The concept of adaptation is employed in many fields such as biology, psychology, cognitive sciences, robotics, social sciences, even literacy and art,1 and its meaning varies quite evidently according to the particular research context in which it is applied. We expect to find a particularly rich catalogue of meanings within evolutionary biology, where adaptation has held a particularly central role since Darwin’s The Origin of Species (1859) throughout important epistemological shifts and scientific findings that enriched and diversified the concept. Accordingly, (...) a conceptual taxonomy of adaptation in evolutionary biology may help to disambiguate it. Interdisciplinary researches focused on adaptation would benefit from such a result. In the present work we recognize and define seven different meanings of adaptation: (1) individual fitness; (2) adaptation of a population; (3) adaptation as the process of natural selection; (4) adaptive traits; (5) molecular adaptation; (6) adaptation as structural tinkering; (7) plasticity. For convenience here, we refer to them as W-, P-, NS-, T-, M-, S- and PL-ADAPTATION. We present the seven meanings in some detail, hinting at their respective origins and conceptual developments in the history of evolutionary thought (references are offered for further deepening). However, it is important to point out that evolution researchers seldom if ever refer to a single meaning purified from the others. This applies also to the authors we cite as representatives of one of the seven meanings. In Discussion and Conclusion draw from our work some future perspectives for adaptation within evolutionary biology. (shrink)

This inaugural handbook documents the distinctive research field that utilizes history and philosophy in investigation of theoretical, curricular and pedagogical issues in the teaching of science and mathematics. It is contributed to by 130 researchers from 30 countries; it provides a logically structured, fully referenced guide to the ways in which science and mathematics education is, informed by the history and philosophy of these disciplines, as well as by the philosophy of education more generally. The first handbook to cover the (...) field, it lays down a much-needed marker of progress to date and provides a platform for informed and coherent future analysis and research of the subject. -/- The publication comes at a time of heightened worldwide concern over the standard of science and mathematics education, attended by fierce debate over how best to reform curricula and enliven student engagement in the subjects There is a growing recognition among educators and policy makers that the learning of science must dovetail with learning about science; this handbook is uniquely positioned as a locus for the discussion. -/- The handbook features sections on pedagogical, theoretical, national, and biographical research, setting the literature of each tradition in its historical context. Each chapter engages in an assessment of the strengths and weakness of the research addressed, and suggests potentially fruitful avenues of future research. A key element of the handbook’s broader analytical framework is its identification and examination of unnoticed philosophical assumptions in science and mathematics research. It reminds readers at a crucial juncture that there has been a long and rich tradition of historical and philosophical engagements with science and mathematics teaching, and that lessons can be learnt from these engagements for the resolution of current theoretical, curricular and pedagogical questions that face teachers and administrators. (shrink)

ArgumentContrary to common portrayals of social Darwinism as a transference of laissez-faire values, the widely read evolutionism of Japan's foremost Darwinist of the early twentieth-century, Oka Asajirō (1868–1944), reflects a statist outlook that regards capitalism as the beginning of the nation's degeneration. The evolutionary theory of orthogenesis that Oka employed in his 1910 essay, “The Future of Humankind,” links him to a pre-Darwinian idealist tradition that depicted the state as an organism that develops through life-cycle stages. For Oka, laissez-faire capitalism (...) marked the moment when the state began to decline toward extinction due to the orthogenetic overdevelopment of hitherto subordinate individual egos. Because conservative bureaucrat-intellectuals had been drawing upon this same organicist-developmental tradition since the 1880s in an attempt to forestall the social ills of industrialism, Oka's call for statist measures, including eugenics, to lessen and delay the atomizing, enervating, and corrupting influence of capitalism articulated the political vision of officialdom. Statist evolutionism, not social Darwinism, might be the term that best describes Oka's approach. (shrink)

