This chapter analyzes the notion of human nature and the concept of inner nature from the perspective of developmental systems theory. It explores the folkbiology of human nature and looks at three features associated with traits that are expressions of the inner nature that organisms inherit from their parents: fixity, typicality, teleology.

What kind mechanisms one deems central for the evolutionary process deeply influences one's understanding of the nature of organisms, including cognition. Reversely, adopting a certain approach to the nature of life and cognition and the relationship between them or between the organism and its environment should affect one's view of evolutionary theory. This paper explores this reciprocal relationship in more detail. In particular it argues that the view of living and cognitive systems, especially humans, as deeply integrated beings (...) embedded in and transformed by their genetic, epigenetic, behavioral, ecological, socio-cultural and cognitive-symbolic legacies calls for an extended evolutionary synthesis that goes beyond either a theory of genes juxtaposed against a theory of cultural evolution and or even more sophisticated theories of gene-culture coevolution and niche construction. Environments, particularly in the form of developmental environments, do not just select for variation, they also create new variation by influencing development through the reliable transmission of non-genetic but heritable information. This paper stresses particularly views of embodied, embedded, enacted and extended cognition, and their relationship to those aspects of extended inheritance that lie between genetic and cultural inheritance, the still gray area of epigenetic and behavioral inheritance systems that play a role in parental effect. These are the processes that can be regarded as transgenerational developmental plasticity and that I think can most fruitfully contribute to, and be investigated by, developmental psychology. (shrink)

Developmental systems theory (DST) is a wholeheartedly epigenetic approach to development, inheritance and evolution. The developmental system of an organism is the entire matrix of resources that are needed to reproduce the life cycle. The range of developmental resources that are properly described as being inherited, and which are subject to natural selection, is far wider than has traditionally been allowed. Evolution acts on this extended set of developmental resources. From a (...) class='Hi'>developmental systems perspective, development does not proceed according to a preformed plan; what is inherited is much more than DNA; and evolution is change not only in gene frequencies, but in entire developmental systems. (shrink)

In 1809--the year of Charles Darwin's birth--Jean-Baptiste Lamarck published Philosophie zoologique, the first comprehensive and systematic theory of biological evolution. The Lamarckian approach emphasizes the generation of developmental variations; Darwinism stresses selection. Lamarck's ideas were eventually eclipsed by Darwinian concepts, especially after the emergence of the Modern Synthesis in the twentieth century. The different approaches--which can be seen as complementary rather than mutually exclusive--have important implications for the kinds of questions biologists ask and for the type (...) of research they conduct. Lamarckism has been evolving--or, in Lamarckian terminology, transforming--since Philosophie zoologique's description of biological processes mediated by "subtle fluids." Essays in this book focus on new developments in biology that make Lamarck's ideas relevant not only to modern empirical and theoretical research but also to problems in the philosophy of biology. Contributors discuss the historical transformations of Lamarckism from the 1820s to the 1940s, and the different understandings of Lamarck and Lamarckism; the Modern Synthesis and its emphasis on Mendelian genetics; theoretical and experimental research on such "Lamarckian" topics as plasticity, soft (epigenetic) inheritance, and individuality; and the importance of a developmental approach to evolution in the philosophy of biology. The book shows the advantages of a "Lamarckian" perspective on evolution. Indeed, the development-oriented approach it presents is becoming central to current evolutionary studies--as can be seen in the burgeoning field of Evo-Devo. Transformations of Lamarckism makes a unique contribution to this research. (shrink)

Organisms inherit various kinds of developmental information and cues from their parents. The study of inheritance systems is aimed at identifying and classifying the various mechanisms and processes of heredity, the types of hereditary information that is passed on by each, the functional interaction between the different systems, and the evolutionary consequences of these properties. We present the discussion of inheritance systems in the context of several debates. First, between proponents of monism about heredity (...) (gene-centric views), holism about heredity (Developmental Systems Theory), and those stressing the role of multiple systems of inheritance. Second, between those analyzing inheritance solely in terms of replication and transmission, and views that stress the multi-generation reproduction of phenotypic traits. A third debate is concerned with different criteria that have been proposed for identifying and delimiting inheritance systems. A fourth controversy revolves around the significance of the “Lamarckian” aspects of some of the inheritance systems that have been identified, such as epigenetic inheritance and behavioral inheritance, that allow the transmission of environmentally induced characters (i.e., “soft inheritance”). (shrink)

Recent theories in cognitive science have begun to focus on the active role of organisms in shaping their own environment, and the role of these environmental resources for cognition. Approaches such as situated, embedded, ecological, distributed and particularly extended cognition look beyond ‘what is inside your head’ to the old Gibsonian question of ‘what your head is inside of’ and with which it forms a wider whole—its internal and external cognitive niche. Since these views have been treated as a radical (...) departure from the received view of cognition, their proponents have looked for support to similar extended views within (the philosophy of) biology, most notably the theory of niche construction. This paper argues that there is an even closer and more fruitful parallel with developmental systems theory and developmental niche construction. These ask not ‘what is inside the genes you inherited’, but ‘what the inherited genes are inside of’ and with which they form a wider whole—their internal and external ontogenetic niche, understood as the set of epigenetic, social, ecological, epistemic and symbolic legacies inherited by the organism as necessary developmental resources. To the cognizing agent, the epistemic niche presents itself not just as a partially self-engineered selective niche, as the niche construction paradigm will have it, but even more so as a partially self-engineered ontogenetic niche, a problem-solving resource and scaffold for individual development and learning. This move should be beneficial for coming to grips with our own (including cognitive) nature: what is most distinctive about humans is their developmentally plastic brains immersed into a well-engineered, cumulatively constructed cognitive–developmental niche. (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)

