We may now ask the question: In what historical perspective should we place the work of Richard Goldschmidt? There is no doubt that in the period 1910–1950 Goldschmidt was an important and prolific figure in the history of biology in general, and of genetics in particular. His textbook on physiological genetics, published in 1938, was an amazing compendium of ideas put forward in the previous half-century about how genes influence physiology and development. His earlier studies on the (...) genetic and geographic distribution of the various species and races of Lymantria contributed soundly to an understanding of the inheritance and development of sex, as well as the genetics of speciation. He was a curious mixture of the experimentalist, empiricist, and theorizer, who could grasp both the large and the small at once and wrap them into the same integrated and coherent package.In insisting on trying to relate genetics to development, Goldschmidt was continually forced to speculate and to forsake experimental fact for generalized theory. In refusing to reconcile a physiological and developmental conception with a corpuscular theory of the gene, he forced himself to substitute for a large body of experimentally verified evidence, vague and general speculations which did not lead in any precise direction. L. C. Dunn summarizes succinctly the effect of Goldschmidt's thinking on the development of genetic theory: Although Goldschmidt's ideas had a considerable influence in directing attention to the developmental processes intervening between genes and characters, they did not lead to the establishment of a general theory of development in genetical terms. In fact, at the end of the period, as signalized by Goldschmidt's book of 1938 [Physiological Gentics] doubt remained whether there was a field with a defined problem which could be identified as developmental genetics.Yet the frequency with which Goldschmidt is cited by later writers suggests that there is a more positive side to his contribution than this. The early Mendelians had recognized the genic constitution of organisms and the Drosophila workers had uncovered in detail the chromosomal mechanism of heredity. These great discoveries were of classical character, solidly based on a multitude of well established facts. By themselves, however, they would not have been sufficient to place genetics in the central position within biology which it was to assume. Beyond the “static” analysis of gene transmission an insight was needed into the dynamic role of genes in cellular physiology, biochemistry, and development. Goldschmidt recognized this aspect and outlined in brilliant generalizations a physiological theory of genetics. Necessarily the prophetic nature of this achievement — romantic in the sense of Ostwald's classification of great scientists and their discoveries — transcended its factual basis. It was the audacity of the theoretician unimpeded by the still scanty data which gave a new focus to biological thought.Some have called Goldschmidt an “obstructionist” in the history of genetics—one whose efforts were largely directed toward useless and continual criticism — challenge for the sake of challenge. As this line of argument goes, time wasted answering Goldschmidt's poorly conceived criticisms could have been better used to pursue new lines of thought within the classical gene theory. Sometimes historians are upbraided for studying the work of such “obstructionists,” or even the work of those whose ideas have later proved to be inadequate. This argument misses the point of history and the lessons which it can teach. Perhaps Goldschmidt did take the time of a number of the classical geneticists who tried to answer his objections to the gene theory; and perhaps many of his own ideas about genes were, by today's view, “wrong.” These are always easier judgments to make by hindsight than at the moment. But that is not the point. Goldschmidt had a considerable influence on his contemporaries, and to dismiss him as an obstructionist because his opponents later proved to be more nearly right, distorts history and obscures significant questions which we might well be asking. It might be valuable to know why Goldschmidt felt compelled to be an iconoclast —and how that impulse, psychological in orgin, led him to overlook critical items of evidence which his opponents considered more carefully. It might also be valuable to try and understand how a contemporary of Goldschmidt or. Morgan really viewed the gene theory — what were its strong and weak point? Such questions are not readily answered by studying the work of only those men who were correct by later standards.Answers to some of the above questions can be gleaned from this study of Goldschmidt — though the first, and most psychologically oriented, question is largely untouched here.On the one hand, Goldschmidt forced classical geneticists to reexamine the foundation of their ideas about the nature, inviolability, and even physical dimensions of genes. He also kept alive the very real problems of gene physiology and its relations to development — a relationship that was given less attention by the Drosophila school in their pursuit of the transmission process. On a broader level he emphasized to the biological community a principle that was in danger of being lost in the enthusiasm surrounding the expansion of the Mendelian-chromosome theory. The point was that all theories in science are transient, logical constructs which should not be held as sacred. Time and again, as the history of science has shown, theories become rigid structures which retard new modes of thought. From Goldschmidt's perspective, the gene theory was in danger of performing just that function, for it was preventing workers from viewing the gene in a functional as well as a structural light.On the other hand, examining the work of critics (or even of outright “obstructionists”) gives the historian an opportunity to observe how the more established community of scholars at some point in time reacts to challenges of its cherished ideas. The fact that many workers in his own day (such as Beadle, Luria, Delbruck, Sturtevant, Bridges, and others), as well as most geneticists today, saw Goldschmidt as having been largely “obstructionist” is useful historical (and/or sociological) data. Clearly, as this paper has tried to show, Goldschmidt's ideas were not all obstructionist, and his viewpoint that genes must be considered functionally was one which was clearly valid, premature as it may have been from an experimental and technique point of view. Perhaps Goldschmidt's major fault was that he developed alternative theories which were not easily testable. Nonetheless, his challenge to a fundamental and established idea brought forth a reaction from many members of the genetic community which indicates how unwilling they were to reconsider the formalized theory on which their work was based. While many of Goldschmidt's criticisms — and particularly his way of making them — arose out of his own idiosyncrasies, they were also a product of his times. His view of the nature of theory-construction, and his rejection of old-style reductionism and mechanism, were part of a growing awareness within the world scientific community that explanations based on reducing complex phenomena to ultimate particles or units were clearly inadequate. The success of the Mendelian-chromosome theory had obscured that fact for many geneticists, but to Goldschmidt it was a point that could not be allowed to pass unnoticed. (shrink)

