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’ towards ‘evolvability’ as the move from a concept that is a mere tool of criticism to a concept that establishes a positive explanatory project. However, by taking a look at how the concept of constraint was employed in the 1980s, I argue that developmental constraint was not just seen as restricting possibilities (‘constraining’), but also as facilitating morphological change in several ways. Accounting for macroevolutionary transformation and the origin of novel form was an aim of these developmental approaches to evolution. Thus, the concept of developmental constraint was part of a positive explanatory agenda long before the advent of Evo-devo as a genuine scientific discipline. In the 1980s, despite the lack of a clear disciplinary identity, this concept coordinated research among paleontologists, morphologists, and developmentally inclined evolutionary biologists. I discuss the different functions that scientific concepts can have, highlighting that instead of classifying or explaining natural phenomena, concepts such as ‘developmental constraint’ and ‘evolvability’ are more important in setting explanatory agendas so as to provide intellectual coherence to scientific approaches. The essay concludes with a puzzle about how to conceptually distinguish evolvability and selection. (shrink)

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 an organism’s developmental constitution results in different homologues/characters as units that can evolve independently of each other. The explanans of an account of homology is developmental, yet the very explanandum is an evolutionary phenomenon: evolvability in a character-by-character fashion, which manifests itself in phylogenetic patterns as recognized by phylogenetic approaches to homology. While developmental constraints and selection have often been viewed as antagonistic forces, I argue that both are complementary as they concern different parts of the evolutionary process. Developmental constraints, conceived of as the presence of the same set of homologues across phenotypic change, pertain to how heritable variation can be generated in the first place (evolvability), while natural selection operates subsequently on the produced variation. (shrink)

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.

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 of the received view and propose a hierarchical view of constraints. (shrink)

In this paper, I argue that Kant adopted, throughout his career, a position that is much more akin to classical accounts of epigenesis, although he does reject the more radical forms of epigenesis proposed in his own time, and does make use of preformationist sounding terms. I argue that this is because Kant thinks of what is pre-formed as a species, not an individual or a part of an individual; has no qualm with the idea of a specific, teleological principle (...) or force underlying generation, and conceives of germs and predispositions as specific constraints on such a principle or force. Neither of these conceptions of what is “preformed”, I argue, is in strict opposition to classical epigenesis. I further suggest that Kant’s lingering use of preformationist terminology is due to his belief that this is required to account for the specificity of the specific generative force; his resistance towards the unrestricted plasticity of the generative force in radical epigenesis, which violates species-fixism; and his insistence on the internal, organic basis of developmental plasticity and variation within species. I conclude by suggesting that this terminological and interpretative peculiarity is partly due to a larger shift in the natural philosophical concerns surrounding the debate on epigenesis and preformation. Specifically, it is a sign that the original reasons for resisting epigenesis, namely its use of specific, teleological principles and its commitment to the natural production of biological structure, became less of a concern, whereas unrestricted plasticity and its undermining of fixism became a real issue, thereby also becoming the focal point of the debate. (shrink)

In this paper, I argue that Kant adopted, throughout his career, a position that is much more akin to classical accounts of epigenesis, although he does reject the more radical forms of epigenesis proposed in his own time, and does make use of preformationist sounding terms. I argue that this is because Kant (1) thinks of what is pre-formed as a species, not an individual or a part of an individual; (2) has no qualm with the idea of a specific, (...) teleological principle or force underlying generation, and conceives of germs and predispositions as specific constraints on such a principle or force. Neither of these conceptions of what is “preformed”, I argue, is in strict opposition to classical epigenesis. I further suggest that Kant’s lingering use of preformationist terminology is due to (1) his belief that this is required to account for the specificity of the specific generative force; (2) his resistance towards the unrestricted plasticity of the generative force in radical epigenesis, which violates species-fixism; and (3) his insistence on the internal, organic basis of developmental plasticity and variation within species. I conclude by suggesting that this terminological and interpretative peculiarity is partly due to a larger shift in the natural philosophical concerns surrounding the debate on epigenesis and preformation. Specifically, it is a sign that the original reasons for resisting epigenesis, namely its use of specific, teleological principles and its commitment to the natural production of biological structure, became less of a concern, whereas unrestricted plasticity and its undermining of fixism became a real issue, thereby also becoming the focal point of the debate. (shrink)

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. (shrink)