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 into gene and non-gene and the vestiges of the ‘black-boxing’ of developmental processes in the modern synthesis, such as the asymmetric use of the concept of information. Phenomena that are marginalized in current gene-centric conceptions, such as extra-genetic inheritance, niche construction and phenotypic plasticity are placed center stage. (shrink)

What are the effects of our environment on human development and the next generation? Numerous studies have provided ample evidence that a healthy environment and lifestyle of the mother is important for her offspring. Biological mechanisms underlying these environmental influences have been proposed to involve alterations in the epigenome. Is there enough evidence to suggest a similar contribution from the part of the father? Animal models provide proof of a transgenerational epigenetic effect through the paternal germ line, but (...) can this be translated to humans? To date, literature on fathers is scarce. Human studies do not always incorporate appropriate tools to evaluate paternal influences or epigenetic effects. In reviewing the literature, I stress the need to explore and recognize paternal contributions to offspring's health within the Developmental Origins of Health and Disease hypothesis, and coin this new concept the Paternal Origins of Health and Disease paradigm. A better understanding of preconceptional origins of disease through the totality of paternal exposures, or the paternal exposome, will provide evidence-based public health recommendations for future fathers. A father's hereditary contribution exceeds that of his DNA sequence. The environment can regulate systems such as the sperm epigenome, which is inheritable as well and may steer the genome and health status of his offspring. We here suggest a new concept, the Paternal Origins of Health and Disease, as part of the DOHaD theory. (shrink)

Developmental Systems Theory (DST) emphasises the importance of non-genetic factors in development and their relevance to evolution. A common, deflationary reaction is that it has long been appreciated that non-genetic factors are causally indispensable. This paper argues that DST can be reformulated to make a more substantive claim: that the special role played by genes is also played by some (but not all) non-genetic resources. That special role is to transmit inherited representations, in the sense of Shea (...) (2007: Biology and Philosophy, 22, 313-331). Formulating DST as the claim that there are non-genetic inherited representations turns it into a striking, empirically-testable hypothesis, driving the sort of investigations that are only now beginning to appear in the scientific literature. DST’s characteristic rejection of a gene vs. environment dichotomy is preserved, but without dissolving all potentially explanatory distinctions into an interactionist causal soup, as some have alleged. (shrink)

Parental effects are a major source of phenotypic plasticity and may influence offspring phenotype in concert with environmental demands. Studies of “environmental epigenetics” suggest that (1) DNA methylation states are variable and that both demethylation and remethylation occur in post‐mitotic cells, and (2) that remodeling of DNA methylation can occur in response to environmentally driven intracellular signaling pathways. Studies of mother‐offspring interactions in rodents suggest that parental signals influence the DNA methylation, leading to stable changes in (...) gene expression. If parental effects do indeed enhance the “match” between prevailing environmental demands and offspring phenotype, then the potential for variation in environmental conditions over time would suggest a mechanism that requires active maintenance across generations through parental signaling. We suggest that parental regulation of DNA methylation states is thus an ideal candidate mechanism for parental effects on phenotypic variation. (shrink)

The dichotomy between Nature and Nurture, which has been dismantled within the framework of development, remains embodied in the notions of plasticity and evolvability. We argue that plasticity and evolvability, like development and heredity, are neither dichotomous nor distinct: the very same mechanisms may be involved in both, and the research perspective chosen depends to a large extent on the type of problem being explored and the kinds of questions being asked. Epigenetic inheritance leads to transgenerationally (...) extended plasticity, and developmentally-induced heritable epigenetic variations provide additional foci for selection that can lead to evolutionary change. Moreover, hereditary innovations may result from developmentally induced large-scale genomic repatterning events, which are akin to Goldschmidtian “systemic mutations”. The epigenetic mechanisms involved in repatterning can be activated by both environmental and genomic stress, and lead to phylogenetic as well as ontogenetic changes. Hence, the effects and the mechanisms of plasticity directly contribute to evolvability. (shrink)

The article proposes to further develop the ideas of the Extended Evolutionary Synthesis by including into evolutionary research an analysis of phenomena that occur above the organismal level. We demonstrate that the current Extended Synthesis is focused more on individual traits (genetically or non-genetically inherited) and less on community system traits (synergetic/organizational traits) that characterize transgenerational biological, ecological, social, and cultural systems. In this regard, we will consider various communities that are made up of interacting populations, and for which (...) the individual members can belong to the same or to different species. Examples of communities include biofilms, ant colonies, symbiotic associations resulting in holobiont formation, and human societies. The proposed model of evolution at the level of communities revises classic theorizing on the major transitions in evolution by analyzing the interplay between community/social traits and individual traits, and how this brings forth ideas of top-down regulations of bottom-up evolutionary processes (collaboration of downward and upward causation). The work demonstrates that such interplay also includes reticulate interactions and reticulate causation. In this regard, we exemplify how community systems provide various non-genetic ‘scaffoldings’, ‘constraints’, and ‘affordances’ for individual and sociocultural evolutionary development. Such research complements prevailing models that focus on the vertical transmission of heritable information, from parent to offspring, with research that instead focusses on horizontal, oblique and even reverse information transmission, going from offspring to parent. We call this reversed information transfer the ‘offspring effect’ to contrast it from the ‘parental effect’. We argue that the proposed approach to inheritance is effective for modelling cumulative and distributed developmental process and for explaining the biological origins and evolution of language. (shrink)

