Homology

While the homology concept has taken on importance in thinking about the nature of psychological kinds, no one has shown how comparative psychological and behavioral evidence can distinguish between competing homology claims. I adapt the operational criteria of homology to accomplish this. I consider two competing homology claims that compare human anger with putative aggression systems of nonhuman animals, and demonstrate the effectiveness of these criteria in adjudicating between these claims.

A central topic in research at the intersection of development and evolution is the origin of novel traits. Despite progress on understanding how developmental mechanisms underlie patterns of diversity in the history of life, the problem of novelty continues to challenge researchers. Here we argue that research on evolutionary novelty and the closely associated phenomenon of co-option can be reframed fruitfully by: (1) specifying a conceptual model of mechanisms that underwrite character identity, (2) providing a richer and more empirically precise (...) notion of co-option that goes beyond common appeals to “deep homology”, and (3) attending to the nature of experimental interventions that can determine whether and how the co-option of identity mechanisms can help to explain novel character origins. This reframing has the potential to channel future investigation to make substantive progress on the problem of evolutionary novelty. To illustrate this potential, we apply our reframing to two case studies: treehopper helmets and beetle horns. (shrink)

Given the pervasiveness of gene sharing in evolution and the extent of homology across the tree of life, why is everything not homologous with everything else? The continuity and overlapping genetic contributions to diverse traits across lineages seem to imply that no discrete determination of homology is possible. Although some argue that the widespread overlap in parts and processes should be acknowledged as “partial” homology, this threatens a broad base of presumed comparative morphological knowledge accepted by most biologists. Following a (...) long scientific tradition, we advocate a strategy of “theoretical articulation” that introduces further distinctions to existing concepts to produce increased contrastive resolution among the labels used to represent biological phenomena. We pursue this strategy by drawing on successful patterns of reasoning from serial homology at the level of gene sequences to generate an enriched characterization of serial homology as a hierarchical, phylogenetic concept. Specifically, we propose that the concept of serial homology should be applied primarily to repeated but developmentally individualized body parts, such as cell types, differentiated body segments, or epidermal appendages. For these characters, a phylogenetic history can be reconstructed, similar to families of paralogous genes, endowing the notion of serial homology with a hierarchical, phylogenetic interpretation. On this basis, we propose a five-fold theoretical classification that permits a more fine-grained mapping of diverse trait-types. This facilitates answering the question of why everything is not homologous with everything else, as well as how novelty is possible given that any new character possesses evolutionary precursors. We illustrate the fecundity of our account by reference to debates over insect wing serial homologs and vertebrate paired appendages. (shrink)

The term ‘homology’ is persistently polysemous, defying the expectation that extensive scientific research should yield semantic stability. A common response has been to seek a unification of various prominent definitions. This paper proposes an alternative strategy, based on the insight that scientific concepts function as tools for research: When analyzing various conceptualizations of homology, we should preserve those distinguishing features that support particular research goals. We illustrate the fruitfulness of our strategy by application to two cases. First, we revisit E. (...) Ray Lankester’s celebrated (1870) evolutionary reappraisal of homology and argue that his analysis has been distorted by assimilation to modern agendas. His ‘homogeny’ does not mean the same thing as modern evolutionary ‘homology,’ and his ‘homoplasy’ is no mere antonym. Instead, Lankester uses both new terms to pose a question that remains strikingly relevant—how do mechanistic and historical causes of morphological resemblance interact? Second, we examine the puzzle of avian digit homology, which exemplifies disciplinary differences in homology conceptualization and assessment. Recent progress has been fueled by the development of new tools within the relevant disciplines (paleontology and developmental biology) and especially by increasing interdisciplinary cooperation. Conceptual unification has played very little role in this work, which instead seeks concrete evolutionary scenarios that integrate all the available evidence. Together these cases indicate the complex relationship between concepts and other tools in homology research. (shrink)

Serial homology, i.e., homology between repetitive structures in the same individual organism, is a debated concept in evolutionary developmental biology. The central question is the evolutionary interpretation of “sameness” in the context of the same body. This essay provides a synthetic analysis of the main issues involved in the debate, connecting conceptual problems with current experimental research. It is argued that a concept of serial homology that is not of the all-or-nothing kind can smooth several theoretical inconsistencies, while being more (...) in line with what we know about evolutionary change and the way we investigate it. (shrink)

