No full-text available
To read the full-text of this research,
you can request a copy directly from the authors.
Against unifying homology concepts: Redirecting the debate
Abstract
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 Lankester's celebrated 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.
... The question about what homology is should, however, be complemented by the question how we adequately treat homology as term and concept(s) so ubiquitous in our daily work. With no unification in sight, obviously no authority forcing a certain concept upon us, and maybe not even the desire/use for unification (Cracraft, 2023;Gouveâ and Brigandt, 2023;Minelli, 2023), I attempt to make a case for conceptual explicitness similar to what is advocated by the ontology community. ...
... As becomes obvious from the usage of homology and similar terms in the literature, the linguistic problem of morphology (Vogt et al., 2010) does not only apply to terms used to refer to morphological entities but in fact to the very terms and concepts building the conceptual framework of morphology. Homology, a term so central and so frequently used, is a polyseme given as label to numerous different concepts (Brigandt, 2003;Ballego-Campos et al., 2023;Gouveâ and Brigandt, 2023); incongruence of homology statements is inevitable. Attempts of unification will unlikely gain universal acceptance. ...
Morphology, the scientific discipline dealing with description and comparison of organismal form, is one of the oldest disciplines in biology and traditionally strongly linked to the concept of homology. With morphological data being used and morphological knowledge being applied in other (younger) biological disciplines, morphology has often been degraded to an only auxiliary discipline or a mere set of methods serving those other disciplines. While this notion has been wrong all along, the last decades have seen a renaissance of morphology mostly due to significant leaps in imaging techniques and the advent of 3D digital data. Modern large-scale morphological endeavors in what is called phenomics and new means of functional analyses underline the fruitfulness of morphological research. Furthermore, morphology has been revisited on a conceptual level leading to a “re-philosophication” of morphology acknowledging its nature as explanatory science. Based on Richter and Wirkner’s research program of Evolutionary Morphology, this essay expands the conceptual framework to identify entities and processes vital for morphology as independent scientific discipline. With no unified homology concept in sight (and maybe not even desired), following the emergence of bio-ontologies in morphology, a plea is made for conceptual explicitness which acknowledges the plurality of homology concepts but enables intersubjective transfer.
... The concept was further genealogically developed by Ray Lankester in 1870, who tried to replace the term of homology with a more mechanistic name, 'homogeny' [avoiding the idealistic meaning of "ology" -see (Gouvêa and Brigandt, 2023)]. "Structures which are genetically related, in so far as they have a single representative in a common ancestor, may be called homogenous"- (Lankester, 1870). ...
... The name 'homogeny' did not survive in scientific literature. Lankester also coined the term homoplasy: the similarity of traits not due to common ancestry (see historical summary in (Gouvêa and Brigandt, 2023). Thus, homoplasy is the alternative scenario to the 'homology' hypothesis-non-homologyimplying convergent evolution. ...
Here, we provide a brief historical overview of the homology concept and then will discuss its applications to diverse nervous systems of invertebrates targeting the level of individual functionally characterized neurons, controlling specific behaviors.
Essays on evolvability from the perspectives of quantitative and population genetics, evolutionary developmental biology, systems biology, macroevolution, and the philosophy of science.
Evolvability—the capability of organisms to evolve—wasn't recognized as a fundamental concept in evolutionary theory until 1990. Though there is still some debate as to whether it represents a truly new concept, the essays in this volume emphasize its value in enabling new research programs and facilitating communication among the major disciplines in evolutionary biology. The contributors, many of whom were instrumental in the development of the concept of evolvability, synthesize what we have learned about it over the past thirty years. They focus on the historical and philosophical contexts that influenced the emergence of the concept and suggest ways to develop a common language and theory to drive further evolvability research.
The essays, drawn from a workshop on evolvability hosted in 2019–2020 by the Center of Advanced Study at the Norwegian Academy of Science and Letters, in Oslo, provide scientific and historical background on evolvability. The contributors represent different disciplines of evolutionary biology, including quantitative and population genetics, evolutionary developmental biology, systems biology, and macroevolution, as well as the philosophy of science. This plurality of approaches allows researchers in disciplines as diverse as developmental biology, molecular biology, and systems biology to communicate with those working in mainstream evolutionary biology. The contributors also discuss key questions at the forefront of research on evolvability.
Contributors:J. David Aponte, W. Scott Armbruster, Geir H. Bolstad, Salomé Bourg, Ingo Brigandt, Anne Calof, James M. Cheverud, Josselin Clo, Frietson Galis, Mark Grabowski, Rebecca Green, Benedikt Hallgrímsson, Thomas F. Hansen, Agnes Holstad, David Houle, David Jablonski, Arthur Lander, Arnaud LeRouzic, Alan C. Love, Ralph Marcucio, Michael B. Morrissey, Laura Nuño de la Rosa, Øystein H. Opedal, Mihaela Pavličev, Christophe Pélabon, Jane M. Reid, Heather Richbourg, Jacqueline L. Sztepanacz, Masahito Tsuboi, Cristina Villegas, Marta Vidal-García, Kjetil L. Voje, Andreas Wagner, Günter P. Wagner, Nathan M. Young
Essays on evolvability from the perspectives of quantitative and population genetics, evolutionary developmental biology, systems biology, macroevolution, and the philosophy of science.
