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.

Biology and Philosophy, forthcoming 2007.

“Functional homology” appears regularly in different areas of biological research and yet it is apparently a contradiction in terms—homology concerns identity of structure regardless of form and function. I argue that despite this conceptual tension there is a legitimate conception of ‘homology of function’, which can be recovered by utilizing a distinction from pre-Darwinian physiology (use versus activity) to identify an appropriate meaning of ‘function’. This account is directly applicable to molecular developmental biology and shares a connection to the theme (...) of hierarchy in homology. I situate ‘homology of function’ within existing definitions and criteria for structural assessments of homology, and introduce a criterion of ‘organization’ for judging function homologues, which focuses on hierarchically interconnected interdependencies (similar to relative position and connection for skeletal elements in structural homology). This analysis of biological concepts has at least three broad philosophical consequences: (1) it provides the grounds for the study of behavior and psychological categories as homologues; (2) it demonstrates that philosophers who take selected effect function as primary effectively ignore large portions of comparative, structural, and experimental research, thereby misconstruing biological reasoning and knowledge; and, (3) it underwrites causal generalizations, which illuminates inferences made from model organisms in experimental biology. (shrink)

Philosophical discussions of biological classification have failed to recognise the central role of homology in the classification of biological parts and processes. One reason for this is a misunderstanding of the relationship between judgments of homology and the core explanatory theories of biology. The textbook characterisation of homology as identity by descent is commonly regarded as a definition. I suggest instead that it is one of several attempts to explain the phenomena of homology. Twenty years ago the ‘new experimentalist’ movement (...) in philosophy of science drew attention to the fact that many experimental phenomena have a ‘life of their own’: the conviction that they are real is not dependent on the theories used to characterise and explain them. I suggest that something similar can be true of descriptive phenomena, and that many homologies are phenomena of this kind. As a result the descriptive biology of form and function has a life of its own—a degree of epistemological independence from the theories that explain form and function. I also suggest that the two major ‘homology concepts’ in contemporary biology, usually seen as two competing definitions, are in reality complementary elements of the biological explanation of homology. (shrink)

Editors' introduction to the special issue on homology (Biology and Philosophy Vol. 22, Issue 5, 2007).

Abstract: The massive redeployment hypothesis (MRH) is a theory about the functional topography of the human brain, offering a middle course between strict localization on the one hand, and holism on the other. Central to MRH is the claim that cognitive evolution proceeded in a way analogous to component reuse in software engineering, whereby existing components-originally developed to serve some specific purpose-were used for new purposes and combined to support new capacities, without disrupting their participation in existing programs. If the (...) evolution of cognition was indeed driven by such exaptation, then we should be able to make some specific empirical predictions regarding the resulting functional topography of the brain. This essay discusses three such predictions, and some of the evidence supporting them. Then, using this account as a background, the essay considers the implications of these findings for an account of the functional integration of cognitive operations. For instance, MRH suggests that in order to determine the functional role of a given brain area it is necessary to consider its participation across multiple task categories, and not just focus on one, as has been the typical practice in cognitive neuroscience. This change of methodology will motivate (even perhaps necessitate) the development of a new, domain-neutral vocabulary for characterizing the contribution of individual brain areas to larger functional complexes, and direct particular attention to the question of how these various area roles are integrated and coordinated to result in the observed cognitive effect. Finally, the details of the mix of cognitive functions a given area supports should tell us something interesting not just about the likely computational role of that area, but about the nature of and relations between the cognitive functions themselves. For instance, growing evidence of the role of “motor” areas like M1, SMA and PMC in language processing, and of “language” areas like Broca’s area in motor control, offers the possibility for significantly reconceptualizing the nature both of language and of motor control. (shrink)

The specialization of visual function within biological function is reason for introducing “homology thinking” into explanations of the visual system. It is argued that such specialization arises when organisms evolve by differentiation from their predecessors. Thus, it is essentially historical, and visual function should be regarded as a lineage property. The colour vision of birds and mammals do not function the same way as one another, on this account, because each is an adaptation to special needs of the visual functions (...) of predecessors—very different kinds of predecessors in each case. Thus, history underlies function. We also see how homology thinking figures in the hierarchical classification of visual systems, and how it supports the explanation of visual function by functional role analysis. (shrink)

What are 'emotions'? This book offers a balanced survey of facts and theory.