Studies performed in absence of gravitational constraint show that a living system is unable to choose between two different phenotypes, thus leading cells to segregate into different, alternative stable states. This finding demonstrates that the genotype does not determine by itself the phenotype but requires additional, physical constraints to finalize cell differentiation. Constraints belong to two classes: holonomic (independent of the system's dynamical states, as being established by the space‐time geometry of the field) and non‐holonomic (modified during those (...) biological processes to which they contribute in shaping). This latter kind of “constraints”, in which dynamics works on the constraint to recreate them, have emerged as critical determinants of self‐organizing systems, by manifesting a “closure of constraints.” Overall, the constraints act by harnessing the “randomness” represented by the simultaneous presence of equiprobable events restraining the system within one attractor. These results cast doubt on the mainstream scientific concept and call for a better understanding of causation in cell biology. (shrink)

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].

Recently at least two papers1,2have appeared that look at cancer from an evolutionary perspective. That cancer has a negative effect on fitness needs no argument. However, cancer origination is not an isolated process, but the potential for it is linked in diverse ways to other genetically determined developmental events, complicating the way selection acts on it, and through it on the evolution of development. The two papers take a totally different line. Kavanagh argues that anti‐cancer selection has led to (...) developmental constraints. Leroi et al. argue that cancer is a side‐effect of recent evolutionary changes that usually will disappear over time through anti‐cancer selection. Here we place the papers in a wider perspective, and in so doing discuss various alternative developmental links cancer may have together with their evolutionary implications. BioEssays 25:1035–1039, 2003. © 2003 Wiley Periodicals, Inc. (shrink)

This chapter contains sections titled: Introduction The Evolving Evolutionary Synthesis Population Genetics vs. Developmental Genetics Evo‐Devo's Central Target: The Study of Evolvability Origins? Conclusion Postscript: Counterpoint Acknowledgments References.

Evolutionary developmental biology (Evo-devo) is a loose conglomeration of research programs in the life sciences with two main axes: (a) the evolution of development, or inquiry into the pattern and processes of how ontogeny varies and changes over time; and, (b) the developmental basis of evolution, or inquiry into the causal impact of ontogenetic processes on evolutionary trajectories—both in terms of constraint and facilitation. Philosophical issues are found along both axes surrounding concepts such as evolvability, novelty, and modularity. (...) The developmental basis of evolution has garnered much attention because it speaks to the possibility of revising a standard construal of evolutionary theory, but the evolution of development harbors its own conceptual questions. This article addresses the heterogeneity of Evo-devo’s conglomerate structure (including disagreements over its individuation), as well as the concepts and controversies of philosophical interest pertaining to the evolution of development and the developmental basis of evolution. Future research will benefit from a shift away from global theorizing toward the scientific practices of Evo-devo. (shrink)

The concept of scaffolding has wide resonance in several scientific fields. Here we attempt to adopt it for the study of development. In this perspective, the embryo is conceived as an integral whole, comprised of several hierarchical modules as in a recurrent circularity of emerging patterns. Within the developmental hierarchy, each module yields an inter-level relationship that makes it possible for the scaffolding to mediate the production of selectable variations. A wide range of genetic, cellular and morphological mechanisms allows (...) the scaffolding to integrate these modular variations into a functionally coordinate unit. A genetic scaffolding accounts for the inherited invariance of pattern formation during the embryo’s growth. At higher level, cells behave as agents endowed with the capacity to interpret any scaffolding variation as signs. The full hierarchy of a multi-level scaffolding is eventually attained when the embryo acquires the capacity to impose a number of developmental constraints on its constituting parts in a top-down direction. The acquisition of this capacity allows a semiotic threshold to emerge between the living cellular world and the underlying non-living molecular world. As this boundary is gradually defined during development, cells enter into new functional relationships, while, at the same time, are relieved from their physical determinism. The resulting constraints can thus become the driving forces that upgrade embryonic scaffolding from the simple molecular signalling to the complexity of sign recognition proper of a cellular community. In this semiotic perspective, the apparent goal directness of any developmental strategy should no longer be accounted for by a predetermined genetic program, but by the gradual definition of the relationships selected amongst the ones historically explored. (shrink)

