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The subcategory of Developmental Modularity consists of papers that engage with the study of development as consisting of discrete units (modules) and the extent to which these modules are integrated (see Developmental Systems Theory for a contrasting position). The specific developmental phenomena that are studied include heterochrony and allometry, duplication and divergence of modules (as seen in segmentation), and the co-option of modules (often called exaptation).

The unifying philosophical theme is ontological; the landscape consists of differing views on how a developing organism is divided into functional or structural subunits. The internal disagreements are over what the basic units are, and how they are identified and organized.  However, all papers in this section examine the extent to which an organism’s development can be explained in terms of interactions between (reasonably) autonomous morphogenetic (developmental) modules.

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  1. Wilfried Allaerts (1999). Local and Global Patterns During Morphogenesis of the Retinotectal Topographical Mapping in the Vertebrate Brain. Acta Biotheoretica 47 (2):99-122.
    The highly ordered neuronal projections from the retina to the tectum mesencephali (optic tectum) in several vertebrate groups have been intensively studied. Several hypotheses so far have been proposed, suggesting mechanisms to explain the topographical and biochemical specificity of the retinotectal projections during ontogeny. In the present paper we compare the main hypotheses of retinotectal development with respect to the nature of specificity envisaged, the activity-dependence versus inheritance criterium and the strategy of argument, in casu the descriptive versus interferential type (...)
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  2. Ron Amundson, Accounting For Vertebrate Limbs: From Owen's Homology To Novelty In Evo-Devo.
    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 (...)
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  3. Sergio Balari & Guillermo Lorenzo (2015). The End of Development. Biological Theory 10 (1):60-72.
    Recently, there has been a growing interest, both within theoretical biology and the philosophy of biology, in the possibility and desirability of a theory of development. Among the many issues raised within this debate, the questions of the spatial and temporal boundaries of development have received particular attention. In this article, noting that so far the discussion has mostly centered on the processes of morphogenesis and organogenesis, we argue that an important missing element in the equation, namely the development of (...)
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  4. Paul B. Baltes (1998). Testing the Limits of the Ontogenetic Sources of Talent and Excellence. Behavioral and Brain Sciences 21 (3):407-408.
    Experiential factors such as long-term deliberate practice are powerful and necessary conditions for outstanding achievement. Nevertheless, to be able to reject the role of biology based individual differences (including genetic ones) in the manifestation of talent requires designs that expose heterogeneous samples to so-called testing-the-limits conditions, allowing asymptotic levels of performance to be analyzed comparatively. When such research has been conducted, as in the field of lifespan cognition, individual differences, including biology based ones, come to the fore and demonstrate that (...)
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  5. C. D. N. Barel (1993). Concepts of an Architectonic Approach to Transformation Morphology. Acta Biotheoretica 41 (4):345-381.
    This paper is about a general methodology for pattern transformation. Patterns are network representations of the relations among structures and functions within an organism. Transformation refers to any realistic or abstract transformation relevant to biology, e.g. ontogeny, evolution and phenotypic clines. The main aim of the paper is a methodology for analyzing the range of effects on a pattern due to perturbing one or more of its structures and/or functions (transformation morphology). Concepts relevant to such an analysis of pattern transformation (...)
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  6. Ingo Brigandt (2009). Accounting for Vertebrate Limbs: From Owen's Homology to Novelty in Evo-Devo. Philosophy & Theory in Biology 1 (20130604):e004.
    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 (...)
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  7. Laurel Cooper, Ramona Walls, Justin Elser, Maria A. Gandolfo, Dennis W. Stevenson & Barry Smith (2013). The Plant Ontology as a Tool for Comparative Plant Anatomy and Genomic Analyses. Plant and Cell Physiology 54:1-23..
    The Plant Ontology (PO; http://www.plantontology.org/) is a publicly-available, collaborative effort to develop and maintain a controlled, structured vocabulary (“ontology”) of terms to describe plant anatomy, morphology and the stages of plant development. The goals of the PO are to link (annotate) gene expression and phenotype data to plant structures and stages of plant development, using the data model adopted by the Gene Ontology. From its original design covering only rice, maize and Arabidopsis, the scope of the PO has been expanded (...)
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  8. Kim J. Dale & Olivier Pourquié (2000). A Clock‐Work Somite. Bioessays 22 (1):72-83.
    Somites are transient structures which represent the most overt segmental feature of the vertebrate embryo. The strict temporal regulation of somitogenesis is of critical developmental importance since many segmental structures adopt a periodicity based on that of the somites. Until recently, the mechanisms underlying the periodicity of somitogenesis were largely unknown. Based on the oscillations of c-hairy1 and lunatic fringe RNA, we now have evidence for an intrinsic segmentation clock in presomitic cells. Translation of this temporal periodicity into a spatial (...)