The peripheral nervous system has classically been separated into a somatic division composed of both afferent and efferent pathways and an autonomic division containing only efferents. J. N. Langley, who codified this asymmetrical plan at the beginning of the twentieth century, considered different afferents, including visceral ones, as candidates for inclusion in his concept of the “autonomic nervous system”, but he finally excluded all candidates for lack of any distinguishing histological markers. Langley's classification has been enormously influential in shaping modern (...) ideas about both the structure and the function of the PNS. We survey recent information about the PNS and argue that many of the sensory neurons designated as “visceral” and “somatic” are in fact part of a histologically distinct group of afferents concerned primarily autonomic function. These afferents have traditionally been known as “small dark” neurons or B-neurons. In this target article we outline an association between autonomic and B-neurons based on ontogeny, cell phenotype, and functional relations, grouping them together as part of a common reflex system involved in homeostasis. This more parsimonious classification of the PNS, made possible by the identification of a group of afferents associated primarily with the ANS, avoids a number of confusions produced by the classical orientation. It may also have practical implications for an understanding of nociception, homeostatic reflexes, and the evolution of the nervous system. (shrink)

In this paper I suggest that play is a distinctive behavioural category whose adaptive significance calls for explanation. Play primarily affords juveniles practice toward the exercise of later skills. Its benefits exceed its costs when sufficient practice would otherwise be unlikely or unsafe, as is particularly true with physical skills and socially competitive ones. Manipulative play with objects is a byproduct of increased intelligence, specifically selected for only in a few advanced primates, notably the chimpanzee.The adaptiveness of play in pongid (...) evolution is traced through the probable changes in selective pressures that occurred in hominid evolution. It is argued that fantasy was an emergent property in hominids, made possible by symbolic intelligence and language, and serving to make play complex enough to continue to provide useful practice for increasingly complex later skills.The advent of organised instruction and education has meant that play's unplanned, intrinisic goal-setting could be replaced by extrinsic goal-setting in the systematic development of particular skills. However, the need to ensure adequate motivation has continued to give play educational value. In addition, its capacity to enhance innovative behaviour seems to be a residual function of play which has acquired a new cultural importance. (shrink)

Despite its importance, the linkage between genetic and cultural evolution has until now been little explored. An understanding of this linkage is needed to extend evolutionary theory so that it can deal for the first time with the phenomena of mind and human social history. We characterize the process of gene-culture coevolution, in which culture is shaped by biological imperatives while biological traits are simultaneously altered by genetic evolution in response to cultural history. A case is made from both theory (...) and evidence that genetic and cultural evolution are inseverable, and that the human mind has tended to evolve so as to bias individuals toward certain patterns of cognition and choice rather than others. With the aid of mathematical models we trace the coevolutionary circuit: The genes prescribe structure in developmental pathways that lay down endocrine and neural systems, imposing regularities in the development of cognition and behavior; these regularities translate upward into holistic patterns of culture, which can be predicted in the form of probability density distributions ; natural selection acts within human history to favor certain epigenetic rules over others; and the selection alters the frequencies of the underlying genes. The effects of genetic and cultural changes reverberate throughout the circuit and are consequently tested with the passage of each life cycle. In addition to modeling gene-culture coevolution, we apply methods from island biogeography and information theory to examine the cultural capacity of the genes, the factors determining the magnitude of cultural diversity, and the possible reasons for the uniqueness of the human achievement. (shrink)

Despite its importance, the linkage between genetic and cultural evolution has until now been little explored. An understanding of this linkage is needed to extend evolutionary theory so that it can deal for the first time with the phenomena of mind and human social history. We characterize the process of gene-culture coevolution, in which culture is shaped by biological imperatives while biological traits are simultaneously altered by genetic evolution in response to cultural history. A case is made from both theory (...) and evidence that genetic and cultural evolution are inseverable, and that the human mind has tended to evolve so as to bias individuals toward certain patterns of cognition and choice rather than others. With the aid of mathematical models we trace the coevolutionary circuit: The genes prescribe structure in developmental pathways that lay down endocrine and neural systems, imposing regularities in the development of cognition and behavior; these regularities translate upward into holistic patterns of culture, which can be predicted in the form of probability density distributions ; natural selection acts within human history to favor certain epigenetic rules over others; and the selection alters the frequencies of the underlying genes. The effects of genetic and cultural changes reverberate throughout the circuit and are consequently tested with the passage of each life cycle. In addition to modeling gene-culture coevolution, we apply methods from island biogeography and information theory to examine the cultural capacity of the genes, the factors determining the magnitude of cultural diversity, and the possible reasons for the uniqueness of the human achievement. (shrink)