Evolvability—the capability of organisms to evolve—wasn't recognized as a fundamental concept in evolutionary theory until 1990. Though there is still some debate as to whether it represents a truly new concept, the essays in this volume emphasize its value in enabling new research programs and facilitating communication among the major disciplines in evolutionary biology. The contributors, many of whom were instrumental in the development of the concept of evolvability, synthesize what we have learned about it over the past thirty years. They focus on the historical and philosophical contexts that influenced the emergence of the concept and suggest ways to develop a common language and theory to drive further evolvability research.
The essays, drawn from a workshop on evolvability hosted in 2019–2020 by the Center of Advanced Study at the Norwegian Academy of Science and Letters, in Oslo, provide scientific and historical background on evolvability. The contributors represent different disciplines of evolutionary biology, including quantitative and population genetics, evolutionary developmental biology, systems biology, and macroevolution, as well as the philosophy of science. This plurality of approaches allows researchers in disciplines as diverse as developmental biology, molecular biology, and systems biology to communicate with those working in mainstream evolutionary biology. The contributors also discuss key questions at the forefront of research on evolvability.
Contributors:J. David Aponte, W. Scott Armbruster, Geir H. Bolstad, Salomé Bourg, Ingo Brigandt, Anne Calof, James M. Cheverud, Josselin Clo, Frietson Galis, Mark Grabowski, Rebecca Green, Benedikt Hallgrímsson, Thomas F. Hansen, Agnes Holstad, David Houle, David Jablonski, Arthur Lander, Arnaud LeRouzic, Alan C. Love, Ralph Marcucio, Michael B. Morrissey, Laura Nuño de la Rosa, Øystein H. Opedal, Mihaela Pavličev, Christophe Pélabon, Jane M. Reid, Heather Richbourg, Jacqueline L. Sztepanacz, Masahito Tsuboi, Cristina Villegas, Marta Vidal-García, Kjetil L. Voje, Andreas Wagner, Günter P. Wagner, Nathan M. Young
Essays on evolvability from the perspectives of quantitative and population genetics, evolutionary developmental biology, systems biology, macroevolution, and the philosophy of science.
Evolvability—the capability of organisms to evolve—wasn't recognized as a fundamental concept in evolutionary theory until 1990. Though there is still some debate as to whether it represents a truly new concept, the essays in this volume emphasize its value in enabling new research programs and facilitating communication among the major disciplines in evolutionary biology. The contributors, many of whom were instrumental in the development of the concept of evolvability, synthesize what we have learned about it over the past thirty years. They focus on the historical and philosophical contexts that influenced the emergence of the concept and suggest ways to develop a common language and theory to drive further evolvability research.
The essays, drawn from a workshop on evolvability hosted in 2019–2020 by the Center of Advanced Study at the Norwegian Academy of Science and Letters, in Oslo, provide scientific and historical background on evolvability. The contributors represent different disciplines of evolutionary biology, including quantitative and population genetics, evolutionary developmental biology, systems biology, and macroevolution, as well as the philosophy of science. This plurality of approaches allows researchers in disciplines as diverse as developmental biology, molecular biology, and systems biology to communicate with those working in mainstream evolutionary biology. The contributors also discuss key questions at the forefront of research on evolvability.
Contributors:J. David Aponte, W. Scott Armbruster, Geir H. Bolstad, Salomé Bourg, Ingo Brigandt, Anne Calof, James M. Cheverud, Josselin Clo, Frietson Galis, Mark Grabowski, Rebecca Green, Benedikt Hallgrímsson, Thomas F. Hansen, Agnes Holstad, David Houle, David Jablonski, Arthur Lander, Arnaud LeRouzic, Alan C. Love, Ralph Marcucio, Michael B. Morrissey, Laura Nuño de la Rosa, Øystein H. Opedal, Mihaela Pavličev, Christophe Pélabon, Jane M. Reid, Heather Richbourg, Jacqueline L. Sztepanacz, Masahito Tsuboi, Cristina Villegas, Marta Vidal-García, Kjetil L. Voje, Andreas Wagner, Günter P. Wagner, Nathan M. Young
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 homologues and vertebrate paired appendages. This article is protected by copyright. All rights reserved.
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.