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  9. Jamie A. Davies (2002). Do Different Branching Epithelia Use a Conserved Developmental Mechanism? Bioessays 24 (10):937-948.
  10. Jose F. de Celis (2003). Pattern Formation in the Drosophila Wing: The Development of the Veins. Bioessays 25 (5):443-451.
  11. Jose F. de Celis (1999). The Function of Vestigial in Drosophila Wing Development: How Are Tissue‐Specific Responses to Signalling Pathways Specified? Bioessays 21 (7):542-545.
  12. Laura Nuño de la Rosa (forthcoming). Becoming Organisms: The Organisation of Development and the Development of Organisation. History and Philosophy of the Life Sciences.
    Despite the radical importance of embryology in the development of organicism, developmental biology remains philosophically underexplored as a theoretical and empirical resource to clarify the nature of organisms. This paper discusses how embryology can help develop the organisational definition of the organism as a differentiated, functionally integrated, and autonomous system. I distinguish two conceptions of development in the organisational tradition that yield two different conceptions of the organism: the life-history view claims that organisms can be considered as such during their (...)
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  13. Arnold De Loof (1992). Problems and Paradigms. All Animals Develop From a Blastula: Consequences of an Undervalued Definition for Thinking on Development. Bioessays 14 (8):573-575.
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  14. François Delaportex (1983). Theories of Osteogenesis in the Eighteenth Century. Journal of the History of Biology 16 (3):343 - 360.
  15. Herman Denis (1994). A Parallel Between Development and Evolution: Germ Cell Recruitment by the Gonads. Bioessays 16 (12):933-938.
  16. David J. Depew (2010). Is Evolutionary Biology Infected With Invalid Teleological Reasoning? Philosophy and Theory in Biology 2 (20130604).
    John Reiss is a practicing evolutionary biologist (herpetology) who by his own account happened to be in the right place (Harvard’s Museum of Comparative Zoology) at the right time (the 1980s) to hear echoes of the debate about sociobiology that had been raging there between E. O. Wilson and, on the other side, Stephen Jay Gould and Richard Lewontin (xiv). Reiss is not concerned with sociobiology, at least in this book, but with the adaptationism that Gould and Lewontin saw in (...)
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  17. Denis Duboule (1992). The Vertebrate Limb: A Model System to Study the Hox/Hom Gene Network During Development and Evolution. Bioessays 14 (6):375-384.
  18. B. Duchaine (2011). Developmental Prosopagnosia: Cognitive, Neural, and Developmental Investigations. In Andy Calder, Gillian Rhodes, Mark Johnson & Jim Haxby (eds.), Oxford Handbook of Face Perception. OUP Oxford 821--838.
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  19. Richard P. Elinson (1989). Microtubules and Specification of the Dorsoventral Axis in Frog Embryos. Bioessays 11 (5):124-127.
  20. Raphael Falk (2004). Long Live the Genome! So Should the Gene. History and Philosophy of the Life Sciences 26 (1):105 - 121.
    Developments in the sequencing of whole genomes and in simultaneously surveying many thousands of transcription and translation products of specific cells have ushered in a conceptual revolution in genetics that rationally introduces top-down, holistic analyses. This emphasized the futility of attempts to reduce genes to structurally discrete entities along the genome, and the need to return to Johannsen's definition of a gene as 'something' that refers to an invariant entity of inheritance and development. We may view genes either as generic (...)
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  21. Bernardino Fantini (2000). L'embryologie, la 'géographie chimique' de la cellule et la synthèse entre morphologie et chimie (1930-1950). History and Philosophy of the Life Sciences 22 (3):353 - 380.
    Chemical embryology was born in 1931 with the publication of Chemical Embryology by Joseph Needham. In the following two decades it became an innovative research project aiming at the description of the construction of the embryological structure and differentiation in biochemical terms. This research programme produced a vast amount of experimental evidence and theories on the chemical dynamics of the embryo: particularly chemical characterization of the zygote and the developing embryo, the chemical exchanges between the nucleus and the cytoplasm, the (...)
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  22. Emmanuel Farge (2013). Mechano-Sensing in Embryonic Biochemical and Morphologic Patterning: Evolutionary Perspectives in the Emergence of Primary Organisms. [REVIEW] Biological Theory 8 (3):232-244.
    Embryogenesis involves biochemical patterning as well as mechanical morphogenetic movements, both regulated by the expression of the regulatory genes of development. The reciprocal interplay of morphogenetic movements with developmental gene expression is becoming an increasingly intense subject of investigation. The molecular processes through which differentiation patterning closely regulates the development of morphogenetic movements are today becoming well understood. Conversely, experimental evidence recently revealed the involvement of mechanical cues due to morphogenetic movements in activating mechano-transduction pathways that control both the differentiation (...)