Obesity is the focus of multiple lines of inquiry that have -- together and separately -- produced many deep insights into the physiology of weight gain and maintenance. We examine three such streams of research and show how they are oriented to obesity intervention through multilevel integrated approaches. The first research programme is concerned with the genetics and biochemistry of fat production, and it links metabolism, physiology, endocrinology and neurochemistry. The second account of obesity is developmental and draws (...) together epigenetic and environmental explanations that can be embedded in an evolutionary framework. The third line of research focuses on the role of gut microbes in the production of obesity, and how microbial activities interact with host genetics, development and metabolism. These interwoven explanatory strategies are driven by an orientation to intervention, both for experimental and therapeutic outcomes. We connect the integrative and intervention-oriented aspects of obesity research through a discussion of translation, broadening the concept to capture the dynamic, iterative processes of scientific practice and therapy development. This system-oriented analysis of obesity research expands the philosophical scrutiny of contemporary developments in the biosciences and biomedicine, and has the potential to enrich philosophy of science and medicine. (shrink)

The logic of genetic discovery has changed little over time, but the focus of biology is shifting from simple genotype–phenotype relationships to complex metabolic, physiological, developmental, and behavioral traits. In light of this, the traditional reductionist view of individual genes as privileged difference‐making causes of phenotypes is re‐examined. The scope and nature of genetic effects in complex regulatory systems, in which dynamics are driven by regulatory feedback and hierarchical interactions across levels of organization are considered. This review argues (...) that it is appropriate to treat genes as specific actual difference‐makers for the molecular regulation of gene expression. However, they are often neither stable, proportional, nor specific as causes of the overall dynamic behavior of regulatory networks. Dynamical models, properly formulated and validated, provide the tools to probe cause‐and‐effect relationships in complex biological systems, allowing to go beyond the limitations of genetic reductionism to gain an integrative understanding of the causal processes underlying complex phenotypes. (shrink)

Three early 20th-century attempts at unifying separate areas of biology, in particular development, genetics, physiology, and evolution, are compared in regard to their success and fruitfulness for further research: Jacques Loeb’s reductionist project of unifying approaches by physico-chemical explanations; Richard Goldschmidt’s anti-reductionist attempts to unify by integration; and Sewall Wright’s combination of reductionist research and vision of hierarchical genetic systems. Loeb’s program, demanding that all aspects of biology, including evolution, be studied by the methods of the experimental sciences, proved (...) highly successful and indispensible for higher level investigations, even though evolutionary change and properties of biological systems up to now cannot be fully explained on the molecular level alone. Goldschmidt has been appraised as pioneer of physiological and developmental genetics and of a new evolutionary synthesis which transcended neo-Darwinism. However, this study concludes that his anti-reductionist attempts to integrate genetics, development and evolution have to be regarded as failures or dead ends. His grand speculations were based on the one hand on concepts and experimental systems that were too vague in order to stimulate further research, and on the other on experiments which in their core parts turned out not to be reproducible. In contrast, Sewall Wright, apart from being one of the architects of the neo-Darwinian synthesis of the 1930s, opened up new paths of testable quantitative developmental genetic investigations. He placed his research within a framework of logical reasoning, which resulted in the farsighted speculation that examinations of biological systems should be related to the regulation of hierarchical genetic subsystems, possibly providing a mechanism for development and evolution. I argue that his suggestion of basing the study of systems on clearly defined properties of the components has proved superior to Goldschmidt’s approach of studying systems as a whole, and that attempts to integrate different fields at a too early stage may prove futile or worse. © 2011 Elsevier Inc. (shrink)

Model organisms became an indispensable part of experimental systems in molecular developmental and cell biology, constructed to investigate physiological and pathological processes. They are thought to play a crucial role for the elucidation of gene function, complementing the sequencing of the genomes of humans and other organisms. Accordingly, historians and philosophers paid considerable attention to various issues concerning this aspect of experimental biology. With respect to the representational features of model organisms, that is, their status as models, the (...) main focus was on generalization of phenomena investigated in such experimental systems. Model organisms have been said to be models for other organisms or a higher taxon. This, however, presupposes a representation of the phenomenon in question. I will argue that prior to generalization, model organisms allow researchers to built generative material models of phenomena – structures, processes or the mechanisms that explain them – through their integration in experimental set-ups that carve out the phenomena from the whole organism and thus represent them. I will use the history of zebrafish biology to show how model organism systems, from around 1970 on, were developed to construct material models of molecular mechanisms explaining developmental or physiological processes. (shrink)

Among all organisms, the size of each body part or organ scales with overall body size, a phenomenon called allometry. The study of shape and form has attracted enormous interest from biologists, but the genetic, developmental and physiological mechanisms that control allometry and the proportional growth of parts have remained elusive. Recent progress in our understanding of body‐size regulation provides a new synthetic framework for thinking about the mechanisms and the evolution of allometric scaling. In particular, insulin/IGF signaling, (...) which plays major roles in longevity, diabetes and the regulation of cell, organ and body size, might also be centrally involved in regulating organismal shape. Here we review recent advances in the fields of growth regulation and endocrinology and use them to construct a developmental model of static allometry expression in insects. This model serves as the foundation for a research program that will result in a deeper understanding of the relationship between growth and form, a question that has fascinated biologists for centuries. BioEssays 29:536–548, 2007. © 2007 Wiley Periodicals, Inc. (shrink)

When considering the question of possible transformation and disappearance of scientific objects, it is useful to distinguish between epistemic and technical objects. This paper presents preliminary observations and offers a typology of obsolescence. It is based on several case studies drawn from the history of life sciences. The paper proceeds as follows: first, the dynamics of epistemic objects is considered through the examples of Carl Correns’ study of “xenia”, Alfred Kühn’s work on physiological developmental genetics, and Paul (...) Zamecnik’s research on the protein biosynthesis. Second, the phasing out of technical objects is then separately discussed and illustrated by the example of radioactive tracing in biology – until recently, an established technique of visualization. (shrink)