The frameshift hypothesis is a widely accepted model of bird wing evolution. This hypothesis postulates a shift in positional values, or molecular-developmental identity, that caused a change in digit phenotype. The hypothesis synthesized developmental and paleontological data on wing digit homology. The “most anterior digit” (MAD) hypothesis presents an alternative view based on changes in transcriptional regulation in the limb. The molecular evidence for both hypotheses is that the MAD expresses Hoxd13 but not Hoxd11 and Hoxd12. This digit I “signature” is thought to characterize all amniotes. Here, we studied Hoxd expression patterns in a phylogenetic sample of 18 amniotes. Instead of a conserved molecular signature in digit I, we find wide variation of Hoxd11, Hoxd12, and Hoxd13 expression in digit I. Patterns of apoptosis, and Sox9 expression, a marker of the phalanx-forming region, suggest that phalanges were lost from wing digit IV because of early arrest of the phalanx-forming region followed by cell death. Finally, we show that multiple amniote lineages lost phalanges with no frameshift. Our findings suggest that the bird wing evolved by targeted loss of phalanges under selection. Consistent with our view, some recent phylogenies based on dinosaur fossils eliminate the need to postulate a frameshift in the first place. We suggest that the phenotype of the Archaeopteryx lithographica wing is also consistent with phalanx loss. More broadly, our results support a gradualist model of evolution based on tinkering with developmental gene expression.
Key words: limb development, evo-devo, hox genes, apoptosis, phalanx-forming region, frameshift theory, bird, reptile.
Advances in developmental genetics and evo-devo in the last several decades have enabled the growth of novel developmental approaches to the classic theme of homology. These approaches depart from the more standard phylogenetic view by contending that homology between morphological characters depends on developmental-genetic individuation and explanation. This paper provides a systematic re-examination of the relationship between developmental and phylogenetic homology in light of current evidence from developmental and evolutionary genetics and genomics. I present a qualitative model of the processes that cause de-coupling of morphological and molecular evolution by developmental system drift (DSD), and hypothesize that DSD is a widespread and regular stochastic process that is predictable from parameters such as population size, pleiotropy, and regulatory redundancy. These theoretical findings support an integrative approach in which models of DSD aid in determining when developmental-genetic explanations of homology apply and when other explanations such as stabilizing selection are needed. I argue that a phylogenetic monism about the definition and criteria of homology supports the integration of different explanations into a theory of homology better than pluralism does.
There have been repeated attempts in the history of comparative biology to provide a mechanistic account of morphological homology. However, it is well-established that homologues can develop from diverse sets of developmental causes, appearing not to share any core causal architecture that underwrites character identity. We address this challenge with a new conceptual model of Character Identity Mechanisms (ChIMs). ChIMs are cohesive mechanisms with a recognizable causal profile that allows them to be traced through evolution as homologues despite having a diverse etiological organization. Our model hypothesizes that anatomical units at different levels of organization -- cell types, tissues, and organs -- have level-specific ChIMs with different conserved parts, activities, and organization. Relying on a methodology of conceptual engineering, we show how the ChIM concept advances our understanding of the developmental basis of morphological characters, while forging an important link between comparative and mechanistic biology.
In crown group tetrapods, individual digits are homologized in relation to a pentadactyl ground plan. However, testing hypotheses of digit homology is challenging because it is unclear whether digits represent distinct and conserved gene regulatory states. Here we show dramatic evolutionary dynamism in the gene expression profiles of digits, challenging the notion that five digits have conserved developmental identities across amniotes. Transcriptomics shows diversity in the patterns of gene expression differentiation of digits, although the anterior-most digit of the pentadactyl limb has a unique, conserved expression profile. Further, we identify a core set of transcription factors that are differentially expressed among the digits of amniote limbs; their spatial expression domains, however, vary between species. In light of these results, we reevaluate the frame shift hypothesis of avian wing evolution and conclude only the identity of the anterior-most digit has shifted position, suggesting a 1,3,4 digit identity in the bird wing.
The last half century of paleornithological research has transformed the way that biologists perceive the evolutionary history of birds. This transformation has been driven, since 1969, by a series of exciting fossil discoveries combined with intense scientific debate over how best to interpret these discoveries. Ideally, as evidence accrues and results accumulate, interpretive scientific agreement forms. But this has not entirely happened in the debate over avian origins: the accumulation of scientific evidence and analyses has had some effect, but not a conclusive one, in terms of resolving the question of avian origins. Although the majority of biologists have come to accept that birds are dinosaurs, there is lingering and, in some quarters, strident opposition to this view. In order to both understand the ongoing disagreement about avian origins and generate a prediction about the future of the debate, here we use a revised model of scientific practice to assess the current and historical state of play surrounding the topic of bird evolutionary origins. Many scientists are familiar with the metascientific scholars Sir Karl Popper and Thomas Kuhn, and these are the primary figures that have been appealed to so far, in prior attempts to assess the dispute. But we demonstrate that a variation of Imre Lakatos’s model of progressive versus degenerative research programmes provides a novel and productive assessment of the debate. We establish that a refurbished Lakatosian account both explains the intractability of the dispute and predicts a likely outcome for the debate about avian origins. In short, here, we offer a metascientific tool for rationally assessing competing theories—one that allows researchers involved in seemingly intractable scientific disputes to advance their debates.