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  23. Gary Felsenfeld (2014). The Evolution of Epigenetics. Perspectives in Biology and Medicine 57 (1):132-148.
    Since the early days of embryology, a central puzzle for biologists has been how a fertilized egg can execute a clearly defined and reproducible program that leads ultimately to a complex organism. It was clear that all of the information necessary to create the adult must already reside in the zygote, but how that information was translated into a complex organism was obscure. Even as recently as the late 1940s, the molecular mechanisms associated with early development were unknown and, in (...)
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  24. Jean-Louis Fischer & Julian Smith (1984). French Embryology and the « Mechanics of Development » From 1887 to 1910: L. Chabry, Y. Delage & E. Bataillon. History and Philosophy of the Life Sciences 6 (1):25 - 39.
  25. Loïc Forest & Jacques Demongeot (2008). A General Formalism for Tissue Morphogenesis Based on Cellular Dynamics and Control System Interactions. Acta Biotheoretica 56 (1):51-74.
    Morphogenesis is a key process in developmental biology. An important issue is the understanding of the generation of shape and cellular organisation in tissues. Despite of their great diversity, morphogenetic processes share common features. This work is an attempt to describe this diversity using the same formalism based on a cellular description. Tissue is seen as a multi-cellular system whose behaviour is the result of all constitutive cells dynamics. Morphogenesis is then considered as a spatiotemporal organization of cells activities. We (...)
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  26. Loïc Forest, Jaime San Martín, Fernando Padilla, Fabrice Chassat, Françoise Giroud & Jacques Demongeot (2004). Morphogenetic Processes: Application to Cambial Growth Dynamics. Acta Biotheoretica 52 (4):415-438.
    Both the physiological and the pathological morphogenetic processes that we can meet in embryogenesis, neogenesis and degenerative dysgenesis present common features: they are ruled by three different kinds of mechanisms, one related to cell migration, the second to cell differentiation and the third to cell proliferation. We deal here with an application to the cambial growth which essentially involves the third type of mechanism.Woody plants produce secondary tissue (secondary xylem and phloem) from a meristematic tissue called vascular cambium, responsible for (...)
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  27. Charles Galperin (1998). From Cell Lineage to Developmental Genetics. History and Philosophy of the Life Sciences 20 (3):301 - 350.
    One of the bases of developmental genetics resides in the alliance of clonal analysis and genetic analysis. But the study of cell lineage — cells which have their genealogical relationship — and the study of the cellular labelled progeny, have their own history. We have tried to follow it since its foundation with C.O. Whitman (1878) and E.B. Wilson (1892). A.H. Sturtevant (1929) and C. Stern (1936) the first tools to study the 'cell lineage' in Drosophila. We stress the contribution (...)
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  28. Claudia Lorena García (2007). Cognitive Modularity, Biological Modularity and Evolvability. Biological Theory: Integrating Development, Evolution and Cognition (KLI) 2 (1):62-73.
    There is an argument that has recently been deployed in favor of thinking that the mind is mostly (or even exclusively) composed of cognitive modules; an argument that draws from some ideas and concepts of evolutionary and of developmental biology. In a nutshell, the argument concludes that a mind that is massively composed of cognitive mechanisms that are cognitively modular (henceforth, c-modular) is more evolvable than a mind that is not c-modular (or that is scarcely c-modular), since a cognitive mechanism (...)
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  29. Scott F. Gilbert (2011). Expanding the Temporal Dimensions of Developmental Biology: The Role of Environmental Agents in Establishing Adult-Onset Phenotypes. Biological Theory 6 (1):65-72.
  30. Scott F. Gilbert (2006). The Generation of Novelty: The Province of Developmental Biology. Biological Theory 1 (2):209-212.
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  31. B. Goodwin (1985). What Are the Causes of Morphology. Bioessays 5:32-36.
  32. Paul E. Griffiths & Russell D. Gray (2004). The Developmental Systems Perspective: Organism-Environment Systems as Units of Development and Evolution. In Massimo Pigliucci & Katherine Preston (eds.), Phenotypic Integration: Studying the Ecology and Evolution of Complex Phenotypes. Oxford University Press 409--431.
    Developmental systems theory is an attempt to sum up the ideas of a research tradition in developmental psychobiology that goes back at least to Daniel Lehrman’s work in the 1950s. It yields a representation of evolution that is quite capable of accommodating the traditional themes of natural selection and also the new results that are emerging from evolutionary developmental biology. But it adds something else - a framework for thinking about development and evolution without the distorting dichotomization of biological processes (...)
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  33. Clifford Grobstein (1985). The Early Development of Human Embryos. Journal of Medicine and Philosophy 10 (3):213-236.