Literature on maternal exposures and the risk of epigenetic changes or diseases in the offspring is growing. Paternal contributions are often not considered. However, some animal and epidemiologic studies on various contaminants, nutrition, and lifestyle‐related conditions suggest a paternal influence on the offspring's future health. The phenotypic outcomes may have been attributed to DNA damage or mutations, but increasing evidence shows that the inheritance of environmentally induced functional changes of the genome, and related disorders, are (also) driven by (...) epigenetic components. In this essay we suggest the existence of epigenetic windows of susceptibility to environmental insults during sperm development. Changes in DNA methylation, histone modification, and non‐coding RNAs are viable mechanistic candidates for a non‐genetic transfer of paternal environmental information, from maturing germ cell to zygote. Inclusion of paternal factors in future research will ultimately improve the understanding of transgenerational epigenetic plasticity and health‐related effects in future generations. (shrink)

That there is a heritable or familial component of susceptibility to chronic non‐communicable diseases such as type 2 diabetes, obesity and cardiovascular disease is well established, but there is increasing evidence that some elements of such heritability are transmitted non‐genomically and that the processes whereby environmental influences act during early development to shape disease risk in later life can have effects beyond a single generation. Such heritability may operate through epigenetic mechanisms involving regulation of either imprinted or non‐imprinted (...) genes but also through broader mechanisms related to parental physiology or behaviour. We review evidence and potential mechanisms for non‐genomic transgenerational inheritance of ‘lifestyle’ disease and propose that the ‘developmental origins of disease’ phenomenon is a maladaptive consequence of an ancestral mechanism of developmental plasticity that may have had adaptive value in the evolution of generalist species such as Homo sapiens. BioEssays 29: 145–154, 2007. © 2007 Wiley Periodicals, Inc. (shrink)

The themes, problems and challenges of developmental systems theory as described in Cycles of Contingency are discussed. We argue in favor of a robust approach to philosophical and scientific problems of extended heredity and the integration of behavior, development, inheritance, and evolution. Problems with Sterelny's proposal to evaluate inheritance systems using his `Hoyle criteria' are discussed and critically evaluated. Additional support for a developmental systems perspective is sought in evolutionary studies of performance (...) and behavior modulation of fitness. (shrink)

Since the 1990s, the terms “Lamarckism” and “Lamarckian” have seen a significant resurgence in biological publications. The discovery of new molecular mechanisms have been interpreted as evidence supporting the reality and efficiency of the inheritance of acquired characters, and thus the revival of Lamarckism. The present paper aims at giving a critical evaluation of such interpretations. I argue that two types of arguments allow to draw a clear distinction between the genuine Lamarckian concept of inheritance of (...) acquired characters and transgenerational epigenetic inheritance. The first concerns the explanandum of the processes under consideration: molecular mechanisms of transgenerational epigenetic inheritance are understood as evolved products of natural selection. This means that the kind of inheritance of acquired characters they might be responsible for is an obligatory emergent feature of evolution, whereas traditional Lamarckisms conceived the inheritance of acquired characters as a property inherent in living matter itself. The second argument concerns the explanans of the inheritance of acquired characters: in light of current knowledge, epigenetic mechanisms are not able to drive adaptive evolution by themselves. Emergent Lamarckian phenomena would be possible if and only if individual epigenetic variation allowed the inheritance of acquired characters to be a factor of unlimited change. This implies specific requirements for epigenetic variation, which I explicitly define and expand upon. I then show that given current knowledge, these requirements are not empirically grounded. (shrink)

Critically significant parental effects in behavioral genetics may be partly understood as a consequence of maternal brain structure and function of caregiving systems recently studied in humans as well as rodents. Key parental brain areas regulate emotions, motivation/reward, and decision making, as well as more complex social-cognitive circuits. Additional key environmental factors must include socioeconomic status and paternal brain physiology. These have implications for developmental and evolutionary biology as well as public policy.

Many scientists and philosophers of science are troubled by the relative isolation of developmental from evolutionary biology. Reconciling the science of development with the science of heredity preoccupied a minority of biologists for much of the twentieth century, but these efforts were not corporately successful. Mainly in the past fifteen years, however, these previously dispersed integrating programmes have been themselves synthesized and so reinvigorated. Two of these more recent synthesizing endeavours are evolutionary developmental biology and developmental (...) class='Hi'>systems theory. While the former is a bourgeoning and scientifically well-respected biological discipline, the same cannot be said of DST, which is virtually unknown among biologists. In this review, we provide overviews of DST and EDB, summarize their key tenets, examine how they relate to one another and to the study of epigenetics, and survey the impact that DST and EDB have had on biological theory and practice. (shrink)

In the visual representation of ontologies, in particular of part-whole relationships, it is customary to use graph theory as the representational background. We claim here that the standard graph-based approach has a number of limitations, and we propose instead a new representation of part-whole structures for ontologies, and describe the results of experiments designed to show the effectiveness of this new proposal especially as concerns reduction of visual complexity. The proposal is developed to serve visualization of ontologies conformant to (...) the Basic Formal Ontology. But it can be used also for more general applications, particularly in the biomedical domain. (shrink)