Understanding the evolution of biodiversity on Earth is a central aim in biology. Currently, various disciplines of science contribute to unravel evolution at all levels of life, from individual organisms to species and higher ranks, using different approaches and specific terminologies. The search for common origin, traditionally called homology, is a connecting paradigm of all studies related to evolution. However, it is not always sufficiently taken into account that defining homology depends on the hierarchical level studied (organism, population, species), which can cause confusion. Therefore, we propose a framework to define homologies making use of existing terms, which refer to homology in different fields, but restricting them to an unambiguous meaning and a particular hierarchical level. We propose to use the overarching term 'homology' only when 'morphological homology', 'vertical gene transfer' and 'phylogenetic homology' are confirmed. Consequently, neither phylogenetic nor morphological homology are equal to homology. This paper is intended for readers with different research backgrounds. We challenge their traditional approaches, inviting them to consider the proposed framework and offering them a new perspective for their own research.
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 (a) an abstract genealogical account that unifies the application of the term to all types of characters used in phylogenetic systematics and (b) 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 (e.g. genes and proteins, or body parts), 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 (e.g. serial homology, paralogy, and xenology) 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.
The origin of the avian hand, with its reduced and fused carpals and digits, from the five-fingered hands and complex wrists of early dinosaurs represents one of the major transformations of manus morphology among tetrapods. Much attention has been directed to the later part of this transition, from four- to three-fingered taxa. However, earlier anatomical changes may have influenced these later modifications, possibly paving the way for a later frameshift in digit identities. We investigate the five- to four-fingered transition among early dinosaurs, along with changes in carpus morphology. New three-dimensional reconstructions from computed tomography data of the manus of the Triassic and Early Jurassic theropod dinosaurs Coelophysis bauri and Megapnosaurus rhodesiensis are described and compared intra- and interspecifically. Several novel findings emerge from these reconstructions and comparisons, including the first evidence of an ossified centrale and a free intermedium in some C. bauri specimens, as well as confirmation of the presence of a vestigial fifth metacarpal in this taxon. Additionally, a specimen of C. bauri and an unnamed coelophysoid from the Upper Triassic Hayden Quarry, New Mexico, are to our knowledge the only theropods (other than alvarezsaurs and birds) in which all of the distal carpals are completely fused together into a single unit. Several differences between the manus of C. bauri and M. rhodesiensis are also identified. We review the evolution of the archosauromorph manus more broadly in light of these new data, and caution against incorporating carpal characters in phylogenetic analyses of fine-scale relationships of Archosauromorpha, in light of the high degree of observed polymorphism in taxa for which large sample sizes are available, such as the theropod Coelophysis and the sauropodomorph Plateosaurus. We also find that the reduction of the carpus and ultimate loss of the fourth and fifth digits among early dinosaurs did not proceed in a neat, stepwise fashion, but was characterized by multiple losses and possible gains of carpals, metacarpals and phalanges. Taken together, the high degree of intra- and interspecific variability in the number and identities of carpals, and the state of reduction of the fourth and fifth digits suggest the presence of a ‘zone of developmental variability’ in early dinosaur manus evolution, from which novel avian-like morphologies eventually emerged and became channelized among later theropod clades.
Since the late 1970s, the field of evolutionary biology has undergone empirical and theoretical developments that have threaten the pillars of evolutionary theory. Some evolutionary biologists have recently argued that evolutionary biology is not experiencing a paradigm shift, but an expansion of the modern synthesis. Philosophers of biology focusing on scientific practices seem to agree with this pluralistic interpretation and have argued that evolutionary theory should rather be seen as an organized network of multiple problem agendas with diverse disciplinary contributors. In this paper, I apply a computational analysis to study the dynamics and conceptual structure of one of the main emerging problem agendas in evolutionary biology: evolvability. I have used CiteSpace, an application for visualizing and analyzing trends and patterns in scientific literature that applies cocitation analysis to identify scientific specialities. I analyze the main clusters of the evolvability cocitation network with the aim to identify the main research lines and the interdisciplinary relationships that structure this research front. I then compare these results with the existing classifications of evolvability concepts, and identify four main conceptual tensions within the definitions of evolvability. Finally, I argue that there is a lot of usefulness in the inconsistency in which the term evolvability is used in biological research. I claim that evolvability research has set up "trading zones" in biology that make possible interdisciplinary exchanges.
It is time to escape the constraints of the Systematics Wars narrative and pursue new questions that are better positioned to establish the relevance of the field in this time period to broader issues in the history of biology and history of science. To date, the underlying assumptions of the Systematics Wars narrative have led historians to prioritize theory over practice and the conflicts of a few leading theorists over the less-polarized interactions of systematists at large. We show how shifting to a practice-oriented view of methodology, centered on the trajectory of mathematization in systematics, demonstrates problems with the common view that one camp (cladistics) straightforwardly “won” over the other (phenetics). In particular, we critique David Hull’s historical account in Science as a Process by demonstrating exactly the sort of intermediate level of positive sharing between phenetic and cladistic theories that undermines their mutually exclusive individuality as conceptual systems over time. It is misleading, or at least inadequate, to treat them simply as holistically opposed theories that can only interact by competition to the death. Looking to the future, we suggest that the concept of workflow provides an important new perspective on the history of mathematization and computerization in biology after World War II.