Recent studies on neural pathways in a broad spectrum of vertebrates suggest that, in addition to migration and an increase in the number of certain select neurons, a significant aspect of neural evolution is a “parcellation” (segregation-isolation) process that involves the loss of selected connections by the new aggregates. A similar process occurs during ontogenetic development. These findings suggest that in many neuronal systems axons do not invade unknown territories during evolutionary or ontogenetic development but follow in their ancestors' paths (...) to their ancestral targets; if the connection is later lost, it reflects the specialization of the circuitry.The pattern of interspecific variability suggests (1) that overlap of circuits is a more common feature in primitive (generalized) than in specialized brain organizations and (2) that most projections, such as the retinal, thalamotelencephalic, corticotectal, and tectal efferent ones, were bilateral in the primitive condition. Specialization of these systems in some vertebrate groups has involved the selective loss of connections, resulting in greater isolation of functions. The parcellation process may also play an important role in cell diversification.The parcellation process as described here is thought to be one of several underlying mechanisms of evolutionary and ontogenetic differentiation. (shrink)

Responses are strengthened by consequences having to do with the survival of individuals and species. With respect to the provenance of behavior, we know more about ontogenic than phylogenic contingencies. The contingencies responsible for unlearned behavior acted long ago. This remoteness affects our scientific methods, both experimental and conceptual. Until we have identified he variables responsible for an event, we tend to invent causes. Explanatory entities such as “instincts,” “drives,” and “traits” still survive. Unable to show how organisms can behave (...) effectively under complex circumstances, we endow them with special abilities permitting them to do so.Behavior exhibited by most members of a species is often accepted as inherited if all members were not likely to have been exposed to relevant ontogenic contingencies. When contingencies are not obvious, it is perhaps unwise to call any behavior either inherited or acquired, as the examples of churring in honey guides and following in imprinted ducklings show. Nor can the relative importance of phylogenic and ontogenic contingencies be argued from instances in which unlearned or learned behavior intrudes or dominates. Intrusions occur in both directions.Behavior influenced by its consequences seems directed toward the future, but only past effects are relevant. The mere fact that behavior is adaptive does not indicate whether phylogenic or ontogenic processes have been responsible for it. Examples include the several possible provenances of imitation, aggression, and communication. The generality of such concepts limits their usefulness. A more specific analysis is needed if we are to deal effectively with the two kinds of contingencies and their products. (shrink)

Human behavior is the joint product of (i) contingencies of survival responsible for natural selection, and (ii) contingencies of reinforcement responsible for the repertoires of individuals, including (iii) the special contingencies maintained by an evolved social environment. Selection by consequences is a causal mode found only in living things, or in machines made by living things. It was first recognized in natural selection: Reproduction, a first consequence, led to the evolution of cells, organs, and organisms reproducing themselves under increasingly diverse (...) conditions. The behavior functioned well, however, only under conditions similar to those under which it was selected.Reproduction under a wider range of consequences became possible with the evolution of processes through which organisms acquired behavior appropriate to novel environments. One of these, operant conditioning, is a second kind of selection by consequences: New responses could be strengthened by events which followed them. When the selecting consequences are the same, operant conditioning and natural selection work together redundantly. But because a species which quickly acquires behavior appropriate to an environment has less need for an innate repertoire, operant conditioning could replace as well as supplement the natural selection of behavior.Social behavior is within easy range of natural selection, because other members are one of the most stable features of the environment of a species. The human species presumably became more social when its vocal musculature came under operant control. Verbal behavior greatly increased the importance of a third kind of selection by consequences, the evolution of social environments or cultures. The effect on the group, and not the reinforcing consequences for individual members, is responsible for the evolution of culture. (shrink)

In our commentary we elaborate on Barresi & Moore's use of language as a tool. In particular, we highlight the importance of cognitive linguistic research with its emphasis on the relation between morpnosyntax and intentional schemes. We also speculate about how language itself might play a role in children's integration of first and third person knowledge.

Immediate imitation is likely to be a major, direct input to Barresi & Moore's level 2 competence, but deferred imitation is unlikely to play a key role in the transition to level 3, because (1) the attribution of first person knowledge is neither a necessary cause nor an obvious consequence of deferred imitation, and (2) deferred imitation does not correlate phylogenetically with capacities that more plausibly either yield or reflect a concept of intentional agency.

