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Developmental constraints are restrictions on the development of a particular phenotypic trait. These are physical, mechanical or structural limitations as well as irreversible commitments at key developmental stages that will limit or bias the phenotypes that selection can work with. Accordingly, developmental constraints (and not stabilizing selection) explain why certain traits, such as the structure of the tetrapod limb, are highly conserved in different evolving lineages with different adaptive pressures (e.g.  whale fins and frog legs).

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  1. L. Almeida & J. Demongeot (2012). Predictive Power of “A Minima” Models in Biology. Acta Biotheoretica 60 (1-2):3-19.
    Many apparently complex mechanisms in biology, especially in embryology and molecular biology, can be explained easily by reasoning at the level of the “efficient cause” of the observed phenomenology: the mechanism can then be explained by a simple geometrical argument or a variational principle, leading to the solution of an optimization problem, for example, via the co-existence of a minimization and a maximization problem . Passing from a microscopic level to the macroscopic level often involves an averaging effect that gives (...)
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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. Ron Amundson (1994). Two Concepts of Constraint: Adaptationism and the Challenge From Developmental Biology. Philosophy of Science 61 (4):556-578.
    The so-called "adaptationism" of mainstream evolutionary biology has been criticized from a variety of sources. One, which has received relatively little philosophical attention, is developmental biology. Developmental constraints are said to be neglected by adaptationists. This paper explores the divergent methodological and explanatory interests that separate mainstream evolutionary biology from its embryological and developmental critics. It will focus on the concept of constraint itself; even this central concept is understood differently by the two sides of the dispute.
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  4. Andre Ariew (1999). Innateness is Canalization: In Defense of a Developmental Account of Innateness. In Philosophy of Science. MIT Press, Cambridge, MA S19-S27.
    Lorenz proposed in his (1935) articulation of a theory of behavioral instincts that the objective of ethology is to distinguish behaviors that are “innate” from behaviors that are “learned” (or “acquired”). Lorenz’s motive was to open the investigation of certain “adaptive” behaviors to evolutionary theorizing. Accordingly, since innate behaviors are “genetic”, they are open to such investigation. By Lorenz’s light an innate/acquired or learned dichotomy rested on a familiar Darwinian distinction between genes and environments. Ever since Lorenz, ascriptions of innateness (...)
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  5. Tudor M. Baetu (2012). Mechanistic Constraints on Evolutionary Outcomes. Philosophy of Science 79 (2):276-294.
  6. Jonathan B. L. Bard (2008). Waddington's Legacy to Developmental and Theoretical Biology. Biological Theory 3 (3):188-197.
    Conrad Hal Waddington was a British developmental biologist who mainly worked in Cambridge and Edinburgh, but spent the late 1930s with Morgan in California learning about Drosophila. He was the first person to realize that development depended on the then unknown activities of genes, and he needed an appropriate model organism. His major experimental contributions were to show how mutation analysis could be used to investigate developmental mechanisms in Drosophila, and to explore how developmental mutation could drive evolution, his other (...)
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  7. William Bechtel (ed.) (1986). Integrating Scientific Disciplines. University of Chicago Press.
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  8. W. Berdel & G. Nass (1958). Über Die Bedeutung Des Auslesefaktors Im Rekapitulationsmechanismus der Phylogenetisch-Ontogenetischen Parallele. Acta Biotheoretica 12 (4):195-210.
    Haeckels theory of recapitulation shall be extended by the following rule: During the ontogenetical recapitulation of the phenotypical effects, the recapitulation of the phylogenetical natural selection factors according to the genotypical potentials is a condition of manifestation. The phylogenetical natural selectionfactor produces the activation of the gen as ontogenetical manifestation-stimulus. Factor of natural selection is the one of the extern or intern environment to which has happened the adaption in the phylogenesis. Concerning the intern environments the phenotypical effect of the (...)
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  9. Irene Berra (2014). An Evolutionary Ockham's Razor to Reciprocity. Frontiers in Theoretical and Philosophical Psychology 5:01258.