Adherents of the current orthodoxy of a derivation of birds from theropod dinosaurs, criticize the commentary by Feduccia (2013, Auk, 130) [1 - 12] entitled "Bird Origins Anew" as well as numerous papers by Lingham-Soliar on theropod dermal fibers, using numerous mischaracterizations and misstatements of content, and illustrate their own misconceptions of the nature of the debate, which are here clarified. While there is general agreement with the affinity of birds and maniraptorans, the widely accepted phylogeny, advocating derived earth-bound maniraptorans giving rise to more primitive avians (i.e. Archaeopteryx), may be "topsyturvy." The current primary debate concerns whether maniraptorans are ancestral or derived within the phylogeny, and whether many maniraptorans and birds form a clade distinct from true theropods. Corollaries of the current scheme show largely terrestrial maniraptoran theropods similar to the Late Cretaceous Velociraptor giving rise to avians, and flight originating via a terrestrial (cursorial) "gravity-resisted," as opposed to an arboreal "gravity-assisted" model. The current dogma posits pennaceous flight remiges in earth-bound theropods having evolved in terrestrial theropods that never flew. As part of the orthodoxy, fully feathered maniraptorans such as the tetrapteryx gliders Microraptor and allies, are incorrectly reconstructed as terrestrial cursors, when in reality their anatomy and elongate hindlimb feathers would be a hindrance to terrestrial locomotion.The same is true of many early birds, exemplified by reconstruction of the arboreally adapted Confuciusornis as a terrestrial predator, part of the overall theropodan scheme of birds evolving from terrestrial dinosaurs, and flight from the ground up. Both sides of this contentious debate must be constantly aware that new fossil or even molecular discoveries on birds may change current conclusions.
Recent discoveries of spectacular dinosaur fossils overwhelmingly support the hypothesis that birds are descended from maniraptoran theropod dinosaurs, and furthermore, demonstrate that distinctive bird characteristics such as feathers, flight, endothermic physiology, unique strategies for reproduction and growth, and a novel pulmonary system originated among Mesozoic terrestrial dinosaurs. The transition from ground-living to flight-capable theropod dinosaurs now probably represents one of the best-documented major evolutionary transitions in life history. Recent studies in developmental biology and other disciplines provide additional insights into how bird characteristics originated and evolved. The iconic features of extant birds for the most part evolved in a gradual and stepwise fashion throughout archosaur evolution. However, new data also highlight occasional bursts of morphological novelty at certain stages particularly close to the origin of birds and an unavoidable complex, mosaic evolutionary distribution of major bird characteristics on the theropod tree. Research into bird origins provides a premier example of how paleontological and neontological data can interact to reveal the complexity of major innovations, to answer key evolutionary questions, and to lead to new research directions. A better understanding of bird origins requires multifaceted and integrative approaches, yet fossils necessarily provide the final test of any evolutionary model.
Copyright © 2014, American Association for the Advancement of Science.
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.
A philosophical account of human nature that defends the concept against dehumanization, Darwinian, and developmentalist challenges.
Human nature has always been a foundational issue for philosophy. What does it mean to have a human nature? Is the concept the relic of a bygone age? What is the use of such a concept? What are the epistemic and ontological commitments people make when they use the concept? In What's Left of Human Nature? Maria Kronfeldner offers a philosophical account of human nature that defends the concept against contemporary criticism. In particular, she takes on challenges related to social misuse of the concept that dehumanizes those regarded as lacking human nature (the dehumanization challenge); the conflict between Darwinian thinking and essentialist concepts of human nature (the Darwinian challenge); and the consensus that evolution, heredity, and ontogenetic development result from nurture and nature.
After answering each of these challenges, Kronfeldner presents a revisionist account of human nature that minimizes dehumanization and does not fall back on outdated biological ideas. Her account is post-essentialist because it eliminates the concept of an essence of being human; pluralist in that it argues that there are different things in the world that correspond to three different post-essentialist concepts of human nature; and interactive because it understands nature and nurture as interacting at the developmental, epigenetic, and evolutionary levels. On the basis of this, she introduces a dialectical concept of an ever-changing and “looping” human nature. Finally, noting the essentially contested character of the concept and the ambiguity and redundancy of the terminology, she wonders if we should simply eliminate the term “human nature” altogether.