'...a challenging and useful book, both because it provokes a careful scrutiny of one's own basic ideas regarding evolutionary theory, and because it cuts across so many biological disciplines.' -The Quarterly Review of Biology 'In my view, this work exemplifies Theoretical Biology at its best...here is rampant speculation that is consistently based on cautious reasoning from the available data. Even more refreshing is the absence of sloganeering, grandstanding, and 'isms'.' -Biology and Philosophy 'Epigenetics is fundamental to understanding both development (...) and gene expression, and not surprisingly, evolutionary biologists have long been fascinated with its proper place in evolutionary theory...Enter Jablonka and Lamb, who provide a thoughtful review of the recent molecular literature and suggest a number of potential consequences.' -EvolutionSince first publication of this controversial book, much of the initial opposition to the ideas it contained has been replaced by a general, although often grudging, acceptance of them. Advances in knowledge, especially at the molecular level, have enhanced general awareness and interest in epigenetics and the evolution of systems that store and transmit information and put any of the authors' speculations on a more solid basis. This paperback edition contains a new Preface that sets out the major changes in the scientific world and in the authors' own thinking that have occurred since the book was published. A new Appendix provides a selected bibliography of the many books and articles about epigenetic inheritance and its role in evolution that have appeared since first publication. (shrink)

In reworking a variety of biological concepts, Developmental Systems Theory (DST) has made frequent use of parity of reasoning. We have done this to show, for instance, that factors that have similar sorts of impact on a developing organism tend nevertheless to be invested with quite different causal importance. We have made similar arguments about evolutionary processes. Together, these analyses have allowed DST not only to cut through some age-old muddles about the nature of development, but also (...) to effect a long-delayed reintegration of development into evolutionary theory. Our penchant for causal symmetry, however (or 'causal democracy', as it has recently been termed), has sometimes been misunderstood. This paper shows that causal symmetry is neither a platitude about multiple influences nor a denial of useful distinctions, but a powerful way of exposing hidden assumptions and opening up traditional formulations to fruitful change. (shrink)

Using the evolution of the stickleback family of subarctic fish as a touchstone, we explore the effect of new discoveries about regulatory genetics, developmental plasticity, and epigenetic inheritance on the conceptual foundations of the Modern Evolutionary Synthesis. Identifying the creativity of natural selection as the hallmark of the Modern Synthesis, we show that since its inception its adherents have pursued a variety of research projects that at first seemed to conflict with its principles, but were accommodated. (...) We situate challenges coming from developmental biology in a dialectic between innovation and tradition, suggesting on the basis of past episodes that even if developmental plasticity and epigenetic inheritance are aligned with its principles the Modern Synthesis will be significantly affected. (shrink)

I argue that too much attention has been paid to the Baldwin effect. George Gaylord Simpson was probably right when he said that the effect is theoretically possible and may have actually occurred but that this has no major implications for evolutionary theory. The Baldwin effect is not even central to Baldwin's own account of social heredity and biology-culture co-evolution, an account that in important respects resembles the modern ideas of epigenetic inheritance and niche-construction.

It is now widely accepted that a scientifically credible conception of human nature must reject the folkbiological idea of a fixed, inner essence that makes us human. We argue here that to understand human nature is to understand the plastic process of human development and the diversity it produces. Drawing on the framework of developmental systems theory and the idea of developmental niche construction we argue that human nature is not embodied in only one input to (...) development, such as the genome, and that it should not be confined to universal or typical human characteristics. Both similarities and certain classes of differences are explained by a human developmental system that reaches well out into the 'environment'. We point to a significant overlap between our account and the ‘Life History Trait Cluster’ account of Grant Ramsey, and defend the developmental systems account against the accusation that trying to encompass developmental plasticity and human diversity leads to an unmanageably complex account of human nature. (shrink)

What role does non-genetic inheritance play in evolution? In recent work we have independently and collectively argued that the existence and scope of non-genetic inheritance systems, including epigenetic inheritance, niche construction/ecological inheritance, and cultural inheritance—alongside certain other theory revisions—necessitates an extension to the neo-Darwinian Modern Synthesis (MS) in the form of an Extended Evolutionary Synthesis (EES). However, this argument has been challenged on the grounds that non-genetic inheritance systems are exclusively (...) proximate mechanisms that serve the ultimate function of calibrating organisms to stochastic environments. In this paper we defend our claims, pointing out that critics of the EES (1) conflate non-genetic inheritance with early 20th-century notions of soft inheritance; (2) misunderstand the nature of the EES in relation to the MS; (3) confuse individual phenotypic plasticity with trans-generational non-genetic inheritance; (4) fail to address the extensive theoretical and empirical literature which shows that non-genetic inheritance can generate novel targets for selection, create new genetic equilibria that would not exist in the absence of non-genetic inheritance, and generate phenotypic variation that is independent of genetic variation; (5) artificially limit ultimate explanations for traits to gene-based selection, which is unsatisfactory for phenotypic traits that originate and spread via non-genetic inheritance systems; and (6) fail to provide an explanation for biological organization. We conclude by noting ways in which we feel that an overly gene-centric theory of evolution is hindering progress in biology and other sciences. (shrink)

Transformations of Lamarckism is an edited volume arising from a workshop to commemorate the bicentenary of the publication of Philosophie Zoologique. The contributed chapters discuss the history of Lamarckism, present new developments in biology that could be considered to vindicate Lamarck, and argue for a revision, if not a revolution, in evolutionary theory. My review argues that twentieth and twenty-first century conceptions of Lamarckism can be considered a reaction to August Weismann’s uncompromising rejection of the (...) inheritance of acquired characters in the late nineteenth century. Weismann rejected the inheritance of acquired characters both as a proximate mechanism of heredity and as an ultimate cause of adaptation. I argue that Weismann’s proximate claim is still valid for the kind of mechanism that he had in mind but that the inheritance of acquired characters has come to refer to many different processes, some of which undoubtedly do occur. However, processes of physiological adaptation and adaptive plasticity, even if transgenerational, do not challenge Weismann’s claim about the ultimate causes of adaptation because these processes can be understood as evolving by natural selection. Finally, I discuss some of the emotional and aesthetic reasons why many find Lamarckism an attractive alternative to hard-core neo-Darwinism. (shrink)