Large‐scale patterns of correlated growth in development are partially driven by competition for metabolic and informational resources. It is argued that competition between organs for limited resources is an important mesoscale morphogenetic mechanism that produces fitness‐enhancing correlated growth. At the genetic level, the growth of individual characters appears independent, or “modular,” because patterns of expression and transcription are often highly localized, mutations have trait‐specific effects, and gene complexes can be co‐opted as a unit to produce novel traits. However, body parts (...) are known to interact over the course of ontogeny, and these reciprocal exchanges can be an important determinant of developmental outcomes. Genetic mechanisms underlie cell and tissue behaviors that allow organs to communicate with one another, but they also create evolutionarily adaptive competitive dynamics that are driven by physiological and biophysical processes. Advances in the understanding of competitive and closely related coordinative interactions across scales will complement existing research programs that emphasize the role of cellular mechanisms in morphogenesis. Study of the large‐scale order produced by competitive dynamics promises to facilitate advances in basic evolutionary and developmental biology, as well as applied research in fields such as bioengineering and regenerative medicine that aim to regulate patterning outcomes. (shrink)

There have been two different schools of thought on the evolution of dominance. On the one hand, followers of Wright [Wright S. 1929. Am. Nat. 63: 274–279, Evolution: Selected Papers by Sewall Wright, University of Chicago Press, Chicago; 1934. Am. Nat. 68: 25–53, Evolution: Selected Papers by Sewall Wright, University of Chicago Press, Chicago; Haldane J.B.S. 1930. Am. Nat. 64: 87–90; 1939. J. Genet. 37: 365–374; Kacser H. and Burns J.A. 1981. Genetics 97: 639–666] have defended the view that (...) dominance is a product of non-linearities in gene expression. On the other hand, followers of Fisher [Fisher R.A. 1928a. Am. Nat. 62: 15–126; 1928b. Am. Nat. 62: 571–574; Bürger R. 1983a. Math. Biosci. 67: 125–143; 1983b. J. Math. Biol. 16: 269–280; Wagner G. and Burger R. 1985. J. Theor. Biol. 113: 475–500; Mayo O. and Reinhard B. 1997. Biol. Rev. 72: 97–110] have argued that dominance evolved via selection on modifier genes. Some have called these “physiological” versus “selectionist,” or more recently [Falk R. 2001. Biol. Philos. 16: 285–323], “functional,” versus “structural” explanations of dominance. This paper argues, however, that one need not treat these explanations as exclusive. While one can disagree about the most likely evolutionary explanation of dominance, as Wright and Fisher did, offering a “physiological” or developmental explanation of dominance does not render dominance “epiphenomenal,” nor show that evolutionary considerations are irrelevant to the maintenance of dominance, as some [Kacser H. and Burns J.A. 1981. Genetics 97: 639–666] have argued. Recent work [Gilchrist M.A. and Nijhout H.F. 2001. Genetics 159: 423–432] illustrates how biological explanation is a multi-level task, requiring both a “top-down” approach to understanding how a pattern of inheritance or trait might be maintained in populations, as well as “bottom-up” modeling of the dynamics of gene expression. (shrink)

We marvel at the social complexity of insects, marked by anatomically and behaviorally distinguishable castes, division of labor and specialization—but how do such systems evolve? Insect societies are composed of individuals, each undergoing its own developmental process and each containing its own genetic information and experiencing its own developmental and experiential environment. Yet societies appear to function as if the colonies themselves are individuals with novel “social genes” and novel social developmental processes. We propose an alternative hypothesis. (...) The origins of complex social behavior, from which insect societies emerge, are derived from ancestral developmental programs. These programs originated in ancient solitary insects and required little evolutionary remodeling. We present evidence from behavioral assays, selective breeding, genetic mapping, functional genomics and endocrinology, and comparative anatomy and physiology. These insights explain how complex social behavior can evolve from heterochronic changes in reproductive signaling systems that govern ubiquitous and ancient relationships between behavior and ovarian development. BioEssays 29:334–343, 2007. © 2007 Wiley Periodicals, Inc. (shrink)

We review the evolution of domestic animals, emphasizing the effect of the earliest steps of domestication on its course. Using the first domesticated species, the dog (Canis familiaris), for illustration, we describe the evolutionary peculiarities during the historical domestication, such as the high level and wide range of diversity. We suggest that the process of earliest domestication via unconscious and later conscious selection of human‐defined behavioral traits may accelerate phenotypic variations. The review is based on the results of a long‐term (...) experiment designed to reproduce early mammalian domestication in the silver fox (Vulpes vulpes) selected for tameability or amenability to domestication. We describe changes in behavior, morphology and physiology that appeared in the fox during its selection for tameability, which were similar to those observed in the domestic dog. Based on the data of the fox experiment and survey of relevant data, we discuss the developmental, genetic and possible molecular genetic mechanisms underlying these changes. We ascribe the causative role in evolutionary transformation of domestic animals to the selection for behavior and to the neurospecific regulatory genes it affects. (shrink)