As it spread through time and into distinct areas of science-from comparative anatomy to evolutionary biology, cladistics, developmental and molecular biology-the homology concept has changed considerably, presenting various meanings. Despite many attempts at developing a comprehensive understanding of the concept, this context-sensitive notion of homology has been a subject of an ongoing debate. Inspired by that and following Kevin de Queiroz and Richard Mayden's view on species concept and delimitation, we presented in this article an attempt to systematize and advance the understanding of the homology problem. Our main goals were: (i) to present a comprehensive checklist of 'concepts of homology'; (ii) to identify which are really concepts with ontological definitions (theoretically rooted in structural correspondence and common ancestry), and which are, in fact, not concepts, but epistemological (empirical and methodological) criteria of homology delimitation; (iii) to provide a synonymy of the concepts and criteria of homology delimitation; (iv) to present a hierarchy of homology concepts within Hennig's hologenetic system; and (v) to endorse the adoption of a unified view of homology by treating homology as a correspondence of spatio-temporal properties (genetic, epigenetic, developmental and positional) at the level of the individual, species or monophyletic group. We found 59 'concepts of homology' in the literature, from which 34 were categorically treated as concepts, 17 as criteria of homology delimitation, Four were excluded from our treatment, and Müller's five concepts were rather treated as approaches to homology. Homology concepts and criteria were synonymized based on structural correspondence, replicability, common ancestry, genetic and epigenetic developmental causes, position and optimization. Regarding the synonymy, we conclusively recognized 21 different concepts of homology, and five empirical and four methodological criteria. Hierarchical ontological aspects of homology were systematized under Hennig's hologenetic system, based on the existence of ontogenetic, tokogenetic and phylogenetic levels of homology. The delimitation of tokogenetic and phylogenetic homologies depends on optimization criteria. The unified view of homology is discussed in the context of the ancestral angiosperm flower.
There is widespread agreement that "homology" designates something of fundamental biological importance, but no consensus as to how precisely that "something" should be defined, recognized, or theorized. Philosophical observers of this situation commonly focus on tensions between historical and mechanistic explications of homological sameness by appeal, respectively, to common ancestry and shared developmental resources. This paper uses select historical episodes to decenter those tensions and challenge standard narratives about how they arose. Haas and Simpson (1946) influentially defined "homology" simply as "similarity due to common ancestry." They claimed historical precedent in Lankester (1870) but seriously oversimplified his views in the process. Lankester did prioritize common ancestry, but he also raised mechanistic questions that resonate with contemporary evo devo work on homology. The rise of genetics inspired similar speculations in twentieth-century workers like Boyden (1943), a zoologist who engaged Simpson in a 15-year debate over homology. Though he shared Simpson's devotion to taxonomy and his interest in evolutionary history, he favored a more operational and less theoretical homology concept. Their dispute is poorly captured by current analyses of the homology problem. It calls for further study of the complex relationship between concepts and the epistemic goals they serve.
Unease with the inclusion of 'sameness' in Owen's definition of homology characterizes a substantial part of the literature on this subject, where this term has acquired an increasingly strict metaphysical flavor. Taken for granted the existence of body features that are "the same", their existence has been explained by appealing to universal laws of form, as the product of common ancestry, or in terms of proximal causes responsible for the emergence of conserved developmental modules. However, a fundamentally different approach is possible, if we shift attention from metaphysics to epistemology. We may reword Owen's statement as follows: organs of different animals, in so far as they can be described as the same despite any difference in form and function, are called homologues. The proposed framework provides an umbrella for both the traditional, all-or-nothing concept of homology, and the less fashionable alternatives of factorial or partial homology, as well as for an extension of homology from form to function. No less attractive is the prospect to handle also ghost homologues, the body parts or organs of which there is non-objective evidence in a given clade, but can nevertheless be represented, in a description that encapsulates some of the traits observable in their extant homologue in the sister clade. Stripped of its different and constraining metaphysical explanations, homology survives as an anchor concept to which different nomadic disciplines and research agendas can be associated. This article is protected by copyright. All rights reserved.
The continued use of the idea of homology is questionable on philosophical and scientific grounds. It is based on the widespread idea that a "homologue" in extant taxa can be "traced back" to a feature in common ancestor. In contrast, Richard Owen, who first used the term in 1846, saw homology (homologue) differently, as "sameness": "the same organ in different animals under every variety of form and function." At that point in time, he was not influenced by evolutionary thinking, and more focused on the details and approaches to biological comparison and description. His was a perceptive approach to comparison. This paper argues that the concept of homology no longer plays a useful role in comparative biology. It is a conceptual idea with little or no empirical implications for modern comparisons among phenotypes. Comparative biology now uses formal phylogenetic analysis in which similar features in individuals of two or more taxa are treated as characters on a tree and tested for historical "sameness" in terms of the concept of synapomorphy. If we are to understand the complexities of phenotypic evolution, applying this method to detailed comparative data will be essential. At the same time, a deep understanding of the phenotype and its history will emerge only through the use of multidisciplinary approaches that address historical changes at different hierarchical levels. This article is protected by copyright. All rights reserved.
Biological science uses multiple species concepts. Order can be brought to this diversity if we recognize two key features. First, any given species concept is likely to have a patchwork structure, generated by repeated application of the concept to new domains. We illustrate this by showing how two species concepts (biological and ecological) have been modified from their initial eukaryotic applications to apply to prokaryotes. Second, both within and between patches, distinct species concepts may interact and hybridize. We thus defend a semantic picture of the species concept as a collection of interacting patchwork structures. Thus, although not all uses of the term pick out the same kind of unit in nature, the diversity of uses reflects something more than mere polysemy. We suggest that the emphasis on the use of species to pick out natural units is itself problematic, because that is not the term’s sole function. In particular, species concepts are used to manage inquiry into processes of speciation, even when these processes do not produce clearly delimited species.