    Reciprocal altruism implies delayed payoffs by definition. It might therefore seem logical to assume that limited memory, calculation, and planning capacities have constrained the evolution of reciprocity in non-human animals. Here I will argue that this is not the case. First, I will show that the emotional track of past interactions is enough to motivate and maintain reciprocity over longer timespans. Second, I will propose a developmental pathway of this system of emotional bookkeeping. In particular, the neuropeptide modulation underlying mother-infant (...)
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  10. James Blachowicz (2013). The Constraint Interpretation of Physical Emergence. Journal for General Philosophy of Science / Zeitschrift für Allgemeine Wissenschaftstheorie 44 (1):21-40.
    I develop a variant of the constraint interpretation of the emergence of purely physical (non-biological) entities, focusing on the principle of the non-derivability of actual physical states from possible physical states (physical laws) alone. While this is a necessary condition for any account of emergence, it is not sufficient, for it becomes trivial if not extended to types of constraint that specifically constitute physical entities, namely, those that individuate and differentiate them. Because physical organizations with these features are in fact (...)
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  11. Ingo Brigandt (2015). From Developmental Constraint to Evolvability: How Concepts Figure in Explanation and Disciplinary Identity. In Alan C. Love (ed.), Conceptual Change in Biology: Scientific and Philosophical Perspectives on Evolution and Development. Springer 305-325.
    The concept of developmental constraint was at the heart of developmental approaches to evolution of the 1980s. While this idea was widely used to criticize neo-Darwinian evolutionary theory, critique does not yield an alternative framework that offers evolutionary explanations. In current Evo-devo the concept of constraint is of minor importance, whereas notions as evolvability are at the center of attention. The latter clearly defines an explanatory agenda for evolutionary research, so that one could view the historical shift from ‘developmental constraint’ (...)
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  12. 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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  13. Ingo Brigandt (2007). Typology Now: Homology and Developmental Constraints Explain Evolvability. Biology and Philosophy 22 (5):709-725.
    By linking the concepts of homology and morphological organization to evolvability, this paper attempts to (1) bridge the gap between developmental and phylogenetic approaches to homology and to (2) show that developmental constraints and natural selection are compatible and in fact complementary. I conceive of a homologue as a unit of morphological evolvability, i.e., as a part of an organism that can exhibit heritable phenotypic variation independently of the organism’s other homologues. An account of homology therefore consists in explaining how (...)
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  14. Gérard Cusset (1991). Les modeLes Sigmoides En Biologie Vegetale. Acta Biotheoretica 39 (3-4):197-205.
    Observed biological growth curves generally are sigmoid in appearance. It is common practice to fit such data with either a Verhulst logistic or a Gompertz curve. This paper critically considers the conceptual bases underlying these descriptive models.The logistic model was developed by Verhulst to accommodate the common sense observation that populations cannot keep growing indefinitely. A justification for using the same equation to describe the growth of individuals, based on considerations from chemical kinetics (autocatalysis of a monomolecular reaction), was put (...)
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  15. 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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  16. 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.
  17. Barker Desjardins & Pearce (eds.) (forthcoming). Entangled Life: Organism and Environment in the Biological and Social Sciences. Springer.
  18. 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.
  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. Barbara L. Finlay, Richard B. Darlington & Nicholas Nicastro (2001). Author's Response: Developmental Structure in Brain Evolution. Behavioral and Brain Sciences 24 (2):298-304.
    First, we clarify the central nature of our argument: our attempt is to apportion variation in brain size between developmental constraint, system-specific change, and change, underlining the unexpectedly large role of developmental constraint, but making no case for exclusivity. We consider the special cases of unusual hypertrophy of single structures in single species, regressive nervous systems, and the unusually variable cerebellum raised by the commentators. We defend the description of the cortex (or any developmentally-constrained structure) as a potential spandrel, and (...)
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  22. John R. Finnerty (2005). Did Internal Transport, Rather Than Directed Locomotion, Favor the Evolution of Bilateral Symmetry in Animals? Bioessays 27 (11):1174-1180.
  23. 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.