There exist many competing philosophical definitions of joint action and no clear criteria to decide between them; so far the search for definitions has by and large been a semantical enterprise rather than an empirical one. This chapter describes and assesses several constraints that could help converge towards a set of necessary and sufficient conditions for joint action. The tightness constraint favours definitions that fit joint actions in which the links between agents are as relaxed as possible, so as (...) to better pinpoint the conceptual core of jointness. The developmental constraint asks for definitions based on realistic psychological states that could be entertained by agents less cognitively developed than ideal human beings. The motor constraint holds that definitions should refer to psychological mechanisms involved in actual human coordination. These first three constraints are discussed and dismissed, mainly because they manage to establish vague limits at best (for various reasons). I then introduce a fourth one, the efficiency constraint, based on the fact that most of our joint actions are generally successful, and according to which definitions should involve conditions that help justify this success. Finally, the rational and evolutionary versions of the efficiency constraint are examined and defended against objections. (shrink)

Throughout the recent history of research at the intersection of evolution and development, notions such as developmental constraint, evolutionary novelty, and evolvability have been prominent, but the term ‘developmental bias’ has scarcely been used. And one may even doubt whether a unique and principled definition of bias is possible. I argue that the concept of developmental bias can still play a vital scientific role by means of setting an explanatory agenda that motivates investigation and guides the formulation (...) of integrative explanatory frameworks. Less crucial is a definition that would classify patterns of phenotypic variation and unify variational patterns involving different traits and taxa as all being ‘bias.’ Instead, what we should want is a concept that generates intellectual identity across various researchers, and that unites the diverse fields and approaches relevant to the study of developmental bias, from paleontology to behavioral biology. I point to some advantages of conducting research specifically under the label of ‘developmental bias,’ compared to employing other, more common terms such as ‘evolvability.’. (shrink)

How micro- and macroevolutionary evolutionary processes produce phenotypic change is without question one of the most intriguing and perplexing issues facing evolutionary biologists. We believe that roadblocks to progress lie A) in the underestimation of the role of the environment, and in particular, that of the interaction of genotypes with environmental factors, and B) in the continuing lack of incorporation of development into the evolutionary synthesis. We propose the integration of genetic, environmental and developmental perspectives on the evolution of (...) the phenotype in the form of the concept of the developmental reaction norm (DRN) The DRN represents the set of multivariate ontogenies that can be produced by a single genotype when it is exposed to environmental variation. It encompasses: 1) the processes that alter the phenotype throughout the ontogenetic trajectory, 2) the recognition that different aspects of the phenotype are (and must be) correlated and 3) the ability of a genotype to produce phenotypes in different environments. This perspective necessitates the explicit study of character expression during development, the evaluation of associations between pairs or groups of characters (e.g., multivariate allometries), and the exploration of reaction norms and phenotypic plasticity. We explicitly extend the concept of the DRN to encompass adjustments made in response to changes in the internal environment as well. Thus, ‘typical’ developmental sequences (e.g., cell fate determination) and plastic responses are simply manifestations of different scales of ‘environmental’ effects along a continuum. We present: (1) a brief conceptual review of three fundamental aspects of the generation and evolution of phenotypes: the changes in the trajectories describing growth and differentiation (ontogeny), the multivariate relationships among characters (allometry), and the effect of the environment (plasticity); (2) a discussion of how these components are merged in the concept of the developmental reaction norm; and (3) a reaction norm perspective of major determinants of phenotypes: epigenesis, selection and constraint. (shrink)

This paper provides an account of the developmental origins of our belief in free will based on research from a range of ages—infants, preschoolers, older children, and adults—and across cultures. The foundations of free will beliefs are in infants' understanding of intentional action—their ability to use context to infer when agents are free to “do otherwise” and when they are constrained. In early childhood, new knowledge about causes of action leads to new abilities to imagine constraints on action. (...) Moreover, unlike adults, young children tend to view psychological causes (i.e., desires) and social causes (i.e., following rules or group norms, being kind or fair) of action as constraints on free will. But these beliefs change, and also diverge across cultures, corresponding to differences between Eastern and Western philosophies of mind, self, and action. Finally, new evidence shows developmentally early, culturally dependent links between free will beliefs and behavior, in particular when choice‐making requires self‐control. (shrink)

Philosophical discussion of molecular and developmental biology began in the late 1960s with the use of genetics as a test case for models of theory reduction. With this exception, the theory of natural selection remained the main focus of philosophy of biology until the late 1970s. It was controversies in evolutionary theory over punctuated equilibrium and adaptationism that first led philosophers to examine the concept of developmental constraint. Developmental biology also gained in prominence in the 1980s as (...) part of a broader interest in the new sciences of self-organization and complexity. The current literature in the philosophy of molecular and developmental biology has grown out of these earlier discussions under the influence of twenty years of rapid and exciting growth of empirical knowledge. Philosophers have examined the concepts of genetic information and genetic program, competing definitions of the gene itself and competing accounts of the role of the gene as a developmental cause. The debate over the relationship between development and evolution has been enriched by theories and results from the new field of 'evolutionary developmental biology'. Future developments seem likely to include an exchange of ideas with the philosophy of psychology, where debates over the concept of innateness have created an interest in genetics and development. (shrink)