IntroductionChildhood maltreatment is a developmental risk factor and can negatively influence later psychological functioning, health, and development in the next generation. A comprehensive understanding of the biopsychosocial underpinnings of CM transmission would allow to identify protective factors that could disrupt the intergenerational CM risk cycle. This study examined the consequences of maternal CM and the effects of psychosocial and biological resilience factors on child attachment and stress-regulatory development using a prospective trans-disciplinary approach.MethodsMother-child dyads participated shortly after parturition, after 3 (...) months, and 12 months later. Mothers’ CM experiences were assessed at t0, attachment representation at t1 and psychosocial risk and social support were assessed at t1 and t2. At t2, dyads participated in the Strange Situation Procedure. Children’s attachmen status were classified as organized vs. disorganized, including their level of disorganized behavior, and heart rate and respiratory sinus arrhythmia were recorded as stress response measures of the autonomic nervous system. Maternal caregiving during SSP was assessed using the AMBIANCE scale. Child’s single nucleotide polymorphisms rs2254298 within the oxytocin receptor and rs2740210 of the oxytocin gene were genotyped using DNA isolated from cord blood.ResultsMaternal CM experiences were significantly associated with an unresolved attachment status, higher perceived stress and more psychological symptoms. These negative effects of CM were attenuated by social support. As expected, maternal unresolved attachment and child disorganized attachment were significantly associated. Maternal caregiving did not mediate the relationship between maternal and child attachment but influenced children’s HR and RSA response and disorganized behavior. Moreover, the rs2254298 genotype of the OXTR gene moderated the stress response of children from mothers with CM. Children carrying the rs2740210 risk allele of the OXT gene showed more disorganized behavior independent from maternal CM experiences.ConclusionWe replicated and extended existing CM and attachment models by co-examining maternal attachment, social support, and child genetic susceptibility on child attachment and cardiovascular stress regulation. The findings contribute to an extended understanding of risk and resilience factors and enable professionals to target adequate services to parents and children at risk. (shrink)

Holobionts are multicellular eukaryotes with multiple species of persistent symbionts. They are not individuals in the genetic sense— composed of and regulated by the same genome—but they are anatomical, physiological, developmental, immunological, and evolutionary units, evolved from a shared relationship between different species. We argue that many of the interactions between human and microbiota symbionts and the reproductive process of a new holobiont are best understood as instances of reciprocal scaffolding of developmental processes and mutual construction of (...) developmental, ecological, and evolutionary niches. Our examples show that mother, fetus, and different symbiotic microbial communities induce or constitute conditions for the development and reproduction of one another. These include the direct induction of maternal or fetus physiological changes, the restructuring of ecological relations between communities, and evolutionary selection against undesirable competitors. The mutual scaffolding and niche constructing processes start early—prior to amniotic rupture. We are evolutionarily, physiologically, and developmentally integrated holobiont systems, strung together through mutual reliance and mutual construction. Bringing the processes of niche construction and developmental scaffolding together to interpret holobiont birth conceptually scaffolds two new directions for research: in niche construction, identifying the evolutionary implications of organisms actively constructing multiple overlapping niches and scaffolds, and in Evolutionary Developmental Biology, characterizing evolutionary and ecological processes as developmental causes. (shrink)

Biological research on aggression is increasingly consulted for possible answers to the social problems of crime and violence. This paper reviews some contrasting approaches to the biological understanding of behavior—behavioral genetic, social-environmental, physiological, developmental—as a prelude to arguing that approaches to aggression are beset by vagueness and imprecision in their definitions and disunity in their measurement strategies. This vagueness and disunity undermines attempts to compare and evaluate the different approaches empirically. Nevertheless, the definitions reveal commitments to particular metaphysical (...) views concerning behavior. Alternative understandings of aggression suggest even different research approaches than those most commonly found in the literature. (shrink)

Psychological individualism is motivated by two taxonomic principles: (i) that psychological states are individuated by their causal powers, and (ii) that causal powers supervene upon intrinsic physiological state. I distinguish two interpretations of individualism--the 'orthodox' and the 'alternative'--each of which is consistent with these motivating principles. I argue that the alternative interpretation is legitimately individualistic on the grounds that it accurately reflects the actual taxonomic practices of bona fide individualistic sciences. The classification of homeobox genes in developmental (...) class='Hi'>genetics provides an illustration. When applied to the taxonomy of psychological kinds, alternative individualism has some surprising consequences. In particular, externalist taxonomies of thought are consistent with the alternative interpretation, and hence consistent with individualism. I conclude, on this basis, that the individualism/externalism dispute which has long preoccupied philosophy of psychology is an empty one. (shrink)

Scholarly opinions vary on what language is, how it evolved, and from where or what it evolved. Long considered uniquely human, today scholars argue for evolutionary continuity between human language and animal communication systems. But while it is generally recognized that language is an evolving communication system, scholars continue to debate from which species language evolved, and what behavioral and cognitive features are the precursors to human language. To understand the nature of protolanguage, some look for homologs in gene functionality, (...) brain areas, or anatomical structures such as the supralaryngeal vocal tract; others point toward primates, their gestural, vocal, multimodal, and in later evolving hominins also their pantomimic communication systems; and still others draw parallels between the musicality that characterizes language and the pitch found in the numerous sounds produced by animals. Accordingly, protolanguage theories today are multiple and diverse, and protolanguages might have also been diverse. This special issue on Evolving (Proto)Language/s for Lingua bundles several of the protolanguage theories that were put forward at the sixth edition of the Ways to Protolanguage conference series, held at the Calouste Gulbenkian Foundation in Lisbon, in 2019. Not aimed at surveying all the different ways there are to conceptualize, study, and model protolanguage/s, this issue provides interested readers with good overviews on the role played in current theorizing on protolanguage/s by (paleo)anthropology, genetics, physiology, developmental, evolutionary, ecological, and pragmatic research lines. (shrink)

Biological research on aggression is increasingly consulted for possible answers to the social problems of crime and violence. This paper reviews some contrasting approaches to the biological understanding of behavior-behavioral genetic, social-environmental, physiological, developmental-as a prelude to arguing that approaches to aggression are beset by vagueness and imprecision in their definitions and disunity in their measurement strategies. This vagueness and disunity undermines attempts to compare and evaluate the different approaches empirically. Nevertheless, the definitions reveal commitments to particular metaphysical (...) views concerning behavior. Alternative understandings of aggression suggest even different research approaches than those most commonly found in the literature. (shrink)