This article argues that at least very many judgments of homology rest on prior attributions of selected‐effect (SE) function, and that many of the “parts” of biological systems that are rightly classified as homologous are constituted by (are so classified in virtue of) their consequence etiologies. We claim that SE functions are often used in the prior identification of the parts deemed to be homologous and are often used to differentiate more restricted homologous kinds within less restricted ones. In doing (...) so, we discuss recent criticism of this view that has been offered (especially that offered by Paul Griffiths). *Received July 2007; revised November 2008. †To contact the authors, please write to: Department of Philosophy, 201 West Duke Building, Box 90743, Durham, NC 27708; e‐mail: [email protected] or [email protected]. (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)

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)

Paul E. Griffiths argues that most research on the emotions has been as misguided as Aristotelian efforts to study "superlunary objects" - objects...

Although there is no in-principle impediment to an EvoDevo of behavior, such an endeavor is not as straightforward as one might think; many of the key terms and concepts used in EvoDevo are tailored to suit its traditional focus on morphology, and are consequently difficult to apply to behavior. In this light, the application of the EvoDevo conceptual toolkit to the behavioral domain requires the establishment of a set of tractable concepts that are readily applicable to behavioral characters. Here, I (...) begin the type of theoretical work that needs to be undertaken in order to achieve this, focusing in particular on the key concept of “novelty.” Building on existing criteria used for the identification of behavioral homology from behavioral ecology, I develop a set of operational criteria for identifying novelty in the behavioral domain. These criteria provide a conceptual foundation for the study of novelty in behavioral traits. (shrink)

I defend the view that many biological categories are defined by homology against a series of arguments designed to show that all biological categories are defined, at least in part, by selected function. I show that categories of homology are `abnormality inclusive'—something often alleged to be unique to selected function categories. I show that classifications by selected function are logically dependent on classifications by homology, but not vice-versa. Finally, I reject the view that biologists must use considerations of selected function (...) to abstract away from variation and pathology to form a canonical description of a class of biological systems. (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)

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)

Emotions are discrete, automatic responses to universally shared, culture-specific and individual-specific events. The emotion terms, such as anger, fear, etcetera, denote a family of related states sharing at least 12 characteristics, which distinguish one emotion family from another, as well as from other affective states. These affective responses are preprogrammed and involuntary, but are also shaped by life experiences.

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.

An emerging class of theories concerning the functional structure of the brain takes the reuse of neural circuitry for various cognitive purposes to be a central organizational principle. According to these theories, it is quite common for neural circuits established for one purpose to be exapted (exploited, recycled, redeployed) during evolution or normal development, and be put to different uses, often without losing their original functions. Neural reuse theories thus differ from the usual understanding of the role of neural plasticity (...) (which is, after all, a kind of reuse) in brain organization along the following lines: According to neural reuse, circuits can continue to acquire new uses after an initial or original function is established; the acquisition of new uses need not involve unusual circumstances such as injury or loss of established function; and the acquisition of a new use need not involve (much) local change to circuit structure (e.g., it might involve only the establishment of functional connections to new neural partners). Thus, neural reuse theories offer a distinct perspective on several topics of general interest, such as: the evolution and development of the brain, including (for instance) the evolutionary-developmental pathway supporting primate tool use and human language; the degree of modularity in brain organization; the degree of localization of cognitive function; and the cortical parcellation problem and the prospects (and proper methods to employ) for function to structure mapping. The idea also has some practical implications in the areas of rehabilitative medicine and machine interface design. (shrink)

I adopt a cladistic view of species, and explore the possibility that there exists an equally valuable cladistic view of organismic traits. This suggestion seems to run counter to the stress on functional views of biological traits in recent work in philosophy and psychology. I show how the tension between these two views can be defused with a multilevel view of biological explanation. Despite the attractions of this compromise, I conclude that we must reject it, and adopt an essentially cladistic (...) conception of biological traits. (shrink)