How does evolution grow bigger brains? It has been widely assumed that growth of individual structures and functional systems in response to niche-specific cognitive challenges is the most plausible mechanism for brain expansion in mammals. Comparison of multiple regressions on allometric data for 131 mammalian species, however, suggests that for 9 of 11 brain structures taxonomic and body size factors are less important than covariance of these major structures with each other. Which structure grows biggest is largely predicted by a (...) conserved order of neurogenesis that can be derived from the basic axial structure of the developing brain. This conserved order of neurogenesis predicts the relative scaling not only of gross brain regions like the isocortex or mesencephalon, but also the level of detail of individual thalamic nuclei. Special selection of particular areas for specific functions does occur, but it is a minor factor compared to the large-scale covariance of the whole brain. The idea that enlarged isocortex could be a a by-product of structural constraints later adapted for various behaviors, contrasts with approaches to selection of particular brain regions for cognitively advanced uses, as is commonly assumed in the case of hominid brain evolution. (shrink)

More and more researchers are examining grammar acquisition from theoretical perspectives that treat it as an emergent phenomenon. In this essay, I argue that a robustly developmental perspective provides a potential explanation for some of the well-known crosslinguistic features of early child language: the process of acquisition is shaped in part by the developmental constraints embodied in von Baer’s law of development. An established model of development, the Developmental Lock, captures and elucidates the probabilistic generalizations at (...) the heart of von Baer’s law. When this model is applied to the acquisition of grammar, it predicts that grammatical achievements that are more generatively entrenched will emerge earlier in development and will be more developmentally resilient than those that are less generatively entrenched. I show that the first prediction is supported by a wealth of psycholinguistic evidence involving typically developing children and that the second prediction is supported by numerous studies involving both children who receive deficient linguistic input and children who experience various language impairments. The success of this model demonstrates the analytic potential of a developmental approach to the study of language acquisition. (shrink)

This book represents an effort to understand very old questions about biological form, function, and the relationships between them. The essays collected here reflect the diversity of approaches in evolutionary developmental biology, including not only studies by prominent scientists whose research focuses on topics concerned with evolution and development, but also historically and conceptually oriented studies that place the scientific work within a larger framework and ask how it can be pushed further. Topics under discussion range from the use (...) of theoretical and empirical biomechanics to understand the evolution of plant form, to detailed studies of the evolution of development and the role of developmental constraints on phenotypic variation. The result is a rich and interdisciplinary volume that will begin a wider conversation about the shape of Evo Devo as it matures as a field. (shrink)

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 should instead be understood in the context of explanatory abstractions in complete causal models of evolutionary change. Once the debate is reframed in this way, the proximate–ultimate distinction’s role in arguments against the theoretical significance of evo-devo is seen to rely on a generally implicit premise: that the variation produced by development is abundant, small and undirected. We show that a “lean version” of the proximate–ultimate distinction can be maintained even when this isotropy assumption does not hold. Finally, we connect these considerations to biological practice. We show that the investigation of developmental constraints in evolutionary transitions has long relied on a methodology which foregrounds the explanatory role of developmental processes. It is, however, entirely compatible with the lean version of the proximate–ultimate distinction. (shrink)

Stochastic gene expression plays a leading developmental role through its contribution to cell differentiation. It is also proposed to promote phenotypic diversification in malignant cells. However, it remains unclear if these two forms of cellular bet‐hedging are identical or rather display distinct features. Here we argue that bet‐hedging phenomena in cancer cells are more similar to those occurring in unicellular organisms than to those of normal metazoan cells. We further propose that the atavistic bet‐hedging strategies in cancer originate from (...) a hijacking of the normal developmental bet‐hedging of metazoans. Finally, we discuss the constraints that may shape the atavistic bet‐hedging strategies of cancer cells. (shrink)

Many natural and technological systems are complex, with organizational structures that exhibit characteristic patterns but defy concise description. One effective approach to analyzing such systems is in terms of repeated topological motifs. Here, we extend the motif concept to characterize the dynamic behavior of complex systems by introducing developmental motifs, which capture patterns of system growth. As a proof of concept, we use developmental motifs to analyze the developmental cell lineage of the nematode Caenorhabditis elegans, revealing a (...) new perspective on its complex structure. We use a family of computational models to explore how biases arising from the dynamics of the developmental gene network, as well as spatial and temporal constraints acting on development, contribute to this complex organization. © 2010 Wiley Periodicals, Inc. Complexity 16: 48‐57, 2011. (shrink)