Willi Hennig (1913–76), founder of phylogenetic systematics, revolutionised our understanding of the relationships among species and their natural classification. An expert on Diptera and fossil insects, Hennig's ideas were applicable to all organisms. He wrote about the science of taxonomy or systematics, refining and promoting discussion of the precise meaning of the term 'relationship', the nature of systematic evidence, and how those matters impinge on a precise understanding of monophyly, paraphyly, and polyphyly. Hennig's contributions are relevant today and are a platform for the future. This book focuses on the intellectual aspects of Hennig's work and gives dimension to the future of the subject in relation to Hennig's foundational contributions to the field of phylogenetic systematics. Suitable for graduate students and academic researchers, this book will also appeal to philosophers and historians interested in the legacy of Willi Hennig.
I propose a new account of homology, according to which homology is a correspondence of developmental mechanisms due to common ancestry, formally defined as an isomorphism of causal graphs over lineages. The semiformal definition highlights the role of homology as a higher-order principle unifying evolutionary models and also provides definite meanings to concepts like constraints, evolvability, and novelty. The novel interpretation of homology suggests a broad perspective that accommodates evolutionary developmental biology and traditional population genetics as distinct but complementary approaches to understand evolution, prompting further empirical and theoretical research. © 2017 by the Philosophy of Science Association. All rights reserved.
Here we describe the forearm and manus of the ceratosaurian theropod dinosaur Ceratosaurus nasicornis Marsh, 1884, from the Upper Jurassic Morrison Formation of the western U.S.A. Recently removed from exhibition and reprepared, the holotype offers important new information on the morphology of this taxon that bears on the evolution of the forelimb in nonavian theropod dinosaurs more generally. The ulna and radius show particular similarities to those of Dilophosaurus and Eoabelisaurus but lack features that characterize derived abelisaurids. In the manus, Ceratosaurus bears short first phalanges, like more derived taxa in the clade, but retains metacarpals that are much more similar to those of Dilophosaurus, Berberosaurus, and Eoabelisaurus. Taken together, and incorporated with existing phylogenetic data on other ceratosaurs, these data are consistent with the placement of Ceratosaurus as close to Abelisauroidea but basal to Eoabelisaurus. More importantly, they strongly imply that the extremely reduced manus of Limusaurus is a derived condition that does not reflect the primitive state for Ceratosauria and therefore that Averostra is not the most likely placement for a shift in manus digit identity during theropod evolution. Finally, digit reduction began in ceratosaurs that still possessed most phalanges and unguals, and we infer that grasping would have been retained as a primary, if reduced, function in these forms.Citation for this article: Carrano, M. T., and J. Choiniere. 2016. New information on the forearm and manus of Ceratosaurus nasicornis Marsh, 1884 (Dinosauria, Theropoda), with implications for theropod forelimb evolution. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2015.1054497.
In the past century, nearly all of the biological sciences have been directly affected by discoveries and developments in genetics, a fast-evolving subject with important theoretical dimensions. in this rich and accessible book, Paul Griffiths and Karola Stotz show how the concept of the gene has evolved and diversified across the many fields that make up modern biology. By examining the molecular biology of the 'environment', they situate genetics in the developmental biology of whole organisms, and reveal how the molecular biosciences have undermined the nature/nurture distinction. Their discussion gives full weight to the revolutionary impacts of molecular biology, while rejecting 'genocentrism' and 'reductionism', and brings the topic right up to date with the philosophical implications of the most recent developments in genetics. Their book will be invaluable for those studying the philosophy of biology, genetics and other life sciences.
Homology-a similar trait shared by different species and derived from common ancestry, such as a seal's fin and a bird's wing-is one of the most fundamental yet challenging concepts in evolutionary biology. This groundbreaking book provides the first mechanistically based theory of what homology is and how it arises in evolution. Günter Wagner, one of the preeminent researchers in the field, argues that homology, or character identity, can be explained through the historical continuity of character identity networks-that is, the gene regulatory networks that enable differential gene expression. He shows how character identity is independent of the form and function of the character itself because the same network can activate different effector genes and thus control the development of different shapes, sizes, and qualities of the character. Demonstrating how this theoretical model can provide a foundation for understanding the evolutionary origin of novel characters, Wagner applies it to the origin and evolution of specific systems, such as cell types; skin, hair, and feathers; limbs and digits; and flowers. The first major synthesis of homology to be published in decades, Homology, Genes, and Evolutionary Innovation reveals how a mechanistically based theory can serve as a unifying concept for any branch of science concerned with the structure and development of organisms, and how it can help explain major transitions in evolution and broad patterns of biological diversity.