Reconstructing ancestral species is a challenging endeavour: fossils are often scarce or enigmatic, and inferring ancestral characters based on novel molecular approaches has long been controversial. A key philosophical challenge pertinent at present is the lack of a theoretical framework capable of evaluating inferences of homology made through integration of multiple kinds of evidence. Here, I present just such a framework. I start with a brief history and critical assessment of attempts at inferring morphological homology through developmental genetics. I then (...) bring attention to a recent model of homology, namely Character Identity Mechanisms, intended partly to elucidate the relationships between morphological characters, developmental genetics, and homology. I utilise and build on this model to construct the evaluative framework mentioned above, which judges the epistemic value of evidence of each kind in each particular case based on three proposed criteria: effectiveness, admissibility, and informativity, as well as providing a generalised guideline on how it can be scientifically operationalised. I then point out the evolution of the eumetazoan body plan as a case in point where the application of this framework can yield satisfactory results, both empirically and conceptually. I will conclude with a discussion on some potential implications for more general philosophy of biology and philosophy of science, especially surrounding evidential integration, models and explanation, and reductionism. (shrink)

We describe an organizing framework for the study of infinitary combinatorics. This framework is Čech cohomology. It describes ZFC principles distinguishing among the ordinals of the form $\omega _n$. More precisely, this framework correlates each $\omega _n$ with an $$ -dimensional generalization of Todorcevic’s walks technique, and begins to account for that technique’s “unreasonable effectiveness” on $\omega _1$.We show in contrast that on higher cardinals $\kappa $, the existence of these principles is frequently independent of the ZFC axioms. Finally, we (...) detail implications of these phenomena for the computation of strong homology groups and higher derived limits, deriving independence results in algebraic topology and homological algebra, respectively, in the process.Abstract prepared by Jeffrey Bergfalk.E-mail: [email protected]. (shrink)

This chapter examines examples from the last half century of foodFood as a rhetorical homologyRhetorical homology—equating foodFoodcultureCultureandFood cultures activism and political discourse. The Black Panther PartyBlack panther party’s Free Breakfast for Children Program is compared to the contemporary indigenous group the I-Collective, whose identityIdentity and activism feature foodFood-focused content both virtually and in person. The two case studies illuminate the powerPower of foodFood as a broader rhetorical tool brandished today by activists and politicians alike. While foodFood has long been used (...) as a political tool, the Black Panther PartyBlack panther party’s Free Breakfast for Children Program is one of the most potent examples of foodFood involving explicit political discourse. Their 1969 national anti-hunger initiative began as a way to make a positive difference in the lives of marginalised children, but also functioned to communicate valuesValue, intention, and political ideology, linking discourses and creating what Barry Brummett called a “rhetorical homologyRhetorical homology”. With the rising influence of social mediaSocial media, and the popularity of foodFood on these platforms we see this happening again in modern discourse. The indigenous activist group I-Collective is a contemporary example of a group using foodFoodas political rhetoricFood rhetoric in a similarly explicit way as did the Black Panther PartyBlack panther party a half century earlier. (shrink)

A given explanatory theory T falls into circular reasoning if the only way to determine its explanandum is through the application of T. To find an underlying theory T′ that determines T′s explanandum helps us save T from this accusation of circularity. We follow the structuralist view of theories in presenting and dealing with this issue, by applying it to particular theories. More specifically, we focus on the relationship between the Darwinian theory of common ancestry and the determination of homologies.

This project begins with the selective sensory experience suggested by lngarden followed by an insensitivity he insinuates to digestive processes. This is juxtaposed with an oenological explanation of phenomenal sedimentation offered by Jean-Luc Marion. It compares the dynamics of time in the former with the those of wine in the latter. Emphasis is given to lngarden's insinuation of time as fluid, liquid, or aquatic. It revisits Ingarden's physiological explanations of partially-open systems by way of the bilateral excretion and absorption of (...) semi-permeable cellular membranes. The importance he eventually grants to inner secretion is considered alongside perspiration and salivation collateral to skin and membranes. It suggests that Ingarden's interest in thermoregulation, partial permeation, and secretion invites alternative conceptions of temporal consciousness in physiological experiences, beyond sequential and linear clock-time and/or Kantian intuition. Temporality experienced as temperance becomes discernible at a permeable point in which the sedimentation of Husserl, the maturation of Marion, and the fluidity and secretion of Ingarden mix and mingle into the taste of time. (shrink)