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 weigh the implications of the spandrel concept for the course of human evolution. The empirical and statistical objections raised in the commentary of Barton are discussed at length. Finally, we catalogue and comment on the suggestions of new ways to study brain evolution, and new aspects of brain evolution to study. (shrink)

The interactivist approach to development generates a framework of types of constraints on what can be constructed. The four constraint types are based on: (1) what the constructed systems are about; (2) the representational relationship itself; (3) the nature of the systems being constructed; and (4) the process of construction itself. We give illustrations of each constraint type. Any developmental theory needs to acknowledge all four types of constraint; however, some current theories conflate different types of constraint, or (...) rely on a single constraint type to explicate development. Such theories will be inherently unable to explain important aspects of development. (shrink)

The interactivist approach to development generates a framework of types of constraints on what can be constructed. The four constraint types are based on: (1) what the constructed systems are about; (2) the representational relationship itself; (3) the nature of the systems being constructed; and (4) the process of construction itself. We give illustrations of each constraint type. Any developmental theory needs to acknowledge all four types of constraint; however, some current theories conflate different types of constraint, or (...) rely on a single constraint type to explicate development. Such theories will be inherently unable to explain important aspects of development. (shrink)

Ethics in business has been an increasingly controversial and important topic of discussion over the last decade. Debate continues about whether ethics should be a part of business, but also includes how business can implement ethical theory in day-to-day operations. Most discussions focus on either traditional moral philosophy, which offers little of practical value for the business community, or psychological theories of moral reasoning, which have been shown to be flawed and incomplete. The theory presented here is called the (...) class='Hi'>Developmental Self-Valuing Theory, and adapts the general psychological theory of Albert Bandura for ethics in business. In this theory, individuals first learn moral values from associations with others who are significant in their lives. Secondly, self-regulation is learned through a process of self-observation, self-judgment and self-reaction. Thirdly, the individual must believe that he can act ethically. Situational constraints and inducements, as well as positive and negative consequences for specific behaviors, will also affect the level of ethical performance. Each of these elements is examined and combined to achieve a practical method for increasing the level of ethics in corporate activity through selection, training and situational enhancement. (shrink)

This article focuses on the question of how we perceive and represent ourselves at the most minimal, pre-reflective level. We first review recent work emphasizing the multisensory basis of our perceptual experiences and the embodied nature of self-awareness. We then focus on interoceptive and tactile signals, as key components of bodily self-consciousness, and discuss one crucial yet overlooked aspect of our embodiment, namely the fact that bodily self-consciousness emerges from the outset within the body of another experiencing subject. Next, we (...) review empirical findings highlighting the developmental primacy of tactile and interoceptive over visual experiences, in shaping the foundations of perceptual awareness. Building upon the influential predictive processing framework in philosophy and theoretical neuroscience, we argue that self-awareness is not only embodied, but also that perceptual experiences are fundamentally 'selfish', that is, imperatively driven by basic constraints of physiological regulation that subserve self-preservation. We then evaluate. (shrink)

Since the advent of the internet, researchers have been interested in the intersubjective possibilities and constraints that digital environments offer users. In the literature, we find some who argue that seemingly disembodied digitally mediated interactions are severely limited when compared to their embodied face-to-face counterparts and others who are more optimistic about the possibilities that such technologies afford. Yet, both camps tend towards offering what we see as static accounts of online intersubjectivity – accounts that attempt to determine the (...) very possibilities and constraints of digital platforms based upon their design. What we think these approaches fail to take into account is how users’ intersubjective capabilities on digital platforms can evolve and change over time. Developmental psychology emphasises the way in which intersubjective capabilities build upon and are interrelated to one another, allowing for more sophisticated styles of intersubjectivity to emerge only once earlier stages are already in place. We find this suggestive for analysing intersubjectivity online as it provides a framework for thinking about how various an individual’s intersubjective capabilities might develop. We argue that in some cases this can happen through gradual adjustments of our basic intersubjective capabilities to digital spaces, especially those that strive to mimic established forms of offline interaction (e.g., Zoom). In other cases, this can involve more substantial processes of learning to walk and talk again online. Consequently, determining the possibilities and limits of online intersubjectivity is, at least in part, relative to a user’s skill, history, and familiarity with the technology in question. (shrink)