The intentional schema may not be sufficiently characterized to make questions about its role in individual and species development intelligible. The idea of metarepresentation may perhaps give it enough content. The importance of metarepresentation itself, however, can be called into question.

Contrary to Müller's claims, and in support of modular theories, genetic factors play a substantial and significant role in language. The finding that some children with specific language impairment (SLI) have nonlinguistic impairments may reflect improper diagnosis of SLI or impairments that are secondary to linguistic impairments. Thus, such findings do not argue against the modularity thesis. The lexical/functional distinction appears to be innate and specifically linguistic and could be instantiated in either symbolic or connectionist systems.

Müller argues that general cognitive skills and linguistic skills are not necessarily independent. However, cross-linguistic evidence from an inherited specific language disorder affecting productive rules suggests significant degrees of modularity, innateness, and universality of language. Confident claims about the overall nature of such a complex system still await more interdisciplinary research.

Müller attempts to downplay cases of dissociation between language and cognition as evidence against the modularity of language. We review cases of associations between verbal and nonverbal abilities as further evidence against the notion of language as an autonomous subsystem. We also point out a discrepancy between his proposal of homologies between nonhuman primates' communication and human language and recent proposals on the evolution of language.

Müller misconstrues autonomy to mean strict locality of brain function, something quite different from the functional autonomy that linguists claim. Similarly, he misperceives the interaction of learned and innate components hypothesized in current generative models. Evidence from sign languages, Creole languages, and neurological studies of rare forms of aphasia also argues against his conclusions.

Despite the burgeoning interest in new and more complex accounts of the organism-environment dyad by biologists and philosophers, little attention has been paid in the resulting discussions to the history of these ideas and to their deployment in disciplines outside biology—especially in the social sciences. Even in biology and philosophy, there is a lack of detailed conceptual models of the organism-environment relationship. This volume is designed to fill these lacunae by providing the first multidisciplinary discussion of the topic of organism-environment (...) interaction. It brings together scholars from history, philosophy, psychology, anthropology, medicine, and biology to discuss the common focus of their work: entangled life, or the complex interaction of organisms and environments. (shrink)

Representations of nonhuman wild animals in children's stories and toys underwent dramatic transformation over the years 1830-1915. During the earlier part of that period, wild animals were presented to children as being savage and dangerous, and that it was necessary for them to be killed or brutally constrained. In the 1890s, an animalcentric discourse emerged in Nature writing, along with an animal-human symbiosis in scientific child study that highlighted childhood innocence, resulting in a valuing of wild animals based upon their (...) similarity to humans. This article will describe the aesthetic devices of children's stories and play materials in relation to the dominant, emerging, and residual ideas about the wild communicated by adults to children through these means. (shrink)

Averageness is purportedly the result of stabilizing selection maintaining the population mean, whereas facial paedomorphosis is a product of directional selection driving the population mean towards an increasingly juvenile appearance. If selection is predominantly stabilizing, intermediate phenotypes reflect high genetic quality and mathematically average faces should be found attractive. If, on the other hand, directional selection is strong enough, extreme phenotypes reflect high genetic quality and juvenilized faces will be found attractive. To compare the effects of stabilizing and directional selection (...) on facial paedomorphosis (juvenilization), graphic morphing and editing techniques were used to alter the appearance of composite faces to make them appear more or less juvenile. Both facial models and judges of attractiveness were from the CSU-Long Beach campus. Although effect sizes for both preferences were large, the effect for averageness was nearly twice that found for juvenilization, an indication that stabilizing selection influences preferences for facial paedomorphosis more so than directional selection in contemporary humans. (shrink)