In this article, I examine the issue of the alleged circularity in the determination of homologies within cladistic analysis. More specifically, I focus on the claims made by the proponents of the dynamic homology approach, regarding the distinction (sometimes made in the literature) between primary and secondary homology. This distinction is sometimes invoked to dissolve the circularity issue, by upholding that characters in a cladistic data matrix have to be only primarily homologous, and thus can be determined independently of phylogenetic (...) hypotheses, by using the classical Owenian criteria (for morphological characters) or via multiple sequence alignment (for sequence data). However, since in the dynamic approach, sequence data can be analyzed without being pre-aligned, proponents have claimed that the distinction between primary and secondary homology has no place within cladistics. I will argue that this is not the case, since cladistic practice within the dynamic framework does presuppose primary homology statements at a higher level. (shrink)

This book offers a novel defence of a highly contested philosophical position: biological natural kind essentialism. This theory is routinely and explicitly rejected for its purported inability to be explicated in the context of contemporary biological science, and its supposed incompatibility with the process and progress of evolution by natural selection. Christopher J. Austin challenges these objections, and in conjunction with contemporary scientific advancements within the field of evolutionary-developmental biology, the book utilises a contemporary neo-Aristotelian metaphysics of "dispositional properties", or (...) causal powers, to provide a theory of essentialism centred on the developmental architecture of organisms and its role in the evolutionary process. By defending a novel theory of Aristotelian biological natural kind essentialism, Essence in the Age of Evolution represents the fresh and exciting union of cutting-edge philosophical insight and scientific knowledge. (shrink)

Bence Nanay has argued that we must abandon the etiological theory of teleological function because this theory explains functions and functional categories in a circular manner. Paul Griffiths argued earlier that we should retain the etiological theory and instead prevent the circularity by making etiologies independent of functional categories. Karen Neander and Alex Rosenberg reply to Nanay similarly, and argue that we should analyze functions in terms of natural selection acting not on functional categories, but merely on lineages. Nanay replies (...) that these lineages cannot be individuated except by reference to functional categories. Worryingly, Neander and Rosenberg themselves have previously argued persuasively that homology often depends on function. This article addresses their arguments and shows how to escape them: Regardless whether the arguments are right about long-term homological categories, they do not apply to generation-to-generation homology. The latter, moreover, is sufficient for individuating the lineages needed to explain teleological functions. (shrink)

There is long-standing conflict between genealogical and developmental accounts of homology. This paper provides a general framework that shows that these accounts are compatible and clarifies precisely how they are related. According to this framework, understanding homology requires both an abstract genealogical account that unifies the application of the term to all types of characters used in phylogenetic systematics and locally enriched accounts that apply only to specific types of characters. The genealogical account serves this unifying role by relying on (...) abstract notions of ‘descent’ and ‘character’. As a result, it takes for granted the existence of such characters. This requires theoretical justification that is provided by enriched accounts, which incorporate the details by which characters are inherited. These enriched accounts apply to limited domains, providing the needed theoretical justification for recognizing characters within that domain. Though connected to the genealogical account of homology in this way, enriched accounts include phenomena that fall outside the scope of the genealogical account. They therefore overlap, but are not nested within, the genealogical account. Developmental accounts of homology are to be understood as enriched accounts of body part homology. Once they are seen in this light, the conflict with the genealogical account vanishes. It is only by understanding the fine conceptual structure undergirding the many uses of the term ‘homology’ that we can understand how these uses hang together. (shrink)

Understanding the organization of an organism by individuating meaningful parts and accounting for organismal properties by studying the interaction of bodily parts is a central practice in many areas of biology. While structures are obvious bodily parts and structure and function have often been seen as antagonistic principles in the study of organismal organization, my tenet is that structures and functions are on a par. I articulate a notion of function (functions as activities), according to which functions are bodily parts (...) just as structures are. Recognizing part-whole relations among an organism’s various structures and functions permits fruitful investigation and multilevel explanation of organismal properties and functioning, across both developmental and evolutionary time. I show how my perspective clarifies debates surrounding homology and evolutionary novelty that stem from an alleged structure-function dichotomy. My approach favors a pluralism about individuation, where the criteria of what counts as a meaningful bodily part depend on the particular epistemic aims pursued in a scientific context. (shrink)

Among philosophical analyses of Darwin’s Origin, a standard view says the theory presented there had no concrete observational consequences against which it might be checked. I challenge this idea with a new analysis of Darwin’s principal geographical distribution observations and how they connect to his common ancestry hypothesis.