Recent epidemiological reports of associations between socioeconomic status and epigenetic markers that predict vulnerability to diseases are bringing to light substantial biological effects of social inequalities. Here, we start the discussion of the moral consequences of these findings. We firstly highlight their explanatory importance in the context of the research program on the Developmental Origins of Health and Disease (DOHaD) and the social determinants of health. In the second section, we review some theories of the moral status (...) of health inequalities. Rather than a complete outline of the debate, we single out those theories that rest on the principle of equality of opportunity and analyze the consequences of DOHaD and epigenetics for these particular conceptions of justice. We argue that DOHaD and epigenetics reshape the conceptual distinction between natural and acquired traits on which these theories rely and might provide important policy tools to tackle unjust distributions of health. (shrink)

The paper aims at proposing an extended notion of epigenesis acknowledging an actual causal import to the phenotypic dimension for the evolutionary diversification of life forms. “Introductory remarks” section offers introductory remarks on the issue of epigenesis contrasting it with ancient and modern preformationist views. In “Transmutation of forms: phenotypic variation, diversification, and complexification” section we propose to intend epigenesis as a process of phenotypic formation and diversification dependent on environmental influences, independent of changes in the genomic nucleotide sequence, and (...) occurring during the whole life span. Then, “Evolvability and phenotypic evolution” section focuses on phenotypic plasticity and offers an overview of basic properties that allows biological systems to be evolvable—i.e. to have the potentiality of producing phenotypic variation. Successively, the emphasis is put on environmentally-induced modifications in the regulation of gene expression giving rise to phenotypic variation and diversification. After some brief considerations on the debated issue of epigenetic inheritance, the issue of culture is considered. The key point is that, in the case of humans and of the evolutionary history of the genus Homo at least, the environment is also, importantly, the cultural environment. Thus, “Bio-cultural feedback” section argues that a bio-cultural feedback should be acknowledged in the “epigenic” processes leading to phenotypic diversification and innovation in Homo evolution. Finally, “Brain plasticity and cultural neural reuse” section introduces the notion of “cultural neural reuse”, which refers to phenotypic/neural modifications induced by specific features of the cultural environment that are effective in human cultural evolution without involving genetic changes. Therefore, cultural neural reuse may be regarded as a key instance of the bio-cultural feedback and ultimately of the extended notion of epigenesis proposed in this work. (shrink)

Parents influence the development of their offspring in many ways beyond the transmission of DNA. This includes transfer of epigenetic states, nutrients, antibodies and hormones, and behavioural interactions after birth. While the evolutionary consequences of such nongenetic inheritance are increasingly well understood, less is known about how inheritance mechanisms evolve. Here, we present a simple but versatile model to explore the adaptive evolution of non-genetic inheritance. Our model is based on a switch mechanism that produces alternative (...) phenotypes in response to different inputs, including genes and non-genetic factors transmitted from parents and the environment experienced during development. This framework shows how genetic and non-genetic inheritance mechanisms and environmental conditions can act as cues by carrying correlational information about future selective conditions. Differential use of these cues is manifested as different degrees of genetic, parental or environmental morph determination. We use this framework to evaluate the conditions favouring non-genetic inheritance, as opposed to genetic determination of phenotype or within-generation plasticity, by applying it to two putative examples of adaptive non-genetic inheritance: maternal effects on seed germination in plants and transgenerational phase shift in desert locusts. Our simulation models show how the adaptive value of non-genetic inheritance depends on its mechanism, the pace of environmental change, and life history characteristics. (shrink)

Some central ideas associated with developmental systems theory are outlined for non-specialists. These ideas concern the nature of biological development, the alleged distinction between “genetic” and “environmental” traits, the relations between organism and environment, and evolutionary processes. I also discuss some criticisms of the DST approach.

Organisms live not as discrete entities on which an independent environment acts, but as members of a reproductive lineage in an ongoing series of interactions between that lineage and a dynamic ecological niche. These interactions continuously shape both systems in a reciprocal manner, resulting in the emergence of reliably co-occurring configurations within and between both systems. The enactive approach to cognition describes this relationship as the structural coupling between an organism and its environment; similarly, Developmental Systems (...) Theory emphasizes the reciprocal nature of structurally coupled systems in its analysis of organisms as developmental processes embedded within a developmental system. Through an enactive-developmental systems framing, this paper identifies the organizational features of cognizing systems in order to motivate a picture of how organism and environment co-determine and co-construct one another. I argue that organisms can be characterized as self-organizing, operationally closed, plastic systems ecologically embedded within a developmental system. In virtue of this organizational makeup, organisms actively engage in the modulation and assessment of their coupling with their environment: cognitive strategies that entail contextualized responses to variations across the web of interactions that comprises the developmental system. (shrink)

DNA methylation plays a key role in neural cell fate and provides a molecular link between early life stress and later-life behavioral phenotypes. Here, studies that combine neuroimaging methods and DNA methylation analysis in pediatric population with a history of adverse experiences were systematically reviewed focusing on: targeted genes and neural correlates; statistical models used to examine the link between DNA methylation and neuroimaging data also considering early life stress and behavioral outcomes. We identified 8 studies that report associations between (...) DNA methylation and brain structure/functions in infants, school age children and adolescents faced with early life stress condition. Results showed that several genes were investigated and different neuroimaging techniques were performed. Statistical model used ranged from correlational to more complex moderated mediation models. Most of the studies considered DNA methylation and neural correlates as mediators in the relationship between early life stress and behavioral phenotypes. Understanding what role DNA methylation and neural correlates play in interaction with early life stress and behavioral outcomes is crucial to promote theory-driven studies as the future direction of this research fields. (shrink)