Modern biology has not yet come to terms with the presence of many organic codes in Nature, despite the fact that we can prove their existence. As a result, it has not yet accepted the idea that the great events of macroevolution were associated with the origin of new organic codes, despite the fact that this is the most parsimonious and logical explanation of those events. This is probably due to the fact that the existence of organic codes in all (...) fundamental processes of life, and in all major transitions in the history of life, has enormous theoretical implications. It requires nothing less than a new theoretical framework, and that kind of change is inevitably slow. There are too many facts to reconsider, too many bits of history to weave together in a new mosaic. But this is what science is about, and the purpose of the present paper is to show that it can be done. More precisely, it is shown that the whole natural history of the brain can be revisited in the light of the organic codes. What is described here is only a bird’s-eye view of brain macroevolution, but it is hoped that the extraordinary potential of the organic codes can nevertheless come through. The paper contains also another message. The organic codes prove that life is based on semiosis, and are in fact the components of organic semiosis, the first and the most diffused form of semiosis on Earth, but not the only one. It will be shown that the evolution of the brain was accompanied by the development of two new types of sign processes. More precisely, it gave origin first to interpretive semiosis, mostly in vertebrates, and then to cultural semiosis, in our species. (shrink)

This thematic issue addresses questions of constraints on the evolution of form—physical, biological, and technical. Here, form is defined as an embodiment of a specific structure, which can be hierarchically different yet emerge from the same processes. The focus of this contribution is about how developmental biology and paleontology can be better integrated and compared in order to produce hypotheses about the evolution of form. The constraints on current EvoDevo research stem from the disconnect in the focus of study for (...) developmental geneticists and evolutionary morphologists; the former being interested in early developmental events at a molecular level in a model animal, the latter in late developmental events or comparison between adult forms, at a structural level in non-model animals. In order to truly integrate information from both fields in our understanding of evolutionary processes, morphology needs to be reintegrated in the study of gene expression, and its time frame needs to be extended beyond early developmental stages. Gene expression in non-model organisms also needs to be studied in order to gain perspective into primitive patterning at evolutionary nodes. Hypotheses formed by the comparison of expression patterns and morphologies seen in extant species can then be tested against forms found in the fossil record, coming closer to understanding the mechanisms underlying evolution. (shrink)

The scientific status of evolutionary theory seems to be more or less perennially under question. I am not referring here (just) to the silliness of young Earth creation- ism (Pigliucci 2002; Boudry and Braeckman 2010), or even of the barely more intel- lectually sophisticated so-called Intelligent Design theory (Recker 2010; Brigandt this volume), but rather to discussions among scientists and philosophers of science concerning the epistemic status of evolutionary theory (Sober 2010). As we shall see in what follows, this debate (...) has a long history, dating all the way back to Darwin, and it is in great part rooted in the fundamental dichotomy between what French biologist and Nobel laureate Jacques Monod (1971) called chance and necessity—i.e., the inevitable and inextricable interplay of deterministic and stochastic mechanisms operating during the course of evolution. (shrink)

It is very curious that C.G. Jung has had so little influence upon the anthropology of consciousness. In this paper, the reasons for this oversight are given. The archetypal psychology of Jung is summarized and shown to be more complex and useful than extreme constructivist accounts would acknowledge. Jung's thinking about consciousness fits very well with a modern neuroscience view of the psyche and acts as a corrective to relativist notions of consciousness and its relation to the self.

The cis-regulatory hypothesis is one of the most important claims of evolutionary developmental biology. In this paper I examine the theoretical argument for cis-regulatory evolution and its role within evolutionary theorizing. I show that, although the argument has some weaknesses, it acts as a useful example for the importance of current scientific debates for science education.

The Modern Synthesis (MS) is the current paradigm in evolutionary biology. It was actually built by expanding on the conceptual foundations laid out by its predecessors, Darwinism and neo-Darwinism. For sometime now there has been talk of a new Extended Evolutionary Synthesis (EES), and this article begins to outline why we may need such an extension, and how it may come about. As philosopher Karl Popper has noticed, the current evolutionary theory is a theory of genes, and we still lack (...) a theory of forms. The field began, in fact, as a theory of forms in Darwin’s days, and the major goal that an EES will aim for is a unification of our theories of genes and of forms. This may be achieved through an organic grafting of novel concepts onto the foundational structure of the MS, particularly evolvability, phenotypic plasticity, epigenetic inheritance, complexity theory, and the theory of evolution in highly dimensional adaptive landscapes. (shrink)