Homology is a natural kind concept, but one that has been notoriously elusive to pin down. There has been sustained debate over the nature of correspondence and the units of comparison. But this continued debate over its meaning has focused on defining homology rather than on its use in practice. The aim of this chapter is to concentrate on the practices of homologizing. I define “homologizing” to be a concept-in-use. Practices of homologizing are kinds of rule following, the satisfaction of (...) which demarcates a category—that of being a homologue. Identifying, explaining, discovering, and understanding are exchanges that connect practice to concept through the performance of a rule by practitioners. These practices are constitutive of natural kinding activities. If homologizing is a kind of kinding, consideration of these practices of discovery, tracking, and identification not only clarifies the meaning, use, and progression of the concept of homology, but provides further understanding of the processes and progression of natural kinds and kinding practices in general. (shrink)

This edited volume of 13 new essays aims to turn past discussions of natural kinds on their head. Instead of presenting a metaphysical view of kinds based largely on an unempirical vantage point, it pursues questions of kindedness which take the use of kinds and activities of kinding in practice as significant in the articulation of them as kinds. The book brings philosophical study of current and historical episodes and case studies from various scientific disciplines to bear on natural kinds (...) as traditionally conceived of within metaphysics. Focusing on these practices reveals the different knowledge-producing activities of kinding and processes involved in natural kind use, generation, and discovery. -/- Specialists in their field, the esteemed group of contributors use diverse empirically responsive approaches to explore the nature of kindhood. This groundbreaking volume presents detailed case studies that exemplify kinding in use. Newly written for this volume, each chapter engages with the activities of kinding across a variety of disciplines. Chapter topics include the nature of kinds, kindhood, kinding, and kind-making in linguistics, chemical classification, neuroscience, gene and protein classification, colour theory in applied mathematics, homology in comparative biology, sex and gender identity theory, memory research, race, extended cognition, symbolic algebra, cartography, and geographic information science. -/- The volume seeks to open up an as-yet unexplored area within the emerging field of philosophy of science in practice, and constitutes a valuable addition to the disciplines of philosophy and history of science, technology, engineering, and mathematics. -/- Contributions from a diverse group of established and junior scholars in the fields of Philosophy and History and Philosophy of Science including Hasok Chang, Jordi Cat, Sally Haslanger, Joyce C. Havstad, Catherine Kendig, Bernhard Nickel, Josipa Petrunic, Samuli Pöyhönen, Thomas A. C. Reydon, Quayshawn Spencer, Jackie Sullivan, Michael Wheeler, and Rasmus Grønfeldt Winther. (shrink)

The origination of novel structures has long been an intriguing topic for biologists. Over the past few decades it has served as a central theme in evolutionary developmental biology. Yet, definitions of evolutionary innovation and novelty are frequently debated and there remains disagreement about what kinds of causal factors best explain the origin of qualitatively new variation in the history of life. Here we examine aspects of these debates, survey three empirical case studies, and reflect on directions for future inquiry (...) that will advance research into the developmental evolution of novel structures. (shrink)

There have been several recent attempts to think about psychological kinds as homologies. Nevertheless, there are serious epistemic challenges for individuating homologous psychological kinds, or cognitive homologies. Some of these challenges are revealed when we look at competing claims of cognitive homology. This paper considers two competing homology claims that compare human anger with putative aggression systems of nonhuman animals. The competition between these hypotheses has been difficult to resolve in part because of what I call the boundary problem: boundaries (...) between instances of psychological kinds (e.g., anger and fear) cannot be directly observed. Thus, there are distinctive difficulties for individuating psychological kinds across lineages. I draw four conclusions from this case study: First, recent evidence from the neuroscience of fear suggests that one of the proposed homologies involves a straightforward conflation of anger and fear. Second, this conflation arises because of the boundary problem. Third, there is an implicit constraint on the operational criteria that is easy to overlook in the psychological case. In this case, ignoring the constraint is part of the problem. Fourth, this is a clear case in which knowledge of homology cannot be accumulated piecemeal. Identifying homologs of human anger requires identifying homologs of fear. (shrink)