  24. 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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  25. Frietson Galis & Johan A. J. Metz (2003). Anti‐Cancer Selection as a Source of Developmental and Evolutionary Constraints. Bioessays 25 (11):1035-1039.
  26. Frietson Galis & Johan A. J. Metz (2003). Anti‐Cancer Selection as a Source of Developmental and Evolutionary Constraints. Bioessays 25 (11):1035-1039.
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  27. 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.
  28. B. Goodwin (1985). What Are the Causes of Morphology. Bioessays 5:32-36.
  29. Stephen Jay Gould (1992). Roots: Ontogeny and Phylogeny – Revisited and Reunited. Bioessays 14 (4):275-279.
  30. 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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  31. 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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  32. J. B. Gurdon (1992). In Search of New Principles of Development Biological Asymmetry and Handedness (1991). Ciba Symposium 162, Ed. Gregory R. Bock AND Joan Marsh. John Wiley. PP.iX+327. £47.40 ISBN 0 471 92961 1. [REVIEW] Bioessays 14 (6):427-427.
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  33. Alex Gurwitsch & Lydia Gurwitsch (1936). Der Feldbegriff in Seiner Anwendung Auf Das Problem der Zellteilung. Acta Biotheoretica 2 (2):77-92.
    The authors show that in certain isolated tissues the mitotic processes continue during at least one hour. They are very strongly stimulated by heat and also by mitogenetic radiation. In the case of the cancer cells new mitoses are promoted in considerable number. A detailed analysis of both energy factors leads to the conclusion that they effect a disturbance of the unstable constellations of the elementary particles in the cell-body. Thus a certain degree of disorganisation of its plasma seems to (...)
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  34. B. Hannigan (1996). Understanding Ageing By Robin Holliday. Bioessays 18:852-852.
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  35. Thomas F. Hansen & David Houle (2004). Evolvability, Stabilizing Selection, and the Problem of Stasis. In Massimo Pigliucci & Katherine Preston (eds.), Phenotypic Integration: Studying the Ecology and Evolution of Complex Phenotypes. Oxford University Press
  36. Claudia Harris & William Brown (1990). Developmental Constraints on Ethical Behavior in Business. Journal of Business Ethics 9 (11):855 - 862.
    Ethical behavior — the conscious attempt to act in accordance with an individually-owned morality — is the product of an advanced stage of the maturing process. Three models of ethical growth derived from research in human development are applied to issues of business ethics.
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  37. J. Herkovits & J. Faber (1978). Shape: Its Development and Regulation Capacity During Embryogenesis. Acta Biotheoretica 27 (3-4):185-200.
    Although several theoretical approaches consider general methods for dealing with shape, recent observations and experimental data show that embryos exhibit marked changes in the properties of the biological material involved in shape development and shape regulation capacity. In vivo experiments have shown that the amphibian embryo gradually develops from a situation in which it is not able to maintain its shape to one in which it can not only maintain its shape but also possesses a maximal tolerance towards deformation together (...)
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  38. Marvin E. Kirsh, Space: Negative Selection, Physical Constraint and Symmetry.
    A descriptive role is suggested for uracil as a temporal divide in the immediate aspects of metabolism verses long term maintained genetic transmission. In particular, details of the mechanism of excision repair of uracil from DNA based on differential parameters of spatial distortion of the planar uracil molecule within the DNA helix verses RNA, when viewed in analogy to a proposed model for space involving the substitution of the act of mirroring for the element of time in processes and a (...)
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  39. Horst Krist (2001). The Internalization of Physical Constraints From a Developmental Perspective. Behavioral and Brain Sciences 24 (4):681-682.
    Shepard's internalization concept is defended against Hecht's criticisms. By ignoring both Shepard's evolutionary perspective and the fact that internalization does not preclude modularization, Hecht advances inconclusive evidence. Developmental research supports Shepard's conclusion that kinematic geometry may be more deeply internalized than physical dynamics. This research also suggests that the internalization concept should be broadened to include representations acquired during ontogeny. [Hecht; Shepard].
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  40. Anne Dambricourt Malassé (1995). Les Attracteurs Inedits de L'Hominisation. Acta Biotheoretica 43 (1-2):113-125.