Although sciences are often conceptualized in terms of theory confirmation and hypothesis testing, an equally important dimension of scientific reasoning is the structure of problems that guide inquiry. This problem structure is evident in several concepts central to evolutionary developmental biology (Evo-devo)—constraints, modularity, evolvability, and novelty. Because problems play an important role in biological practice, they should be included in biological pedagogy, especially when treating the issue of scientific controversy. A key feature of resolving controversy is synthesizing methodologies (...) from different biological disciplines to generate empirically adequate explanations. Concentrating on problem structure illuminates this interdisciplinarity in a way that is often ignored when science is taught only from the perspective of theory or hypothesis. These philosophical considerations can assist life science educators in their continuing quest to teach biology to the next generation. -/- . (shrink)

FOREWORD. Gilbert Gottlieb and the Developmental Point of View. I. INTRODUCTION. 1. Developmental Systems, Nature-Nurture, and the Role of Genes in Behavior and Development: On the Legacy of Gilbert Gottlieb. 2. Normally Occurring Environmental and Behavioral Influences on Gene Activity: From Central Dogma to Probabilistic Epigenesis. II. THEORETICAL FOUNDATIONS FOR THE DEVELOPMENTAL STUDY OF BEHAVIOR AND GENETICS. 3. Historical and Philosophical Perspectives on Behavioral Genetics and Developmental Science. 4. Development and Evolution Revisited. 5. Probabilistic Epigenesis and (...) Modern Behavioral and Neural Genetics. 6. The Roles of Environment, Experience, and Learning in Behavioral Development. 7. Contemporary Ideas in Physics and Biology in Gottlieb’s Psychology. III. EMPIRICAL STUDIES OF BEHAVIORAL DEVELOPMENT AND GENETICS. 8. Behavioral Development during the Mother-Young Interaction in Placental Mammals: The Development of Behavior in the Relationship with the Mother. 9. Amniotic Fluid as an Extended Milieu Interieur. 10. Developmental Effects of Selective Breeding for an Infant Trait. 11. Emergence and Constraint in Novel Behavioral Adaptations. 12. Nonhuman Primate Research Contributions to Understanding Genetic and Environmental Influences on Phenotypic Outcomes across Development. 13. Interactive Contributions of Genes and Early Experience to Behavioural Development: Sensitive Periods and Lateralized Brain and Behaviour. 14. Trans-Generational Epigenetic Inheritance. 15. The Significance of Non-Replication of Gene-Phenotype Associations. 16. Canalization and Malleability Reconsidered: The Developmental Basis of Phenotypic Stability and Variability. IV. APPLICATIONS TO DEVELOPMENT. 17. Gene-Parenting Interplay in the Development of Infant Emotionality. 18. Genetic Research in Psychiatry and Psychology: A Critical Overview. 19. On the Limits of Standard Quantitative Genetic Modeling of Inter-Individual Variation: Extensions, Ergodic Conditions and a New Genetic Factor Model of Intra-Individual Variation. 20. Songs My Mother Taught Me: Gene-Environment Interactions, Brain Development and the Auditory System: Thoughts on Non-Kin Rejection, Part II. 21. Applications of Developmental Systems Theory to Benefit Human Development: On the Contributions of Gilbert Gottlieb to Individuals, Families, and Communities. (shrink)

It is becoming increasingly clear that acquiring cognitive skills is feasible only with significant developmental constraints. However, recent research provides the strongest evidence to date for constructivist development. Here, we examine how these two apparently conflicting perspectives may be reconciled. Specifically, we suggest that subcortical and cortical structures possess divergent developmental strategies, with many subcortical structures satisfying Fodor's criteria for modularity. These structures constitute an early behavioral system that guides the construction of later emerging cortical structures, for (...) which there is little evidence for modularity. Thus, we focus on how the dynamic time course of development itself implicitly constrains the emergence of cortical representations, reducing the requirement for built-in encodings of knowledge in cortical circuits, as on the traditional nativist conception. (shrink)