We hypothesize that under the predicted scenario of climate change epigenetically mediated environmental sex determination could become an epigenetic trap. Epigenetically regulated environmental sex determination is a mechanism by which species can modulate their breeding strategies to accommodate environmental change. Growing evidence suggests that epigenetic mechanisms may play a key role in phenotypic plasticity and in the rapid adaptation of species to environmental change, through the capacity of organisms to maintain a non‐genetic plastic memory of the environmental (...) and ecological conditions experienced by their parents. However, inherited epigenetic variation could also be maladaptive, becoming an epigenetic trap. This is because environmental sex determination can alter sex ratios by increasing the survival of one of the sexes at the expense of negative fitness consequences for the other, which could lead not only to the collapse of natural populations, but also have an impact in farmed animal and plant species. (shrink)

Ideas about heredity and evolution are undergoing a revolutionary change. New findings in molecular biology challenge the gene-centered version of Darwinian theory according to which adaptation occurs only through natural selection of chance DNA variations. In Evolution in Four Dimensions, Eva Jablonka and Marion Lamb argue that there is more to heredity than genes. They trace four "dimensions" in evolution -- four inheritance systems that play a role in evolution: genetic, epigenetic, behavioral, and symbolic. These (...) class='Hi'>systems, they argue, can all provide variations on which natural selection can act. Evolution in Four Dimensions offers a richer, more complex view of evolution than the gene-based, one-dimensional view held by many today. The new synthesis advanced by Jablonka and Lamb makes clear that induced and acquired changes also play a role in evolution.After discussing each of the four inheritance systems in detail, Jablonka and Lamb "put Humpty Dumpty together again" by showing how all of these systems interact. They consider how each may have originated and guided evolutionary history and they discuss the social and philosophical implications of the four-dimensional view of evolution. Each chapter ends with a dialogue in which the authors engage the contrarieties of the fictional "I.M.," or Ifcha Mistabra -- Aramaic for "the opposite conjecture" -- refining their arguments against I.M.'s vigorous counterarguments. The lucid and accessible text is accompanied by artist-physician Anna Zeligowski's lively drawings, which humorously and effectively illustrate the authors' points. (shrink)

The acronym Developmental systems theory (DST) has been introduced into the literature on development in at least three different contexts in recent years – twice for DST, and before that, for Dynamical Systems Theory – and in all cases, to designate a new perspective for understanding development. Subtle but significant differences in argument and aims distinguish these uses, and confound the difficulty of saying just what DST is. My aim in this paper is to disambiguate (...) these different terms – both to call attention to the difference of perspectives, and to carve out a conceptual space for the concrete issues at stake. (shrink)

Our paper aims at bringing to the fore the crucial role that habits play in Charles Darwin’s theory of evolution by means of natural selection. We have organized the paper in two steps: first, we analyse value and functions of the concept of habit in Darwin's early works, notably in his Notebooks, and compare these views to his mature understanding of the concept in the Origin of Species and later works; second, we discuss Darwin’s ideas on habits in the (...) light of today’s theories of epigenetic inheritance, which describe the way in which the functioning and expression of genes is modified by the environment, and how these modifications are transmitted over generations. We argue that Darwin’s lasting and multifaceted interest in the notion of habit, throughout his intellectual life, is both conceptually and methodologically relevant. From a conceptual point of view, intriguing similarities can be found between Darwin’s (early) conception of habit and contemporary views on epigenetic inheritance. From a methodological point of view, we suggest that Darwin’s plastic approach to habits, from his early writings up to the mature works, can provide today’s evolutionary scientists with a viable methodological model to address the challenging task of extending and expanding evolutionary theory, with particular reference to the integration of epigenetic mechanisms into existing models of evolutionary change. Over his entire life Darwin has modified and reassessed his views on habits as many times as required by evidence: his work on this notion may represent the paradigm of a habit of good scientific research methodology. (shrink)

This essay attempts to describe the neo-Lamarckian atmosphere that was dominant in French biology for more than a century. Firstly, we demonstrate that there were not one but at least two French neo-Lamarckian traditions. This implies, therefore, that it is possible to propose a clear definition of a (neo) Lamarckian conception, and by using it, to distinguish these two traditions. We will see that these two conceptions were not dominant at the same time. The first French neo-Lamarckism (1879-1931) was (...) structured by a very mechanic view of natural processes. The main representatives of this first period were scientists such as Alfred Giard (1846-1908), Gaston Bonnier (1853-1922) and Félix Le Dantec (1869-1917). The second Lamarckism - much more vitalist in its inspiration - started to develop under the supervision of people such as Albert Vandel (1894-1980) and Pierre-Paul Grassé (1895-1985). Secondly, this essay suggests that the philosophical inclinations of these neo-Lamarckisms reactivated a very ancient and strong dichotomy of French thought. One part of this dichotomy is a material, physicalist tradition, which started with René Descartes but developed extensively during the 18th and 19th centuries. The other is a spiritual and vitalist reaction to the first one, which also had a very long history, though it is most closely associated with the work of Henri Bergson. Through Claude Bernard, the first neo-Lamarckians tried to construct a mechanical and determinist form of evolutionary theory which was, in effect, a Cartesian theory. The second wave of neo-Lamarckians wanted to reconsider the autonomy and reactivity of life forms, in contrast to purely physical systems. (shrink)