    The recent discovery of a phenomenon of craniofacial growth, called craniofacial contraction, throws a new light on the process of hominization. The main interest of this discovery lies in a growth principle combining the different craniofacial units, that is to say, the neurocranium, the chondrocranium and the splanchnocranium. Until recent years, these different parts were considered as neighbouring element without any morphogenic or morphodynamic connection. But now, we know that the morphogenesis of the base of the skull governs that of (...)
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  41. Lawrence E. Marks & Eric C. Odgaard (2005). Developmental Constraints on Theories of Synesthesia. In Robertson, C. L. & N. Sagiv (eds.), Synesthesia: Perspectives From Cognitive Neuroscience. Oxford University Press
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  42. Trevor Pearce (2011). Evolution and Constraints on Variation: Variant Specification and Range of Assessment. Philosophy of Science 78 (5):739-751.
    There is still a great deal of debate over what counts as a constraint and about how to assess experimentally the relative importance of constraints and selection in evolutionary history. I will argue that the notion of a constraint on variation, and thus the selection-constraint distinction, depends on two specifications: (1) what counts as a variant -- constraints limit or bias the production of what? and (2) range of assessment -- over what range of times or conditions is the variation (...)
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  43. Massimo Pigliucci (2007). Finding the Way in Phenotypic Space: The Origin and Maintenance of Constraints on Organismal Form. Annals of Botany 100:433-438.
    Background: One of the all-time questions in evolutionary biology regards the evolution of organismal shapes, and in particular why certain forms appear repeatedly in the history of life, others only seldom and still others not at all. Recent research in this field has deployed the conceptual framework of constraints and natural selection as measured by quantitative genetic methods. -/- Scope: In this paper I argue that quantitative genetics can by necessity only provide us with useful statistical sum- maries that may (...)
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  44. Massimo Pigliucci (2007). Finding the Way in Phenotypic Space: The Origin and Maintenance of Constraints on Organismal Form. Annals of Botany 100:433-438.
    Background: One of the all-time questions in evolutionary biology regards the evolution of organismal shapes, and in particular why certain forms appear repeatedly in the history of life, others only seldom and still others not at all. Recent research in this field has deployed the conceptual framework of constraints and natural selection as measured by quantitative genetic methods. Scope: In this paper I argue that quantitative genetics can by necessity only provide us with useful statistical sum- maries that may lead (...)
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  45. Massimo Pigliucci (ed.) (2004). Phenotypic Integration: Studying the Ecology and Evolution of Complex Phenotypes. Oxford University Press.
    A new voice in the nature-nurture debate can be heard at the interface between evolution and development. Phenotypic integration is a major growth area in research.
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  46. Massimo Pigliucci & Katherine Preston (eds.) (2004). Phenotypic Integration: Studying the Ecology and Evolution of Complex Phenotypes. Oxford University Press.
    A new voice in the nature-nurture debate can be heard at the interface between evolution and development. Phenotypic integration is a major growth area in research.
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  47. Robertson, C. L. & N. Sagiv (eds.) (2005). Synesthesia: Perspectives From Cognitive Neuroscience. Oxford University Press.
    The research presented in this volume demonstrates that it is no longer reasonable to ask whether or not synesthesia is real--we must now ask how we can account ...
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  48. Roger Sansom (2009). The Nature of Developmental Constraints and the Difference-Maker Argument for Externalism. Biology and Philosophy 24 (4):441-459.
    One current version of the internalism/externalism debate in evolutionary theory focuses on the relative importance of developmental constraints in evolutionary explanation. The received view of developmental constraints sees them as an internalist concept that tend to be shared across related species as opposed to selective pressures that are not. Thus, to the extent that constraints can explain anything, they can better explain similarity across species, while natural selection is better able to explain their differences. I challenge both of these aspects (...)
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  49. William C. Wimsatt (1986). Developmental Constraints, Generative Entrenchment, and the Innate-Acquired Distinction. In William Bechtel (ed.), Integrating Scientific Disciplines. 185--208.
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