In an influential paper, Stephen Jay Gould and Richard Lewontin contrasted selection-driven adaptation with phylogenetic, architectural, and developmental constraints as distinct causes of phenotypic evolution. In subsequent publications Gould has elaborated this distinction into one between a narrow “Darwinian Fundamentalist” emphasis on “external functionalist” processes, and a more inclusive “pluralist” emphasis on “internal structuralist” principles. Although theoretical integration of functionalist and structuralist explanations is the ultimate aim, natural selection and internal constraints are treated as distinct causes of (...) evolutionary change. This distinction is now routinely taken for granted in the literature in evolutionary biology. I argue that this distinction is problematic because the effects attributed to non-selective constraints are more parsimoniously explained as the ordinary effects of selection itself. Although it may still be a useful shorthand to speak of phylogenetic, architectural, and developmental constraints on phenotypic evolution, it is important to understand that such “constraints” do not constitute an alternative set of causes of evolutionary change. The result of this analysis is a clearer understanding of the relationship between adaptation, selection and constraints as explanatory concepts in evolutionary theory. (shrink)

As the accompanying articles in the Special Issue on Semiotic Scaffolding will attest, my colleagues in biosemiotics have done an exemplary job in showing us how to think about the critically generative role that semiotic scaffolding plays “vertically” – i.e., in evolutionary and developmental terms – by “allowing access to the upper floors” of biological complexity, cognition and evolution.In addition to such diachronic considerations of semiotic scaffolding, I wish to offer here a consideration of semiotic scaffolding’s synchronic power, as (...) well – and in particular the ability that it can afford its users to access new and other sign relations “horizontally” as a function of the way that multiple semiotically scaffolded relations intertwine to result in a “definite semantic topology that determines the ways that symbols modify each other’s referential functions in different combinations”.Taking up, in turn, Terrence Deacon’s later challenge that what the sciences of cognition – and biology more generally – needs to “come to grips with [is] the process of semiosis; the dynamic of interpretive activity by which semiotic relationships emerge from other semiotic relationships [as] intrinsically dynamic phases in a generative process”, I attempt here to show how Deacon’s own Peirce-inspired matrix of referential sign relations as presented in The Symbolic Species, when viewed as a semiotic scaffold of interactional constraints and possibility biases, provides the key to understanding the essentially thirdness-manifesting nature of symbol reference, formation and growth. (shrink)

The target article highlights the sources of open-endedness of human communication. However, the authors' perspective does not account for the structure of particular communication systems. To this end, we extend the authors' perspective, in the spirit of evolutionary extended synthesis, with a detailed account of the sources of constraints imposed upon expression in the course of child development.

Infants' own activities create and actively select their learning experiences. Here we review recent models of embodied information seeking and curiosity-driven learning and show that these mechanisms have deep implications for development and evolution. We discuss how these mechanisms yield self-organized epigenesis with emergent ordered behavioral and cognitive developmental stages. We describe a robotic experiment that explored the hypothesis that progress in learning, in and for itself, generates intrinsic rewards: The robot learners probabilistically selected experiences according to their potential (...) for reducing uncertainty. In these experiments, curiosity-driven learning led the robot learner to successively discover object affordances and vocal interaction with its peers. We explain how a learning curriculum adapted to the current constraints of the learning system automatically formed, constraining learning and shaping the developmental trajectory. The observed trajectories in the robot experiment share many properties with those in infant development, including a mixture of regularities and diversities in the developmental patterns. Finally, we argue that such emergent developmental structures can guide and constrain evolution, in particular with regard to the origins of language. (shrink)

Adaptive phenotypic plasticity allows organisms to cope with environmental variability, and yet, despite its adaptive significance, phenotypic plasticity is neither ubiquitous nor infinite. In this review, we merge developmental and population genetic perspectives to explore costs and limits on the evolution of plasticity. Specifically, we focus on the role of modularity in developmental genetic networks as a mechanism underlying phenotypic plasticity, and apply to it lessons learned from population genetic theory on the interplay between relaxed selection and mutation (...) accumulation. We argue that the environmental specificity of gene expression and the associated reduction in pleiotropic constraints drive a fundamental tradeoff between the range of plasticity that can be accommodated and mutation accumulation in alternative developmental networks. This tradeoff has broad implications for understanding the origin and maintenance of plasticity and may contribute to a better understanding of the role of plasticity in the origin, diversification, and loss of phenotypic diversity. (shrink)