Genetic interventions, which include transgenic engineering, gene editing, and other forms of genome modification aimed at altering the information “in” the genetic code, are rapidly increasing in power and scale. Biosemiotics offers unique tools for understanding the nature, risks, scope, and prospects of such technologies, though few in the community have turned their attention specifically in this direction. Bruni is an important exception. In this paper, I examine how we frame the concept of “side effects” that result from genetic (...) interventions and how the concept stands up to current perspectives of the role of organism activity in development. I propose that once the role of living systems in constructing and modifying the informational value of their various developmental resources is taken into account, the concept of a “side effect” will need to be significantly revised. Far from merely a disturbance brought about in a senseless albeit complex system, a biosemiotic view would take “side effects” as at least sometimes the organism’s active re-organization in order to accommodate or make use of novelty. This insight is nascent in the work of developmental plasticity and niche construction theory, but it is brought into sharper focus by the explicitly interpretive perspective offered by biosemiotics. Understanding the “side effects” of genetic interventions depends in part on being able to articulate when and where unexpected consequences are a result of semiotic activity at various levels within the system. While a semiotic interpretation of “side effects” puts into question the naive attitude that would see all unintended side effects as indications of disturbance in system functionality, it certainly does not imply that such side effects are of no concern for the viability of the organisms in the system. As we shall see, the fact that such interventions do not respect the translation of information that occurs in multi-level biological systems ensures that disruption is still likely. But it does unprivilege the human agent as the sole generator of meaning and information in the products of biotechnology, with important consequences on how we understand our relationship with other species. (shrink)

The purpose of this article is to examine stakeholder identification and prioritization by managers using the power, legitimacy, and urgency framework of Mitchell et al. (Academy of Management Review 22, 853–886; 1997). We use a multi-method, comparative case study of two large-scale sporting event organizing committees, with a particular focus on interviews with managers at three hierarchical levels. We support the positive relationship between number of stakeholder attributes and perceived stakeholder salience. Managers’ hierarchical level and role have direct and moderating (...) effects on stakeholder identification and perceived salience. We also found that most stakeholders were definitive, dominant, or dormant types – the other five types were rare. Power has the most important effect on salience, followed by urgency and legitimacy. Based on our case study, we offer several ways to advance the theory of stakeholder identification and salience. (shrink)

In their considered reviews both Thomas Pradeu and Lindell Bromham introduce important topics not sufficiently covered in our book. Pradeu asks us to enlarge on the epigenetic and ecological context of genes, particularly in the form of symbioses. We use the relationship between eukaryotes and their symbiotic organisms as a welcome opportunity to clarify our concept of the developmental niche, and its relationship to the developmental system. Bromham’s comments reveal that she is primarily interested in identifying macroevolutionary (...) patterns. From her vantage point eco-evo-devo, the study of phenotypic plasticity, epigenetic and exogenetic inheritance, have not yet demonstrated the need for any revolutionary change in evolutionary thought. For us they highlight the extent to which proximate developmental mechanisms can inform ultimate biology. (shrink)

This essay attempts to describe the neo-Lamarckian atmosphere that was dominant in French biology for more than a century. Firstly, we demonstrate that there were not one but at least two French neo-Lamarckian traditions. This implies, therefore, that it is possible to propose a clear definition of a (neo)Lamarckian conception, and by using it, to distinguish these two traditions. We will see that these two conceptions were not dominant at the same time. The first French neo-Lamarckism (1879–1931) was structured (...) by a very mechanic view of natural processes. The main representatives of this first period were scientists such as Alfred Giard (1846–1908), Gaston Bonnier (1853–1922) and Félix Le Dantec (1869–1917). The second Lamarckism – much more vitalist in its inspiration – started to develop under the supervision of people such as Albert Vandel (1894–1980) and Pierre-Paul Grassé (1895–1985). Secondly, this essay suggests that the philosophical inclinations of these neo-Lamarckisms reactivated a very ancient and strong dichotomy of French thought. One part of this dichotomy is a material, physicalist tradition, which started with René Descartes but developed extensively during the 18th and 19th centuries. The other is a spiritual and vitalist reaction to the first one, which also had a very long history, though it is most closely associated with the work of Henri Bergson. Through Claude Bernard, the first neo-Lamarckians tried to construct a mechanical and determinist form of evolutionary theory which was, in effect, a Cartesian theory. The second wave of neo-Lamarckians wanted to reconsider the autonomy and reactivity of life forms, in contrast to purely physical systems. (shrink)

Griffiths and Russell D. Gray (1994, 1997, 2001) have argued that the fundamental unit of analysis in developmental systems theory should be a process – the life cycle – and not a set of developmental resources and interactions between those resources. The key concepts of developmental systems theory, epigenesis and developmental dynamics, both also suggest a process view of the units of development. This chapter explores in more depth the features of (...) class='Hi'>developmental systems theory that favour treating processes as fundamental in biology and examines the continuity between developmental systems theory and ideas about process in the work of several major figures in early 20th century biology, most notable C.H Waddington. (shrink)

Cognitive developmental evidence is sometimes conscripted to support ''naturalized epistemology'' arguments to the effect that a general epistemic stance leads children to build theory-like accounts of underlying properties of kinds. A review of the evidence suggests that what prompts conceptual acquisition is not a general epistemic stance but a series of category-specific intuitive principles that constitute an evolved ''natural metaphysics''. This consists in a system of categories and category-specific inferential processes founded on definite biases in prototype formation. Evidence (...) for this system provides a better understanding of the limited ''plasticity'' of ontological commitments as well as a computationally plausible account of their initial state, avoiding ambiguities about innateness. This may provide a starting point for a ''naturalized epistemology'' that takes into account evolved properties of human conceptual structures. (shrink)

