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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).
    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. 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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  4. C. D. N. Barel (1993). Concepts of an Architectonic Approach to Transformation Morphology. Acta Biotheoretica 41 (4).
    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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  5. Ingo Brigandt (2009). Accounting for Vertebrate Limbs: From Owen's Homology to Novelty in Evo-Devo. Philosophy and 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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  6. 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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  7. 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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  8. Scott F. Gilbert (2006). The Generation of Novelty: The Province of Developmental Biology. Biological Theory 1 (2):209-212.
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  9. 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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  10. Ehud Lamm (2014). The Genome as a Developmental Organ. Journal of Physiology 592 (11):2237-2244.
    This paper applies the conceptual toolkit of Evolutionary Developmental Biology (evo‐devo) to the evolution of the genome and the role of the genome in organism development. This challenges both the Modern Evolutionary Synthesis, the dominant view in evolutionary theory for much of the 20th century, and the typically unreflective analysis of heredity by evo‐devo. First, the history of the marginalization of applying system‐thinking to the genome is described. Next, the suggested framework is presented. Finally, its application to the evolution of (...)
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  11. Paula M. Mabee (2006). Integrating Evolution and Development: The Need for Bioinformatics in Evo-Devo. BioScience 56 (4):301-309.
    Abstract -/- This article is an overview of concepts relating to the integration of the genotype and phenotype. One of the major goals of evolutionary developmental biology, or evo-devo, is to understand the transformation of morphology in evolution. This goal can be accomplished by synthesizing the data pertaining to gene regulatory networks and making use of the increasingly comprehensive knowledge of phylogenetic relationships and associated phenotypes. I give several examples of recent success in connecting these different biological levels. These examples (...)
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  12. 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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  13. Philipp Mitteroecker & Simon M. Huttegger (2009). The Concept of Morphospaces in Evolutionary and Developmental Biology: Mathematics and Metaphors. Biological Theory 4 (1):54-67.
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  14. Massimo Pigliucci (2013). Between Holism and Reductionism: A Philosophical Primer on Emergence. Biological Journal of the Linnean Society.
    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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  15. 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.
  16. Raphael Scholl & Massimo Pigliucci (forthcoming). The Proximate–Ultimate Distinction and Evolutionary Developmental Biology: Causal Irrelevance Versus Explanatory Abstraction. Biology and Philosophy:1-18.
    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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  17. 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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  18. 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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  19. 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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