The Mind-Brain Identity Theory lived a short life as a respectable philosophical position in the late 1950s, until Hilary Putnam developed his famous argument on the multiple realizability of mental states. The argument was, and still is, taken as the definitive demonstration of the falsity of Identity Theory and the foundation on which contemporary functionalist computational cognitive science was to be grounded. In this paper, in the wake of some contemporary philosophers, we reopen the case for Identity Theory and offer (...) a solution to the problem of multiple realizabilty. The solution is based on the necessity, at the time of establishing identity relations, of appealing to the notions of “homology” and “analogy” developed in the nineteenth century by Richard Owen. We also suggest that these notions are useful in order to correct certain shortcomings of some recent attempts at rebutting the Multiple Realizability argument. (shrink)

Günter Wagner’s Homology, Genes, and Evolutionary Innovation is a compelling, and empirically well-supported account of the evolution of character identity and character origination which emphasizes the importance of homology and novelty as central explananda for 21st century evolutionary biology. In this essay review, I focus on the similarities and differences between the structuralist picture of evolutionary biology advocated by Wagner, and that presented by standard evolutionary theory. First, I outline the ways in which Wagner’s genetic theory of homology diverges from (...) the account of homology offered by standard evolutionary theory. Then, I consider the motivations for these divergences. Lastly, I discuss a number of concerns with Wagner’s view, and offer some concluding thoughts on the relationship between structuralism and adaptationism. (shrink)

Günter Wagner’s Homology, Genes, and Evolutionary Innovation collects and synthesizes a vast array of empirical data, theoretical models, and conceptual analysis to set out a progressive research program with a central theoretical commitment: the genetic theory of homology. This research program diverges from standard approaches in evolutionary biology, provides sharpened contours to explanations of the origin of novelty, and expands the conceptual repertoire of evolutionary developmental biology. I concentrate on four aspects of the book in this essay review: the genetic (...) theory of homology and character identity networks; the implications for how we explain evolutionary novelties; the expanded set of concepts surrounding homology, and the epistemological conflicts between Wagner’s viewpoint and functionally-oriented evolutionary biology, as well as differences with other Evo–devo researchers. Together these have ramifications for how we interpret different explanatory approaches to evolutionary phenomena and understand relationships between the usefulness of concepts and the reality they represent. (shrink)

This paper responds to the essay reviews by David Haig, Alan Love and Rachel Brown of my recently published book “Homology, Genes and Evolutionary Innovation”. The issues addressed here relate to: the notion of classes and individuals, issues of explanatory value of adaptive and structuralist explanations in evolutionary biology, the role of homology in evolutionary theory, the limits of a pluralist stance vis a vis alternative explanations of homology, as well as the question whether and to what extend the perspective (...) laid out in HGEI can be or should be transferred to other branches of study, like comparative behavioral biology. (shrink)

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

Recent work on inheritance systems can be divided into inclusive conceptions, according to which genetic and non-genetic inheritance are both involved in the development and transmission of nearly all animal behavioral traits, and more demanding conceptions of what it takes for non-genetic resources involved in development to qualify as a distinct inheritance system. It might be thought that, if a more stringent conception is adopted, homologies could not subsist across two distinct inheritance systems. Indeed, it is commonly assumed that homology (...) relations cannot survive a shift between genetic and cultural inheritance systems, and substantial reliance has been placed on that assumption in debates over the phylogenetic origins of hominin behavioral traits, such as male-initiated intergroup aggression. However, in the homology literature it is widely accepted that a trait can be homologous—that is, inherited continuously in two different lineages from a single common ancestor—despite divergence in the mechanisms involved in the trait’s development in the two lineages. In this paper, we argue that even on an extremely stringent understanding of what it takes for developmental resources to form a separate inheritance system, homologies can nonetheless subsist across shifts between distinct inheritance systems. We argue that this result is a merit of this way of characterizing what it is to be an inheritance system, that it has implications for adjudicating between alternative accounts of homology, and that it offers an important cautionary lesson about how to reason with the homology concept, particularly in the context of cultural species. (shrink)