The end-state comfort effect refers to the consistent tendency of healthy adults to end their movements in a comfortable end posture. In children with and without developmental coordination disorder, the results of studies focusing on ESC planning have been inconclusive, which is likely to be due to differences in task constraints. The present pilot study focused on the question whether children with and without DCD were able to change their planning strategy and were more likely to plan for (...) ESC when demanded by complex object manipulations at the end of a task. To this end, we examined ESC planning in 18 children with and without DCD using the previously used sword-task and the newly developed hammer-task. In the sword-task, children had to insert a sword in a wooden block, which could be relatively easily completed with an uncomfortable end-posture. In the hammer-task, children had to strike down a nail in a wooden pounding bench, which required additional force and speed demands, making it relatively difficult to complete the movement with an uncomfortable end-posture. In line with our hypothesis, the results demonstrated that children with and without DCD were more likely to plan for ESC on the hammer-task compared with the sword-task. Thus, while children with and without DCD show inconsistent ESC planning on many previously used tasks, the present pilot study shows that many of them are able to take into account the end-state of their movements if demanded by task constraints. (shrink)

The growing interest and major advances of the last decades in evolutionary developmental biology (EvoDevo) have led to the recognition of the incompleteness of the Modern Synthesis of evolutionary theory. Here we discuss how paleontology makes significant contributions to integrate evolution and development. First, extinct organisms often inform us about developmental processes by showing a combination of features unrecorded in living species. We illustrate this point using the vertebrate fossil record and studies relating bone ossification to life history (...) traits. Second, we discuss exceptionally preserved fossils that document rare ontogenetic sequences and illustrate this case with the patterns of heterochrony observed in Cambrian crustacean larvae preserved three-dimensionally. Third, most fossils potentially document the evolutionary patterns of allometry and modularity, as well as some of the (paleo)ecological factors that had influenced them. The temporal persistence of adaptive patterns in rodent evolution serves to address the importance of ecological constraints in evolution. Fourth, we discuss how the macroevolutionary patterns observed in the tetrapod limb, in the mammal molar proportions, and in the molluscan shell provide independent tests of the validity of morphogenetic models proposed on living species. Reciprocally, these macroevolutionary patterns often act as a source of inspiration to investigate the underlying rules of development, because, at the end, they are the patterns that the neo-Darwinian theory was unable to account for. (shrink)

Cognitive frameworks provide important means for uniting concepts of specificity, cognition, and dynamic change in development. Two points are challenged by evidence from special populations: (1) that boundary constraints such as Residual Normality and a cognitive “endstate” compromise the use of cognitive models; and (2) the developmental process itself automatically rejects either Residual Normality or residual deviance from typical development.

I argue that it is time for many feminists to rethink their attitudes towards evolutionary biology, not because feminists have been wrong to be deeply sceptical about many of its claims, both explicit and implicit, but because biology itself has changed. A new appreciation for the importance of development in biology has become mainstream and a new ontology, associated with developmental systems theory, has been introduced over the last two decades. This turn challenges some of the features of evolutionary (...) biology that have most troubled feminists. DST undermines the idea of biologicales sence and challenges both nature / nurture and nature/ culture distinctions. Freed from these conceptual constraints, evolutionary biology no longer poses the problems that have justified feminist scepticism. Indeed, feminists have already found useful applications for DST and I argue that they should expand their use of DST to support more radical and wide-ranging political theories. Si les féministes n’ont pas eu tort d’être profondément sceptiques face aux nombreuses revendications de la biologie, leur attitude face à cette science doit être remise en question car la biologie s’est transformée au courant des dernières décennies. La «théorie des systèmes de développement» est une théorie qui s’est considérablement développée et qui a pris beaucoup d’ampleur. Cette théorie n’accepte pas le concept d’essence biologique ce qui pose un défi important à la distinction nature/ culture. Une des conséquences de cet apport théorique est que le scepticisme des féministes face à la biologie de l’évolution n’est plus justifié car la biologie ne comporte plus les contraintes essentialistes qui s’avéraient contentieuses. En effet, certaines féministes ont déjà trouvé des applications utiles pour la TDS et nous avançons que les féministes doivent maintenant élargir l’utilisation de la TDS car la porté de celle-ci pourrait être significative dans d’autres domaines tel celui de la théorie politique. (shrink)

Norman presents intriguing arguments in support of a mapping between ecological and constructivist visual cognition, on the one hand, onto the dorsal ventral dual route processing hypothesis, on the other hand. Unfortunately, his account is incompatible with developmental data on the functional emergence of the dorsal and ventral routes. We argue that it is essential for theories of adult visual cognition to take constraints from development seriously.

The principles for universal reading models proposed by Frost correspond to developmental theories, in which neurocognitive constraints and cultural experiences shape development. We question his contention that Hebrew word identification is fundamentally about roots, excluding verbal and nominal word-pattern morphemes; and we propose that readers use all information available in stimuli, adjusting for volume and usefulness.