    The development of the human embryo from the time of fertilization through the eighth post-fertilization week is described for medical policy purposes. During pre-implantation stages, differentiation occurs between precursors of embryonic and extra-embryonic structures. During implantation formation of a fore-hind axis begins within the inner cell mass. By the end of the eighth week, head, face, hands, and feet are suggestive as to species-recognition but not yet definitive. Data from laboratory studies of non-human mammalian embryos elucidate important aspects of human (...)
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  34. Ulrich Krohs (2009). The Cost of Modularity. In Ulrich Krohs & Peter Kroes (eds.), Functions in Biological and Artificial Worlds: Comparative Philosophical Perspectives. MIT Press
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  35. Sandra D. Mitchell (2008). Exporting Causal Knowledge in Evolutionary and Developmental Biology. Philosophy of Science 75 (5):697-706.
    In this article I consider the challenges for exporting causal knowledge raised by complex biological systems. In particular, James Woodward’s interventionist approach to causality identified three types of stability in causal explanation: invariance, modularity, and insensitivity. I consider an example of robust degeneracy in genetic regulatory networks and knockout experimental practice to pose methodological and conceptual questions for our understanding of causal explanation in biology. †To contact the author, please write to: Department of History and Philosophy of Science, University of (...)
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  36. Philipp Mitteroecker & Simon M. Huttegger (2009). The Concept of Morphospaces in Evolutionary and Developmental Biology: Mathematics and Metaphors. Biological Theory 4 (1):54-67.
    Formal spaces have become commonplace conceptual and computational tools in a large array of scientific disciplines, including both the natural and the social sciences. Morphological spaces are spaces describing and relating organismal phenotypes. They play a central role in morphometrics, the statistical description of biological forms, but also underlie the notion of adaptive landscapes that drives many theoretical considerations in evolutionary biology. We briefly review the topological and geometrical properties of the most common morphospaces in the biological literature. In contemporary (...)
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  37. Massimo Pigliucci (2013). Between Holism and Reductionism: A Philosophical Primer on Emergence. Biological Journal of the Linnean Society 112 (2):261-267.
    Ever since Darwin a great deal of the conceptual history of biology may be read as a struggle between two philosophical positions: reductionism and holism. On the one hand, we have the reductionist claim that evolution has to be understood in terms of changes at the fundamental causal level of the gene. As Richard Dawkins famously put it, organisms are just ‘lumbering robots’ in the service of their genetic masters. On the other hand, there is a long holistic tradition that (...)
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  38. Michael Ruse (2006). Scott F. Gilbert?The Generation of Novelty: The Province of Developmental Biology Bare-Knuckle Fighting: EvoDevo Versus Natural Selection. Biological Theory 1 (4):402-403.
  39. Raphael Scholl & Massimo Pigliucci (2014). The Proximate–Ultimate Distinction and Evolutionary Developmental Biology: Causal Irrelevance Versus Explanatory Abstraction. Biology and Philosophy 2014 (5):DOI: 10.1007/s10539-014-9427-1.
    Mayr’s proximate–ultimate distinction has received renewed interest in recent years. Here we discuss its role in arguments about the relevance of developmental to evolutionary biology. We show that two recent critiques of the proximate–ultimate distinction fail to explain why developmental processes in particular should be of interest to evolutionary biologists. We trace these failures to a common problem: both critiques take the proximate–ultimate distinction to neglect specific causal interactions in nature. We argue that this is implausible, and that the distinction (...)
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  40. Kim Sterelny, Symbiosis, Evolvability and Modularity.
    This paper explores the connections between inheritance systems, evolvability and modularity. I argue that the transmission of symbiotic micro-organisms is an inheritance system, and one that is evolutionarily significant because symbionts generate biologically crucial aspects of their hosts’ organisation through modular developmental pathways. More specifically, I develop and defend five theses.
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  41. Rasmus Grønfeldt Winther (2001). Varieties of Modules: Kinds, Levels, Origins, and Behaviors. Journal of Experimental Zoology 291:116-129.
    This article began as a review of a conference, organized by Gerhard Schlosser, entitled “Modularity in Development and Evolution.” The conference was held at, and sponsored by, the Hanse Wissenschaftskolleg in Delmenhorst, Germany in May, 2000. The article subsequently metamorphosed into a literature and concept review as well as an analysis of the differences in current perspectives on modularity. Consequently, I refer to general aspects of the conference but do not review particular presentations. I divide modules into three kinds: structural, (...)
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  42. Rasmus Grønfeldt Winther (2001). August Weismann on Germ-Plasm Variation. Journal of the History of Biology 34 (3):517-555.
    August Weismann is famous for having argued against the inheritance of acquired characters. However, an analysis of his work indicates that Weismann always held that changes in external conditions, acting during development, were the necessary causes of variation in the hereditary material. For much of his career he held that acquired germ-plasm variation was inherited. An irony, which is in tension with much of the standard twentieth-century history of biology, thus exists – Weismann was not a Weismannian. I distinguish three (...)
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