The greatest diversity of butterflies and their host plants occurs in tropical regions. Some groups of butterflies in the tropics exhibit monophagous feeding in the larval stage, exploiting only one family of plants; others are polyphagous, feeding on plants in two or more distinct families. The two major types of tropical habitats for butterflies, namely primary and secondary forests, offer very different evolutionary opportunities for the exploitation of plants as larval food. Butterflies are faced with the major logistical problem, as (...) are many other herbivorous insects, of depositing eggs on the correct plant for successful larval feeding. This paper, using the concepts of phenotype set and spatial patchiness of resources, attemps to make some predictions as to the optimal phenotypic systems for monophagous and polyphagous feeding in tropical butterflies, as related to the spatial patchiness of larval host plants in primary and secondary forests. In addition to the secondary compound chemistry of larval host plants as playing a role in the evolution of monophagy and polyphagy, the assumption is made that the spatial patchiness of host plants within and among different families also acts as a major factor in determining optimal ranges of phenotypes for different patterns of larval feeding. Owing to the high spatial patchiness of primary forest species of canopy trees and vines, it is predicted that butterflies exploiting these will be mostly polyphagous, whereas secondary forests having stable formations of fewer plant species and larger patches of these plants, will have mostly monophagous species. Forest understories may have both monophagous and polyphagous species, depending upon the layer of forest and the general type of understory (i.e. palmaceous or dicotyledonous). Field data on some groups of butterflies from tropical America support these predictions. Polyphagous butterflies are predicted to possess a genetic system of mixed morphs with a population being polymorphic as a whole; monophagous butterflies are predicted to have individuals all more or less similar genetically, and with a high amount of genic variation within individuals. Other forms of monophagy may evolve in species that are essentially monomorphic but with various mechanisms (physiological, developmental, behavioral) of phenotypic flexibility at the individual level. Although the environment is essentially coarse-grained for larvae since most are sedentary and polymorphism is an optimal adaptive strategy, the oviposition strategy of the adult must also be considered and some situations (i.e. forest canopy) have resources (host plants) distributed in a fine-grained fashion. Other forms of limited polyphagy may result from monomorphic genetic systems in which there is considerable phenotypic flexibility. (shrink)

Much contemporary biology consists of identifying the molecular components that associate to perform biological functions, then discovering how these functions are controlled. The concept of control is key to biological understanding, at least of the physiological kind; identifying regulators of processes underpins ideas of causality and allows complicated, multicomponent systems to be summarized in relatively simple diagrams and models. Unfortunately, as this article demonstrates by drawing on published articles, there is a growing tendency for authors to claim that a (...) molecule is a ‘regulator’ of something on evidence that cannot support the conclusion. In particular, gene knockout experiments, which can demonstrate only that a molecule is necessary for a process, are all too frequently being misinterpreted as revealing regulation. This logical error threatens to blur the important distinction between regulation and mere necessity and therefore to weaken one of our strongest tools for comprehending how organisms work. (shrink)

The concept of homology arose from classical studies of comprative morphology, and took on a new signficance with the advent of evolutionary theory. It is currentlyl undergoing antoher metamorphosis: many developmental geneticists now dfine homology as shared patterns of gene expression. However, this ne usage conflaes difinition with criteri, and fails to recognize the meaninful asignments of homology must speify a biologcal level. We argue the although developmental genetic data can help identify homologus structures. they are niether necessary (...) nor sufficient, and do not in any case jutify a new definition of homology. (shrink)

Although features of the dentition figure prominently in discussions of early hominid phylogeny, remarkably little is known of the developmental basis of the variations in occlusal morphology and dental proportions that are observed among taxa. Recent experiments on tooth development in mice have identified some of the genes involved in dental patterning and the control of tooth specification. These findings provide valuable new insight into dental evolution and underscore the strong developmental links that exist among the teeth and (...) the jaws and cranium. The latter has important implications for cladistic studies that traditionally consider features of the skull independently from the dentition. BioEssays 23:481–493, 2001. © 2001 John Wiley & Sons, Inc. (shrink)

The discovery, more than 60 years ago, of a mutant mouse with a short tail led to the birth of the new field of developmental genetics. Over the years since, numerous investigators have probed the biology of the original short‐tail mutation at the T locus, as well the naturally‐occurring t haplotypes that were uncovered as a result of their interaction with this mutation. Although the T locus ranks among the best characterized developmental loci in the mouse, it (...) was not among the first to be cloned. This situation has now been rectified with two recent reports from Herrmann, Lehrach and their colleagues. While the T locus is expressed uniquely in the embryonic tissues predicted from the mutant phenotype, the gene itself, as well as the predicted amino acid sequence of the T product, show no strong homology to any known sequence. For the moment, at least, the mystery behind the function of the T locus still awaits definitive resolution. (shrink)

Physiological experiments conducted over the last 60 years indicate that the plant hormone auxin regulates a diverse set of developmental processes via changes in cell division, cell elongation and cell differentiation. Recent studies using transgenic plants with altered auxin levels support these conclusions and promise to provide more detailed information on the role of auxin during plant development. Although it is possible that all auxin responses are mediated by the same primary biochemical events, the studies described in this (...) review are more consistent with multiple modes of auxin action. The development of molecular and genetic approaches to the study of hormone action should resolve this issue. The accelerated rate of progress in this field suggests that real insight into the mechanism of auxin action may be forthcoming. (shrink)

The role of fixed charges present at the surface of biological membranes is usually described by the Gouy-Chapman-Grahame theory of the electric double-layer where the Grahame equation is applied independently on each side of the membrane and where the capacitive charges are disregarded. In this article, we generalize the Gouy-Chapman-Grahame theory by taking into account both intrinsic charges and capacitive charges, in the density value of the membrane surface charges. In the first part, we show that capacitive charges couple electrostatic (...) potentials present on both sides of the membrane. The intensity of this coupling depends both on the value of the membrane specific capacitance and the transmembrane electric potential difference. In the second part, we suggest some physiological implications of membrane electric double-layers. (shrink)