What counts as an individual in the living world? What does it mean for a living thing to remain the same through time while constantly changing? Immunology, one of the most dynamic fields of today's biology, considers these questions its province, and answers them through its crucial concepts of "self" and "nonself." Though immunology has been dominated since the 1940s by the self-nonself theory, this book argues that this theory is inadequate, because immune responses to self constituents and immune tolerance of foreign entities are the rule, not the exception. An alternative theory, the continuity theory, is advanced instead. This theory offers a new way to answer the question of what triggers an immune response. It also echoes the recent realization that all organisms, and not higher vertebrates only, have an immune system. This book's main thesis is that the self-nonself theory should be abandoned, but that immunology still proves to be decisive for delineating the boundaries of the organism. Articulating an evolutionary and an immunological perspective, it offers an original conception of the organism. Tolerance of the fetus by the mother and of countless bacteria on the body's surfaces proves that every organism is heterogeneous, that is, made of entities of different origins. In other words, every organism appears as a chimera, a mixed living thing the cohesiveness of which is ensured by the constant action of its immune system. The Limits of the Self will be essential reading for anyone interested in the definition of biological individuality and the understanding of the immune system.
I consider the relationship between scientific practice and the philosophical debate surrounding biological individuality. I argue for the sensitivity account, on which biologists do not require a resolution to the individuality debate. This view puts me in disagreement with much of the literature on biological individuality, where it has become common to claim that there is a relationship of dependence between biologists’ conceptions of individuality and the quality of their empirical work. © 2015 by the Philosophy of Science Association. All rights reserved.
Homology (Greek homologica, agreement) is a biological concept with a long and checkered history, summarized in a recent volume of papers (Hall, 1994) devoted to the concept, criteria, and mechanisms of homology and its pivotal importance as the hierarchical basis of comparative biology; what Julian Huxley (1928) referred to as “morphology’s central conception.” Thinking about relationships between homology and embryonic development while writing a larger work on Evolutionary Developmental Biology (Hall, 1992) prompted production of the 1993 volume celebrating the sesquicentennial of Richard Owen having delineated homology from analogy, and the present review on how homology relates to, or is perceived to relate to, embryonic development.
The three digits of birds have been homologized with either the first-second-third or second-third-fourth digits of the primitive tetrapod limb. The disagreement as to the identification of the carpals, metacarpals, and digits of the bird manus is more than a semantic question of numbering digits. First, it is a question of whether birds have followed the digital reduction pattern of other amniotes and whether there is a universal developmental process of carpal, metacarpal, and digital differentiation in amniotes. Second, does the identification of the bird metacarpals and digits also provide evidence for the digits of Archaeopteryx?
In recent years, the concept of evolvability has been gaining in prominence both within evolutionary developmental biology (evo-devo) and the broader field of evolutionary biology. Despite this, there remains considerable disagreement about what evolvability is. This article offers a solution to this problem. I argue that, in focusing too closely on the role played by evolvability as an explanandum in evo-devo, existing philosophical attempts to clarify the evolvability concept have been overly narrow. Within evolutionary biology more broadly, evolvability offers a robust explanation for the evolutionary trajectories of populations. Evolvability is an abstract, robust, dispositional property of populations, which captures the joint causal influence of their internal features on the outcomes of evolution (as opposed to the causal influence of selection, which is often characterized as external). When considering the nature of the physical basis of this disposition, it becomes clear that the many existing definitions of evolvability at play within evo-devo should be understood as capturing only aspects of a much broader phenomenon.
• 1 Introduction
• 2 The Problem of Evolvability
• 3 The Theoretical Role of Evolvability in Evolutionary Biology
• 3.1 The explanatory targets of evolutionary biology
• 3.2 Selection-based explanations
• 3.3 Lineage explanations
• 3.4 Evolvability-based explanations
• 3.5 What properties must evolvability have?
• 4 What Evolvability Really Is
• 4.1 Making sense of f t
• 4.2 Making sense of x and b
• 5 What of the Limbs? The Power of E
• 6 Conclusion
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.
A recent review of the homology concept in cladistics is critiqued in light of the historical literature. Homology as a notion relevant to the recognition of clades remains equivalent to synapomorphy. Some symplesiomorphies are “homologies” inasmuch as they represent synapomorphies of more inclusive taxa; others are complementary character states that do not imply any shared evolutionary history among the taxa that exhibit the state. Undirected character-state change (as characters optimized on an unrooted tree) is a necessary but not sufficient test of homology, because the addition of a root may alter parsimonious reconstructions. Primary and secondary homology are defended as realistic representations of discovery procedures in comparative biology, recognizable even in Direct Optimization. The epistemological relationship between homology as evidence and common ancestry as explanation is again emphasized. An alternative definition of homology is proposed.
© The Willi Hennig Society 2012.
The identification of avian and dinosaurian digits remains one of the major controversies in vertebrate evolution. A long history of morphological interpretations of fossil forms and studies of limb development in embryos has been given as evidence for two differing points of view. From an originally pentadactyl forelimb, either digits I, II and III form in the manus of birds and thus support a dinosaurian ancestry, or digits II, III and IV form in the manus supporting a more ancient ancestry or an evolutionary frame shift. A review of the history of research into the subject is presented here, dating from approximately 1825 to 2009.
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.
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.