The reputations of scientists among their contemporaries depend not only on accomplishment, but also on interactions affected by influence and personality. The historical lore of most fields of scientific endeavor preserve these reputations, often through the identification of founders, innovators, and prolific workers whose contributions are considered fundamental to progress in the field. Historians frequently rely on the historical lore of scientists to guide their studies of the development of ideas, exhibiting justifiable caution in reassessing reputations in the light of (...) current knowledge. However, the transmission of historical lore can obscure the relative importance of accomplishment, influence and personality in shaping contemporary reputations, leaving the historian to either accept reputations at face value or attempt to reconstruct the context in which they were created. The science of taxonomy, because of its rules of priority, leaves a relatively accurate record of historical accomplishment through the persistence of taxa in catalogues and faunal guides. These records allow the modern historian an unbiased means to assess the relative accomplishments of historical figures and therefore a means to critically reassess reputations independent of personality and influence. In the historical lore of North American ichthyology, Louis Agassiz at Harvard and Spencer Baird at the Smithsonian emerge as central figures in the early development of the field during the mid-1800s, contributing not only through the quality and quantity of their science, but also through their roles as institutional leaders and mentors to workers who followed. Charles Girard, originally a student of Agassiz's and later a coworker with Baird, receives little notice in the history of ichthyology, and his reputation is that of a minor player in the initial description of the North American fish fauna, and one whose work appears to have been flawed or even careless when compared to his contemporaries. However, a review of both contemporary and modern taxonomic works reveals that Girard's productivity far exceeded that of either Agassiz or Baird. Furthermore, an examination of the tendency of Girard and his contemporaries to introduce synonymous names into the literature, which might reflect careless or uncritical work, suggests that Girard was among the more accomplished workers of his era, including Agassiz and Baird. Girard's low ranking in the folklore of North American ichthyology, therefore, can not be attributed to discernible shortcomings in his scientific work, but rather to a public and private campaign of criticism waged by Agassiz after Girard's departure from Harvard. While Agassiz's dispute with Girard stemmed from their personal interactions, he expressed them as criticisms of Girard's work, and thus helped shape Girard's scientific reputation as it has been transmitted through the lore of ichthyology. This case study reveals how scientific reputation may not always rest on accomplishment, but can be influenced by personal interactions obscured by time but nonetheless important to history. (shrink)

In 1920, Eugen Steinach and Paul Kammerer reported experiments showing that exposure to high temperatures altered the structure of the gonad and produced hyper-sexuality in "heat rats," presumably as a result of the increased production of sex hormones. Using Steinach's evidence that the gonad is a double gland with distinct sexual and generative functions, they used their findings to explain "racial" differences in the sexuality of indigenous tropical peoples and Europeans. The authors also reported that heat induced anatomical changes in (...) the interstitial cells of the gonad were inherited by the heat rats' descendants. Kammerer used this finding to link endocrinology to his long-standing interest in the inheritance of acquired characteristics. The heat rats supported his hypothesis that the interstitial cells of the double gland were the mechanism of somatic induction in the inheritance of acquired characteristics. The Steinach-Kammerer collaboration, Kammerer's use of Steinach's "puberty gland" to explain somatic induction, and his endocrine analysis of symbiosis reveal Paul Kammerer's late career attempt to integrate endocrinology and genetics with the political ideals of Austrian socialism. With them he developed a bioethics that challenged the growing reliance on race in eugenics and instead promoted cooperation over competition in evolution. I relate his attempt to the controversies surrounding the interstitial cells, to the status of extra-nuclear theories of heredity, and to Kammerer's commitment to Austromarxist social reforms during the interwar period. (shrink)

ve r since the appearance of Ontogeny and Phylogeny a decade ago, Stephen Jay Gould has continued to delight and inform a wide spectrum of readers and, in doing so, to defy C.P. Snow's lament about the "two cultures" of the sciences and the humanities. Gould's monthly column in Natural History magazine, published under the heading "This View of Life," has led to a series of highly praised volumes of essays—Ever Since Darwin (1977), The Panda's Thumb (1980), Hen's Teeth and (...) Horse's Toes (1983), and most recently The Flamingo's Smile (1985). In addition, Gould's The Mismeasure of Man (1981), which won the National Book Critics' Circle Award, analyzed the questionable character of intelligence testing and emphasized the many personal and cultural biases that have led researchers astray in this field. Given the sheer amount of Gould's publications, which include numerous scientific publications as well, Gould's readers have been kept busy indeed absorbing his prodigious output. (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)