems where the explanatory understanding does not consist in tracking a mechanism's operation and productive continuity. I make this case by a discussion of bifurcation analysis in dynamical systems, articulating the distinctive way in which explanatory understanding is provided, especially about the reversibility or irreversibility of molecular processes.

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.

We address the question of whether and to what extent explanatory and modelling strategies in systems biology are mechanistic. After showing how dynamic mathematical models are actually required for mechanistic explanations of complex systems, we caution readers against expecting all systems biology to be about mechanistic explanations. Instead, the aim may be to generate topological explanations that are not standardly mechanistic, or to arrive at design principles that explain system organization and behaviour in general, but not specific mechanisms. These abstraction strategies serve various aims, including prediction and control, that are central to understanding the epistemic diversity of systems biology.

The present paper discusses Kitcher’s framework for studying conceptual change and progress. Kitcher’s core notion of reference potential is hard to apply to concrete cases. In addition, an account of conceptual change as change in reference potential misses some important aspects of conceptual change and conceptual progress. I propose an alternative framework that focuses on the inferences and explanations supported by scientific concepts. The application of my approach to the history of the gene concept offers a better account of the conceptual progress that occurred in the transition from the Mendelian to the molecular gene than Kitcher’s theory.

The paper discusses reference determination from the point of view of conceptual change in science. The first part of the discussion uses the homology concept, a natural kind term from biology, as an example. It is argued that the causal theory of reference gives an incomplete account of reference determination even in the case of natural kind terms. Moreover, even if descriptions of the referent are taken into account, this does not yield a satisfactory account of reference in the case of the homology concept. I suggest that in addition to the factors that standard theories of reference invoke the scientific use of concepts and the epistemic interests pursued with concepts are important factors in determining the reference of scientific concepts. In the second part, I argue for a moderate holism about reference determination according to which the set of conditions that determine the reference of a concept is relatively open and different conditions may be reference fixing depending on the context in which this concept is used. It is also suggested that which features are reference determining in a particular case may depend on the philosophical interests that underlie reference ascription and the study of conceptual change.

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.

Despite the traditional focus on metaphysical issues in discussions of natural kinds in biology, epistemological considerations are at least as important. By revisiting the debate as to whether taxa are kinds or individuals, I argue that both accounts are metaphysically compatible, but that one or the other approach can be pragmatically preferable depending on the epistemic context. Recent objections against construing species as homeostatic property cluster kinds are also addressed. The second part of the paper broadens the perspective by considering homologues as another example of natural kinds, comparing them with analogues as functionally defined kinds. Given that there are various types of natural kinds, I discuss the different theoretical purposes served by diverse kind concepts, suggesting that there is no clear-cut distinction between natural kinds and other kinds, such as functional kinds. Rather than attempting to offer a unique metaphysical account of ‘natural’ kind, a more fruitful approach consists in the epistemological study of how different natural kind concepts are employed in scientific reasoning.

This essay analyzes and develops recent views about explanation in biology. Philosophers of biology have parted with the received deductive-nomological model of scientific explanation primarily by attempting to capture actual biological theorizing and practice. This includes an endorsement of different kinds of explanation (e.g., mathematical and causal-mechanistic), a joint study of discovery and explanation, and an abandonment of models of theory reduction in favor of accounts of explanatory reduction. Of particular current interest are philosophical accounts of complex explanations that appeal to different levels of organismal organization and use contributions from different biological disciplines. The essay lays out one model that views explanatory integration across different disciplines as being structured by scientific problems. I emphasize the philosophical need to take the explanatory aims pursued by different groups of scientists into account, as explanatory aims determine whether different explanations are competing or complementary and govern the dynamics of scientific practice, including interdisciplinary research. I distinguish different kinds of pluralism that philosophers have endorsed in the context of explanation in biology, and draw several implications for science education, especially the need to teach science as an interdisciplinary and dynamic practice guided by scientific problems and explanatory aims.

Essentialism is widely regarded as a mistaken view of biological kinds, such as species. After recounting why (sections 2-3), we provide a brief survey of the chief responses to the “death of essentialism” in the philosophy of biology (section 4). We then develop one of these responses, the claim that biological kinds are homeostatic property clusters (sections 5-6) illustrating this view with several novel examples (section 7). Although this view was first expressed 20 years ago, and has received recent discussion and critique, it remains underdeveloped and is often misrepresented by its critics (section 8).