While it is generally agreed that the concept of homology refers to individuated traits that have been inherited from common ancestry, we still lack an adequate account of trait individuation or inheritance. Here I propose that we utilize a counterfactual criterion of causation to link each trait with a developmental-causal (DC) gene. A DC gene is made up of the genetic information (which might or might not be physically contiguous in the genome) that is needed for the production of the (...) organismic attributes that comprise the trait. I argue that individuated traits—phenes—correspond to organismic features that are caused by DC genes. Using such an approach, we can define a DC map, which shows the relations between each pair of phenes and provides a succinct summary of genotype-phenotype relationships and phenotypic complexity. Phenes in parents and offspring are judged to be homologous if their DC genes are composed of orthologous genetic factors. When comparing more distantly related organisms, traits are homologous when linked by a chain of parent-offspring homologs along the path of ancestry that links the two organisms. There are three possible ways to deal with the potential for multiple equivalent DC genes: maximal, minimal, and consensus homology. Whereas maximal homology has limited utility, the other two approaches have value and can help to guide research at the intersection of evolution and development. (shrink)

The comparative method grants epistemic access to the biological past. Comparing lineages provides empirical traction on both hypotheses about particular lineages and models of trait evolution. Understanding this evidential role is important. Although philosophers have recently turned their attention to relations of descent, little work exists exploring the status of evidence from convergences. I argue that, where they exist, convergences play a central role in the confirmation of adaptive hypotheses. I focus on ‘analogous inferences’, show how such inferences ought to (...) be analysed and suggest three methods for strengthening their evidential weight. 1 Introduction2 Analogous Inferences2.1 Adaptive explanations and analogies2.2 Analogous inferences2.3 Scope, grain, and specificity3 Parallel Modelling, Integrated Explanations, and Convergent Modelling3.1 Parallel modelling3.2 Integrated explanations3.1 Convergent modelling4 Conclusion. (shrink)

The ‘byproduct account’ of female orgasm, a subject of renewed debate since Lloyd (The case of the female orgasm, Harvard University Press, Cambridge, 2005), is universally attributed to Symons (The evolution of human sexuality, Oxford University Press, Oxford, 1979). While this is correct to the extent that he linked it to the adaptive value of male orgasm, I argue that the attribution of the theory as we understand it to Symons is based on a serious and hitherto unrecognised misinterpretation. Symons (...) had a different explanation of why women can orgasm, and beneath this explanation lies an obscure line of argument, including a particularly obscure use of the word ‘homologous’. (shrink)

The aim of this article is to detail some reservations against the beliefs, claims, or presuppositions that current essentialist natural kind concepts (including homeostatic property cluster kinds) model grouping practices in the life sciences accurately and generally. Such concepts fit reasoning into particular preconceived epistemic and semantic patterns. The ability of these patterns to fit scientific practice is often argued in support of homeostatic property cluster accounts, yet there are reasons to think that in the life sciences kind concepts exhibit (...) a diversity of grouping practices that are flattened out by conceptualizing them as natural kinds. Instead this article argues that the process of understanding grouping practices needs to start from a more neutral position independent of any ontological account. Following Love (Acta Biotheor 57:51–75, 2009) this paper suggests that typical natural kind concepts should be broached in the first place as grouping strategies that use a variety of semantic and epistemic tactics to apply group-bound information to tasks of explanation and understanding. (shrink)

This paper explores an important type of biological explanation called ‘homology thinking.’ Homology thinking explains the properties of a homologue by citing the history of a homologue. Homology thinking is significant in several ways. First, it offers more detailed explanations of biological phenomena than corresponding analogy explanations. Second, it provides an important explanation of character similarity and difference. Third, homology thinking offers a promising account of multiple realizability in biology.

Homology is a biological sameness relation that is purported to hold in the face of changes in form, composition, and function. In spite of the centrality and importance of homology, there is no consensus on how we should understand this concept. The two leading views of homology, the genealogical and developmental accounts, have significant shortcomings. We propose a new account, the hierarchical-dependency account of homology, which avoids these shortcomings. Furthermore, our account provides for continuity between special, general, and serial homology.