The mechanisms responsible for the fine tuning of development, where the wildtype phenotype is reproduced with high fidelity, are not well understood. The difficulty in approaching this problem is the identification of mutant phenotypes indicative of a defect in these fine‐tuning control mechanisms. Evolutionary biologists have used asymmetry as a measure of developmental homeostasis. The rationale for this was that, since the same genome controls the development of the left and right sides of a bilaterally symmetrical organism, departures from (...) symmetry can be used to measure genetic or environmental perturbations. This paper examines the relationship between asymmtry and resistance to organophosphorous insecticides in the Australian sheep blowfly, Lucilia cuprina. A resistance gene, Rop‐1, which encodes a carboxylesterase enzyme, also confers a significant increase in asymmetry. Continued exposure of resistant populations to insecticide has selected a dominant suppressor of the asymmetry phenotype. Genetic evidence indicates that the modifier is the L. cuprina Notch homologue. (shrink)

The Epic of Evolution is a course taught at Northern Arizona University. It engages the task of formulating a new epic myth that is based on the physical, natural, social, and cultural sciences. It aims to serve the need of providing meaning for human living in the vast and complex universe that the sciences now depict for us. It is an interdisciplinary effort in an academic setting that is often divided by specializations; it focuses on values in a climate of (...) relativism; and it concentrates on an enterprise for which there are few textbooks at hand. The course is presented in three segments: the cosmos before humans appeared, the human phenomenon, and scenarios for the future of evolution. (shrink)

The last decade has seen a dramatic increase in interest in the extent to which morphological evolution depends on changes in regulatory pathways. Insects provide a fertile ground for study because of their diversity and our high level of understanding of the genetic regulation of development in Drosophila melanogaster. However, comparable genetic approaches are presently possible in only a small number of non‐Drosophilid insects. In a recent paper, Hughes and Kaufman(1) have used a new methodology, RNA interference, in the milkweed (...) bug, Oncopeltus fasciatus, to phenocopy the effects of mutations in Hox genes. RNA interference involves the injection of double‐stranded RNA of the same sequence as the relevant mRNA resulting in a depletion of that transcript.(2) Hughes and Kaufman focused on the gnathal segments, which elaborate specialized appendages important to feeding. Their results indicate that gnathal adaptations in this bug are correlated with changes in Hox gene functions and interactions. BioEssays 23:379–382, 2001. © 2001 John Wiley & Sons, Inc. (shrink)

The unusual occurrence and developmental diversity of asexual eukaryotes remain a puzzle. De novo formation of a functioning asexual genome requires a unique assembly of sets of genes or gene states to disrupt cellular mechanisms of meiosis and gametogenesis, and to affect discrete components of sexuality and produce clonal or hemiclonal offspring. We highlight two usually overlooked but essential conditions to understand the molecular nature of clonal organisms, that is, a nonrecombinant genomic assemblage retaining modifiers of the sexual program, (...) and a complementation between altered reproductive components. These subtle conditions are the basis for physiologically viable and genetically balanced transitions between generations. Genomic and developmental evidence from asexual animals and plants indicates the lack of complementation of molecular changes in the sexual reproductive program is likely the main cause of asexuals' rarity, and can provide an explanatory frame for the developmental diversity and lability of developmental patterns in some asexuals as well as for the discordant time to extinction estimations. (shrink)

In one of the first genetic screens aimed at identifying induced developmental mutants, Nadine Dobrovolskaïa-Zavadskaïa, working at the Pasteur Laboratory in the 1920s, isolated and characterized a mutation affecting Brachyury, a gene that regulates tail and axial development in the mouse. Dobrovolskaïa-Zavadskaïa's analysis of Brachyury and other mutations affecting tail development were among the earliest attempts to link gene action with a tissue-specific developmental process in a vertebrate. Her analyses of genes that interacted with Brachyury led to the (...) discovery of the t-haplotype chromosome of mouse. After 70 years, Brachyury and the multiple genes with which it interacts continue to occupy a prominent focus in developmental biology research. A goal of this review is to identify the contributions that Dobrovolskaïa-Zavadskaïa made to our current thinking about Brachyury and how she helped to shape the dawn of the field of developmental genetics. BioEssays 23:365–371, 2001. © 2001 John Wiley & Sons, Inc. (shrink)

Ludwik Fleck, Edmund Husserl : on the historicity of scientific knowledge -- Gaston Bachelard : the concept of "phenomenotechnique" -- Georges Canguilhem : epistemological history -- Pisum : Carl Correns's experiments on Xenia, 1896-99 -- Eudorina : Max Hartmann's experiments on biological regulation in protozoa, 1914-21 -- Ephestia : Alfred Kähn's experimental design for a developmental physiological -- Genetics, 1924-45 -- Tobacco mosaic virus : virus research at the Kaiser Wilhelm Institutes for Biochemistry and Biology, 1937-45 -- (...) The concept of the gene : molecular biological perspectives -- The liquid scintillation counter : traces of radioactivity -- The concept of information : the writings of François Jacob -- Intersections -- Preparations -- The economy of the scribble. (shrink)

Recent work by Magnuson, Solomon and Grossman(1) adds to a growing body of evidence indicating that microorganisms possess sophisticated signaling systems that enable them to sense and respond to environmental challenges. Typically, this response results in morphological, physiological and even genetic differentiation, paralleling that observed among higher organisms. These signaling systems may be interpreted as adaptations that maximize the reproductive potential of a population.