Whereas an inference (deductive as well as inductive) is usually viewed as being valid in virtue of its argument form, the present paper argues that scientific reasoning is material inference, i.e., justified in virtue of its content. A material inference is licensed by the empirical content embodied in the concepts contained in the premises and conclusion. Understanding scientific reasoning as material inference has the advantage of combining different aspects of scientific reasoning, such as confirmation, discovery, and explanation. This approach explains why these different aspects (including discovery) can be rational without conforming to formal schemes, and why scientific reasoning is local, i.e., justified only in certain domains and contingent on particular empirical facts. The notion of material inference also fruitfully interacts with accounts of conceptual change and psychological theories of concepts.

The ‘death of evidence’ issue in Canada raises the spectre of politicized science, and thus the question of what role social values may have in science and how this meshes with objectivity and evidence. I first criticize philosophical accounts that have to separate different steps of research to restrict the influence of social and other non-epistemic values. A prominent account that social values may play a role even in the context of theory acceptance is the argument from inductive risk. It maintains that the more severe the social consequences of erroneously accepting a theory would be, the more evidence is needed before the theory may be accepted. However, an implication of this position is that increasing evidence makes the impact of social values converge to zero; and I argue for a stronger role for social values. On this position, social values may determine a theory’s conditions of adequacy, which among other things can include co..

The evolutionary embryologist Gavin Rylands de Beer can be viewed as one of the forerunners of modern evolutionary developmental biology in that he posed crucial questions and proposed relevant answers about the causal relationship between ontogeny and phylogeny. In his developmental approach to the phylogenetic phenomenon of homology, he emphasized that homology of morphological structures is to be identified neither with the sameness of the underlying developmental processes nor with the homology of the genes that are in involved in the development of the structures. De Beer’s work on developmental evolution focused on the notion of heterochrony, arguing that paedomorphosis increases morphological evolvability and is thereby an important mode of evolution that accounts for the origin of many taxa, including higher taxa.

The paper discusses concept individuation in the context of scientific concepts and conceptual change in science. It is argued that some concepts can be individuated in different ways. A particular term may be viewed as corresponding to a single concept. But at the same time, we can legitimately individuate in a more fine grained manner, i.e., this term can also be considered as corresponding to two or several concepts. The reason is that there are different philosophical and explanatory interests that underlie a particular study of the change of a scientific term. These interests determine how a concept is to be individuated; and as the same term can be subject to different philosophical studies and interests, its content can be individuated in different ways.

Edited by Alessandro Minelli and Thomas Pradeu, Towards a Theory of Development gathers essays by biologists and philosophers, which display a diversity of theoretical perspectives. The discussions not only cover the state of art, but broaden our vision of what development includes and provide pointers for future research. Interestingly, all contributors agree that explanations should not just be gene-centered, and virtually none use design and other engineering metaphors to articulate principles of cellular and organismal organization. I comment in particular on the issue of how to construe the notion of a ‘theory’ and whether developmental biology has or should aspire to have theories, which four of the contributions discuss in detail while taking opposing positions. Beyond construing a theory in terms of its empirical content, my aim is to shift the focus toward the role that theories have for guiding future scientific theorizing and practice. Such a conception of ‘theory’ is particularly important in the context of development, because arriving at a theoretical framework that provides guidance for the discipline of developmental biology as a whole is more plausible than a unified representation of development across all taxa.

Peculiar to Konrad Lorenz’s view of instinctive behavior is his strong innate-learned dichotomy. He claimed that there are neither ontogenetic nor phylogenetic transitions between instinctive and experience-based behavior components, thus contradicting all former accounts of instinct. The present study discusses how Lorenz came to hold this controversial position by examining the history of Lorenz’s early theoretical development in the crucial period from 1931 to 1937, taking relevant influences into account. Lorenz’s intellectual development is viewed as being guided by four theoretical and practical commitments as to how to study and explain behavior. These four factors, which were part of the general approach of Lorenz but not of other animal psychologists, were crucial in bringing about his specific position on instinctive behavior.

This article reviews the recent reissuing of Richard Owen’s On the Nature of Limbs and its three novel, introductory essays. These essays make Owen’s 1849 text very accessible by discussing the historical context of his work and explaining how Owen’s ideas relate to his larger intellectual framework. In addition to the ways in which the essays point to Owen’s relevance for contemporary biology, I discuss how Owen’s unity of type theory and his homology claims about fins and limbs compare with modern views. While the phenomena studied by Owen are nowadays of major interest to evolutionary developmental biology, research in evo-devo has largely shifted from homology (which was Owen’s concern) towards evolutionary novelty, e.g., accounting for fins as a novelty. Still, I argue that questions about homology are important and raise challenges even for explanations of novelty.