Homology is a central concept of comparative and evolutionary biology, referring to the presence of the same bodily parts (e.g., morphological structures) in different species. The existence of homologies is explained by common ancestry, and according to modern definitions of homology, two structures in different species are homologous if they are derived from the same structure in the common ancestor. Homology has traditionally been contrasted with analogy, the presence of similar traits in different species not necessarily due to common ancestry (...) but due to a similar function or convergent evolution resulting from similar selective pressure in different species. (A more recent contrastive notion is homoplasy, the presence of similar traits in different species without common ancestry, i.e., as an instance of parallel evolution.) This sounds straightforward, but in fact the homology concept has a rich history and currently is the subject of extensive theoretical reflection, resulting in different contemporary approaches to homology. (shrink)

Scientists exhibit different styles in their reasoning about the natural world (e.g., experimental, historical, or statistical). These styles have been characterized, categorized, and combined in many ways throughout the history of science.

Many of the current comparisons of taxic phylogenetic and biological homology in the context of morphology focus on what are seen as categorical distinctions between the two concepts. The first, it is claimed, identifies historical patterns of conservation and variation relating taxa; the second provides a causal framework for the explanation of this conservation and variation. This leads to the conclusion that the two need not be placed in conflict and are in fact compatible, having non-competing epistemic purposes or mapping (...) the same extensions in the form of monophyletic groupings (see Roth, The biological basis of homology 1–26, 1988; Sluys, J Zool Syst Evol Res 34:145–152, 1996; Abouheif, Trends Ecol Evol 12:405–408, 1997; Brigandt, J Exp Zool 299:9–17, 2003, Biol Philos 22:709–725, 2007; Assis and Brigandt, Evol Biol 36:248–255, 2009). This article argues that moves in this direction miss the essential disagreement between these concepts as they have been developed in the context of the debate concerning the best concept for evolutionary investigation. We should rather see these concepts employing a common fundamental methodological approach to homology, but disagreeing about how to apply the methodology effectively. Both concepts employ class reasoning, which pursues homologies as units of generalization—more precisely, as sources of reliable and relevant group-bound information in the form of shared underlying causes. The dispute can be better understood by two poles that structure such reasoning: the need for a reliable basis for projections about the causal history of shared structures, and the desire to identify homologous characters with more informative and specific causal information relevant to generalizing about evolutionary processes. Judgments in favor of one or the other in turn have affected the scope or extension of these competing homology concepts. (shrink)

Planctomycetes, Verrucomicrobia and Chlamydia are prokaryotic phyla, sometimes grouped together as the PVC superphylum of eubacteria. Some PVC species possess interesting attributes, in particular, internal membranes that superficially resemble eukaryotic endomembranes. Some biologists now claim that PVC bacteria are nucleus‐bearing prokaryotes and are considered evolutionary intermediates in the transition from prokaryote to eukaryote. PVC prokaryotes do not possess a nucleus and are not intermediates in the prokaryote‐to‐eukaryote transition. Here we summarise the evidence that shows why all of the PVC traits (...) that are currently cited as evidence for aspiring eukaryoticity are either analogous (the result of convergent evolution), not homologous, to eukaryotic traits; or else they are the result of horizontal gene transfers. (shrink)

In the last 10 years, several authors including Griffiths and Matthen have employed classificatory principles from biology to argue for a radical revision in the way that we individuate psychological traits. Arguing that the fundamental basis for classification of traits in biology is that of ‘homology’ (similarity due to common descent) rather than ‘analogy’, or ‘shared function’, and that psychological traits are a special case of biological traits, they maintain that psychological categories should be individuated primarily by relations of homology (...) rather than in terms of shared function. This poses a direct challenge to the dominant philosophical view of how to define psychological categories, viz., ‘functionalism’. Although the implications of this position extend to all psychological traits, the debate has centered around ‘emotion’ as an example of a psychological category ripe for reinterpretation within this new framework of classification. I address arguments by Griffiths that emotions should be divided into at least two distinct classes, basic emotions and higher cognitive emotions, and that these two classes require radically different theories to explain them. Griffiths argues that while basic emotions in humans are homologous to the corresponding states in other animals, higher cognitive emotions are dependent on mental capacities unique to humans, and are therefore not homologous to basic emotions. Using the example of shame, I argue that (a) many emotions that are commonly classified as being higher cognitive emotions actually correspond to certain basic emotions, and that (b) the “higher cognitive forms” of these emotions are best seen as being homologous to their basic forms. (shrink)