Every cell, tissue, organ and organism is competent to use signs to exchange information reaching common coordinations and organisations of both single cell and group behavior. These sign-mediated interactions we term biological communication. The regulatory system that works in development, morphology, cell fate and identity, physiology, genetic instructions, immunity, memory/learning, physical and mental disease depends on epigenetic marks. The communication of cells, persistent viruses and their defectives such as mobile genetic elements and RNA networks ensures both the transport of regulatory (...) instructions and the reprogramming of these instructions. But how are the different states of the epigenome orchestrated? The epigenetic pathways respond to various signaling cues such as DNA methylation, histone variants, histone modifications, chromatin structure, nucleosome remodeling, and epigenetic interactions. Epigenetic signals are responsible for the establishment, maintenance and reversal of transcriptional states that are fundamental for the cell's ability to memorize past events, such as changes in the external environment, socio-sphere or developmental cues. External signals trigger changes in the epigenome, allowing cells to respond dynamically. Internal signals direct activities that are necessary for body maintenance, and repairing damaged tissues and organs. With the emergence of epigenetic memory, organisms can fix historical and context dependent impressive experiences. Evolution from now on learnt to learn. Learning means organisms can avoid reproduction of always the same. This is key to adaptation. However, inheritance of acquired characteristics is only one of the many examples of the explanatory power of epigenetics. Behavioral epigenetics demonstrates the way in which environmental and social experiences produce individual differences in behaviour, cognition, personality, and mental health. This book assembles experts to outline the various motifs of all kinds of epigenetic regulation of cells throughout their lives. (shrink)

The existence of a genetic program of development was proposed by molecular biologists in the nineteen-sixties. Historians and philosophers of science have since thoroughly criticized this notion. To fully appreciate its significance, it is interesting to consider the research which was pursued during this period by molecular biologists who proposed this notion. This study focuses on François Jacob's work and on the model of development supported by his lab in the early seventies, the T-complex model. This episode of Jacob's scientific (...) activity has since been forgotten. Characterization of this model shows that the notion of program was used in a metaphoric way and that it did not put any constraint on the work pursued in the lab at that time. Some attention is devoted to the origin of this metaphor in the context of the nineteen-seventies. (shrink)

Modern scientific theories of emotional behavior, almost without exception, trace their origin to Charles Darwin, and his publications On the Origin of Species (1859) and The Expression of the Emotions in Man and Animals (1872). The most famous evolutionary dilemma Darwin acknowledged as a challenge to his theory of natural selection was the incomplete sub Cambrian fossil record. However, Darwin struggled with two other rarely referenced theoretical and scientific dilemmas that confounded his theories about emotional behavior. These included (1) the (...) origin of social instincts (e.g., altruism, empathy, reciprocity and cooperation) and the reasons for their conservation in evolution and (2) the peripheral control of heart rate vis-a-vis emotional behavior outside of consciousness. Darwin acknowledged that social instincts are critical to the survival of some species, but had difficulty aligning them with his theory of natural selection in humans. Darwin eventually proposed that heart rate and emotions are controlled via one’s intellect and cortical mechanisms, and that instinctive behavior is genetically-programmed and inherited. Despite ongoing efforts, these two theoretical dilemmas are debated to this day. Simple testable hypotheses have yet to emerge for the biological mechanisms underlying instinctive behavior or the way heart rate is controlled in infants. In this paper, we review attempts to resolve ¬these issues over the past 160 years. We posit that research and theories that supported Darwin’s individualistic brain-centric and genetic model have become an “orthodox” Western view of emotional behavior, one that produced the prevailing behavioral construct of attachment as developed by John Bowlby. We trace research and theories that challenged this orthodoxy at various times, and show how these challenges were repeatedly overlooked, rejected, or misinterpreted. We review two new testable theories, emotional connection theory and calming cycle theory, which we argue resolve the two dilemmas We show emerging scientific evidence from physiology and a wide variety of other fields, as well from clinical trials among prematurely born infants, that supports the two theories. Clinical implications of the new theories and possible new ways to assess risk and intervene in emotional, behavioral and developmental disorders are discussed. (shrink)

This article explores the collaborative research of the Nobel laureate Hans Spemann (1869–1941) and the Swiss zoologist Fritz Baltzer (1884–1974) on problems at the intersection of development and heredity and raises more general questions concerning science and politics in Germany in the interwar period. It argues that Spemann and Baltzer’s collaborative work made a significant contribution to the then ongoing debates about the relation between developmental physiology and hereditary studies, although Spemann distanced himself from _Drosophila_ genetics because of (...) his anti-reductionist position. The article analyzes how Spemann framed the issues of heredity in terms of an epigenetic principle in the context of his work on the “organizer,” and it explores the experimental dynamics of research on newt merogones carried out by Baltzer in a methodological development of Spemann’s constriction experiments. Finally, these research attempts are discussed as part of a broader “prehistory” of the mid-twentieth century cell nuclear transplantation experiments, which provided the basis for later animal cloning. (shrink)

Sociopaths are “outstanding” members of society in two senses: politically, they draw our attention because of the inordinate amount of crime they commit, and psychologically, they hold our fascination because most ofus cannot fathom the cold, detached way they repeatedly harm and manipulate others. Proximate explanations from behavior genetics, child development, personality theory, learning theory, and social psychology describe a complex interaction of genetic and physiological risk factors with demographic and micro environmental variables that predispose a portion of (...) the population to chronic antisocial behavior. More recent, evolutionary and game theoretic models have tried to present an ultimate explanation of sociopathy as the expression of a frequency-dependent life strategy which is selected, in dynamic equilibrium, in response to certain varying environmental circumstances. This paper tries to integrate the proximate, developmental models with the ultimate, evolutionary ones, suggesting that two developmentally different etiologies of sociopathy emerge from two different evolutionary mechanisms. Social strategies for minimizing the incidence of sociopathic behavior in modern society should consider the two different etiologies and the factors that contribute to them. (shrink)