Cognitive developmental evidence is sometimes conscripted to support ''naturalized epistemology'' arguments to the effect that a general epistemic stance leads children to build theory-like accounts of underlying properties of kinds. A review of the evidence suggests that what prompts conceptual acquisition is not a general epistemic stance but a series of category-specific intuitive principles that constitute an evolved ''natural metaphysics''. This consists in a system of categories and category-specific inferential processes founded on definite biases in prototype formation. Evidence (...) for this system provides a better understanding of the limited ''plasticity'' of ontological commitments as well as a computationally plausible account of their initial state, avoiding ambiguities about innateness. This may provide a starting point for a ''naturalized epistemology'' that takes into account evolved properties of human conceptual structures. (shrink)

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 deep interest. Waddington was, however, predominantly a thinker, and set out to provide a coherent framework for understanding the genetic bases of embryogenesis and evolution, developing his ideas in many books. Perhaps his best-known concept is the epigenetic landscape: here a ball rolls down a complex valley, making path choices. The rolling ball represents a cell’s development over time, while the topography represents the changing regulatory environment that controls these choices. In its later forms, the role of each feature in the landscape was controlled by the effects of sets of interacting genes, an idea underpinning contemporary approaches to systems biology. Waddington was the first developmental geneticist and probably the most important developmental biologist of the pre-molecular age. (shrink)

The scarcity of human organs requires the transplant community to make difficult allocation decisions. This process begins at individual medical centers, where transplant teams decide which patients to place on the transplant waiting list. Each transplant center utilizes its own listing criteria to determine if a patient is eligible for transplantation. These criteria have historically considered preexisting affective and psychotic disorders to be relative or absolute contraindications to transplantation. While attitudes within the field appear to be moving away from this (...) practice, there is no data to confirm that eligibility criteria have changed. There are no nationwide guidelines detailing the manner in which affective and psychotic disorders should impact transplant eligibility. Individual transplant centers thus form their own transplant eligibility criteria, resulting in significant inter-institution variability. Data from the 1990s indicates that the majority of transplant programs considered certain psychiatric illnesses, such as active schizophrenia, to be absolute contraindications to transplantation. A review of literature reveals that no comprehensive data has been collected on the topic since that time. Furthermore, the limited data available about current practices suggests that psychiatric illness continues to be viewed as a contraindication to transplantation at some transplant centers. In light of this finding, we review psychiatric literature that examines the impact of affective and psychotic disorders on transplant outcomes and conclude that the presence of these disorders is not an accurate predictor of transplant success. We then discuss the requirements of justice as they relate to the creation of a just organ allocation system. We conclude that transplant eligibility criteria that exclude patients with affective and psychotic disorders on the basis of their psychiatric diagnosis alone are unjust. Just listing criteria must incorporate only those factors that have a causative effect on posttransplant morbidity and mortality. Justice also demands that we eliminate current inter-institution practice variations in favor of national transplant eligibility criteria. Given the limited data available about current practices, we call for an updated study investigating the manner in which affect and psychotic disorders impact transplant eligibility determinations. (shrink)

In the years of the post-Darwinian debate, many American naturalists invoked the name of Lamarck to signal their belief in a purposive and anti-Darwinian view of evolution. Yet Weismann’s theory of germ-plasm continuity undermined the shared tenet of the neo-Lamarckian theories as well as the idea of the interchangeability between biological and social heredity. Edward Drinker Cope, the leader of the so-called “American School,” defended his neo-Lamarckian philosophy against every attempt to redefine the relationship between behavior, development, and (...) heredity beyond the epigenetic model of inheritance. This paper explores Cope’s late-career defense of neo-Lamarckism. Particular attention is dedicated to the debate he had with James Mark Baldwin before the publication of Baldwin’s own “A New Factor in Evolution”. I argue that Cope’s criticism was partly due to the fact that Baldwin’s theory of social heredity threatened Cope’s biologistic stance, as well as his attempt to preserve design in nature. This theoretical attitude had a remarkable impact on Baldwin’s arguments for the theory of organic selection. (shrink)

This exploratory enquiry seeks to examine the largely unexplored potential of semiotics for intercultural training and education. The proposed three-partdiscussion describes the process by which semiotic theoretical principles were selected and progressively refined into an applied model which was then pilotedthrough a 2007 research initiative entitled Tools for Cultural Development. The case study involved six groups of French and Australian trainees from both theacademic and professional sectors, in collaboration with university, government and community partners. The first part of the article (...) summarizes a review of theliterature on approaches to cultural competence training. The study then outlines the transcoding process by which the stated objectives of intercultural educationwere reformulated in semiotic terms, particularly in reference to cultural semiotics on which the theoretical core of the applied model was subsequently based.Relevant principles from other semiotic schools as well as similar theoretical and methodological stances in the social sciences reinforced the established body of theory for the training design. The third part of the study discusses the process by which semiotic principles were further defined as skill-based outcomes and goals for workshop implementation. This pragmatic defining process facilitated development of questionnaires and surveys, thereby allowing participants to evaluate the training experience by examining their perceptions about the workshop outcomes at the beginning and end of the sessions. This article presents the quantitative results of the evaluation and, in discussing the gains and limits of data obtained, provides the context for a follow-up article on the qualitative findingsof the study. (shrink)