Most cognitive scientists nowadays tend to think that at least some of the mind’s capacities are the product of biological evolution, yet important conceptual problems remain for all scientists in order to be able to speak coherently of mental or cognitive systems as having evolved naturally. Two of these important problems concern the articulation of adequate, interesting, and empirically useful concepts of homology and variation as applied to cognitive systems. However, systems in cognitive science are usually understood as functional systems (...) of some sort. Thus, to be able to talk about functional systems being homologous requires having a solid, adequate, and empirically articulated concept of functional homology—and the same is true about functional variation. Here I construct an original concept of functional homology that, in my view, adequately systematizes a number of actual uses of the word “functional homology” in a variety of biological disciplines and in ethology. I also propose a number of criteria for the empirical application of the concept that are analogous to the criteria that are currently used in comparative biology, ethology, and molecular developmental genetics. Then I construct a concept of functional variation on the basis of this concept of homology. (shrink)

The function of a trait token is usually defined in terms of some properties of other (past, present, future) tokens of the same trait type. I argue that this strategy is problematic, as trait types are (at least partly) individuated by their functional properties, which would lead to circularity. In order to avoid this problem, I suggest a way to define the function of a trait token in terms of the properties of the very same trait token. To able to (...) allow for the possibility of malfunctioning, some of these properties need to be modal ones: a function of a trait is to do F just in case its doing F would contribute to the inclusive fitness of the organism whose trait it is. Function attributions have modal force. Finally, I explore whether and how this theory of biological function could be modified to cover artifact function. (shrink)

Over the past decade, it has been discovered that disparate aspects of morphology – often of distantly related groups of organisms – are regulated by the same genetic regulatory mechanisms. Those discoveries provide a new perspective on morphological evolutionary change. A conceptual framework for exploring these research findings is termed ‘deep homology’. A comparative framework for morphological relations of homology is provided that distinguishes analogy, homoplasy, plesiomorphy and synapomorphy. Four examples – three from plants and one from animals – demonstrate (...) that homologous developmental mechanisms can regulate a range of morphological relations including analogy, homoplasy and examples of uncertain homology. Deep homology is part of a much wider range of phenomena in which biological (genes, regulatory mechanisms, morphological traits) and phylogenetic levels of homology can both be disassociated. Therefore, to understand homology, precise, comparative, independent statements of both biological and phylogenetic levels of homology are necessary. (shrink)

Taxa and homologues can in our view be construed both as kinds and as individuals. However, the conceptualization of taxa as natural kinds in the sense of homeostatic property cluster kinds has been criticized by some systematists, as it seems that even such kinds cannot evolve due to their being homeostatic. We reply by arguing that the treatment of transformational and taxic homologies, respectively, as dynamic and static aspects of the same homeostatic property cluster kind represents a good perspective for (...) supporting the conceptualization of taxa as kinds. The focus on a phenomenon of homology based on causal processes (e.g., connectivity, activity-function, genetics, inheritance, and modularity) and implying relationship with modification yields a notion of natural kinds conforming to the phylogenetic-evolutionary framework. Nevertheless, homeostatic property cluster kinds in taxonomic and evolutionary practice must be rooted in the primacy of epistemological classification (homology as observational properties) over metaphysical generalization (series of transformation and common ancestry as unobservational processes). The perspective of individuating characters exclusively by historical-transformational independence instead of their developmental, structural, and functional independence fails to yield a sufficient practical interplay between theory and observation. Purely ontological and ostensional perspectives in evolution and phylogeny (e.g., an ideographic character concept and PhyloCode’s ‘individualism’ of clades) may be pragmatically contested in the case of urgent issues in biodiversity research, conservation, and systematics. (shrink)

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. (shrink)

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. (shrink)

I attempt to raise questions regarding elements of systematics—primarily in the realm of phylogenetic reconstruction—in order to provoke discussion on the current state of affairs in this discipline, and also evolutionary biology in general: e.g., conceptions of homology and homoplasy, hypothesis testing, the nature of and objections to Hennigian “phylogenetic systematics”, and the schism between Darwinian descendants of the “modern evolutionary synthesis” and their supposed antagonists, cladists and punctuationalists.