In lieu of an abstract, here is a brief excerpt of the content:Normalization and the Welfare StateLadelle McWhorterIn Racism and Sexual Oppression in Anglo-America, I argued that as race was absorbed into biology in the nineteenth century, it was recast from a morphological typology to a function of physiological and evolutionary development (McWhorter 2009b). Racial difference became a sign of developmental difference. Racial groups represented stages of human evolution, and raced individuals were to be disciplined and managed in (...) accordance with developmental norms.Twentieth-century race was therefore a category and tool of normalization; to be raced was to be located at a particular point along a developmental curve. Some races were thought to be more or less arrested in their development, while others were merely retarded, and theorists disagreed at times about which were which. But they all agreed that the most highly developed race was the Nordic or the Anglo-Saxon. Anglo-Saxons were the norm for modern human beings— anatomically, physiologically, intellectually, and morally. There were other white races whose status was debated (Mediterraneans and Ashkenazi Jews, for example), and there were disputes over the evolutionary status of Chinese, Hindus, and Mexicans. But most theorists agreed that at the very bottom of the developmental scale we should place Celts, Negroes, and American Indians. These peoples were savages, primitives. They should be trained to the extent possible to contribute to the society that had already superseded them, but their proper destiny was to pass away into extinction as humanity advanced.These evolutionary stragglers, along with the evolutionary throwbacks that appeared sometimes among whites, were considered dangerous for two main reasons. First, a great many of them were expensive. They were shiftless, often incompetent, and frequently lawless. Because of them, it was necessary to [End Page 39] maintain police forces, prisons, poorhouses, and asylums. They were a drain on the economic advance of society, which affected its educational and technological advance and ultimately its evolutionary advance as well. Second, because they had primitive sexual urges unrestrained by reason as well as heritable defects and diseases, they threatened the bloodlines of more advanced human beings. In short, they posed a risk for the ongoing evolutionary advancement of the entire Human Race. To protect the Race, it was necessary to contain these primitives, savages, and throwbacks and to reduce their numbers through careful management of their sexuality. This, I argued in the book, was racism’s cutting edge in the early twentieth century.Since publishing the book, I have done some work on the changes in practices of racism in the wake of the Biological Synthesis—that is, the unification of biology and genetics that occurred in the 1930s through the efforts of renowned scientists such as Theodosius Dobzhansky and Ernst Mayr (McWhorter 2010a, 2010b, 2010c). As I discussed in that work, by 1950, evolutionary biology was less a study of species development than it was a study of the effects of evolutionary forces acting on and within populations. Correlatively, racial practices shifted toward management of statistically delineated populations or ethnic groups. Theorists and policymakers recognized that not all members of a given population have the same genes, meaning that old-style racism (which assumed, for example, that all Negroes were stupid) was false; some Negroes might be quite brilliant, in fact. It was not The Negro who was stupid but the population of Negroes in the aggregate that was measurably stupider than the population of Anglo-Saxons. Accordingly, normalization now occurred at the level of the population. Disciplinary practices designed for cultivating the capacities of groups of individuals were altered or replaced to focus on groups as groups, as aggregates. The point was to shift the statistical norm of the population rather than to alter the bodies or behavior of each and every individual. I have argued elsewhere (McWhorter 2009a, 5) that this merger of disciplinary normalization and population management constitutes the full flowering of what Foucault calls biopower (Foucault 1978, 140).When Lisa Guenther invited me to speak at the plenary on “Re-thinking Whiteness” that took place at the fifth annual meeting of philoSOPHIA at Vanderbilt University, she explained that although many presenters offer polished papers, it has been a... (shrink)

The C. elegans male tail is being studied as a model to understand how genes specify the form of multicellular animals. Morphogenesis of the specialized male copulatory organ takes place in the last larval stages during male development. Genetic analysis is facilitated because the structure is not necessary for male viability or for strain propagation. Analysis of developmental mutants, isolated in several functional and morphological screens, has begun to reveal how fates of cells are determined in the cell lineages, (...) and how the specification of cell fates affects the morphology of the structure. Cytological studies in wild type and in mutants have been used to study the mechanism of pattern formation in the tail peripheral nervous system. The ultimate goal is to define the entire pathway leading to the male copulatory organ. (shrink)

Understanding how traits are integrated at the organismal level remains a fundamental problem at the interface of developmental and evolutionary biology. Hormones, regulatory signaling molecules that coordinate multiple developmental and physiological processes, are major determinants underlying phenotypic integration. The probably best example for this is the lipid-like juvenile hormone (JH) in insects. Here we review the manifold effects of JH, the most versatile animal hormone, with an emphasis on the fruit fly Drosophila melanogaster, an organism amenable to (...) both genetics and endocrinology. JH affects a remarkable number of processes and traits in Drosophila development and life history, including metamorphosis, behavior, reproduction, diapause, stress resistance and aging. While many molecular details underlying JH signaling remain unknown, we argue that studying “hormonal pleiotropy” offers intriguing insights into phenotypic integration and the mechanisms underlying life history evolution. In particular, we illustrate the role of JH as a key mediator of life history trade-offs. BioEssays 27:999–1010, 2005. © 2005 Wiley Periodicals, Inc. (shrink)

The concept of plasticity has always been present in the history of developmental biology, both within the theory of epigenesis and within morphogenesis studies. However this tradition relies also upon a genetic conception of plasticity. Founded upon the concepts of ‘‘phenotypic plasticity’’ and ‘‘reaction norm,’’ this genetic conception focuses on the array of possible phenotypic change in relation to diversified environments. Another concept of plasticity can be found in recent publications by some developmental biologists (Gilbert, West-Eberhard). I argue (...) that these authors adopt a ‘‘broad conception of plasticity’’ that is closely related to a notion of development as something that is ongoing throughout an organism’s lifecycle, and has no clear-cut boundaries. However, I suggest that given a narrow conception of plasticity, one can define temporal boundaries for development that are linked to specific features of the morphological process, which are different from behavioral and physiological processes. (shrink)