According to the received view of evolution, only genes are inherited. From this view it follows that only genetically-caused phenotypic variation is selectable and, thereby, that all selection is at bottom genetic selection. This paper argues that the received view is wrong. In many species, there are intergenerationally-stable phenotypic differences due to environmental differences. Natural selection can act on these nongenetically-caused phenotypic differences in the same way it acts on genetically-caused phenotypic differences. Some selection is at bottom nongenetic selection. The (...) argument against the received view involves a reformulation of the concepts of inheritance and heritability. Inherited factors are all those developmental factors responsible for parent–offspring similarity; some inherited factors are genetic and some are not. Heritable variation is intergenerationally-stable phenotypic variation; some such variation is genetically-caused and some is not. The received view and its critics The possibility of nongenetic selection (the lucky butterfly) The reality of nongenetic selection 3.1 Imprinting mechanisms 3.2 Other learning mechanisms 3.3 Other nongenetic mechanisms Genetic and nongenetic inheritance mechanisms Genetic and nongenetic inherited factors Genetic and nongenetic heritability Conclusions + Current address: Dr Matteo Mameli, Research Fellow, King's College, Cambridge, CB2 1ST, United Kingdom, matteo.mameli{at}kings.cam.ac.uk' + u + '@' + d + ''//-->. (shrink)

Organic selection (the Baldwin Effect) by which an environmentally elicitedphenotypic adaptation comes under genotypic control following selectionwas proposed independently in 1896 by the psychologists James Baldwinand Conwy Lloyd Morgan and by the paleontologist Henry Fairfield Osborn.Modified forms of organic selection were proposed as autonomization bySchmalhausen in 1938, as genetic assimilation by Waddington in 1942, andas an explanation for evolution in changing environments or for speciationby Matsuda and West-Eberhard in the 1980s. Organic selection as amechanism mediating proximate environmental effects on the (...) evolution ofmorphology and behaviour is the topic of this essay. Discussion includesthe context in which organic selection was proposed, Lamarckian or neo-Lamarckian implications of organic selection, Waddingtons experimentalstudies demonstrating the existence and efficacy of genetic assimilation,stabilizing selection and norms of reaction favoured by Schmalhausen, andMatsudas search for a mechanism of organic selection in endocrine changesand in heterochrony. (shrink)

A functional approach to evolutionary morphology is emphasized in this paper. This perspective differs from the current structuralist trend, which emphasizes the constraining role of developmental paths. In addition, the present approach agrees with the adaptationist paradigm. It is further argued that three types of phenomena are better understood in this light: i.- the existence of evolutionary trends, ii.- the maintenance of certain structural features within a given taxon, and iii.- the irreversibility of evolution.

The longstanding species problem in biology has a history that suggests a solution, and that history is not the received history found in many texts written by biologists or philosophers. The notion of species as the division into subordinate groups of any generic predicate was the staple of logic from Aristotle through the middle ages until quite recently. However, the biological species concept during the same period was at first subtly and then overtly different. Unlike the logic sense, which relied (...) on definitions of the essence of both genus and species, biological species from the time of Epicurus were consistently considered to involve a reproductive element: in short, living species relied not on essential definitions, but on the generative cause, which might not be definable. I term this the generative conception of species: species were the generation by reproduction of form. This undercuts the claim that species before Darwin were essentialist, and divorces the notion of a type from that of essence. In fact, as late as the end of the nineteenth century, logicians explicitly treated biological “species” as a homonym only of logical essentialist species, and permitted considerable deviation from the type or form. At every point, species in logic were thought to be a subset, in effect, of some more general notion. I sketch a history of both philosophical and biological traditions of the species concept, before turning to the current conceptions. These are reconsidered in the light of this history, and in particular Mayr’s changing views are shown to be somewhat Whiggish, historiographically. Of the many touted biological species concepts, only one of which (Mayr’s) is called _the_ Biological Species Concept, none appears to capture all the relevant facts, intuitions, and operational requirements of biology. Cladistic conceptions, however, have much in common with the older philosophical literature, in that the natural group of cladism is the clade, or monophyletic group. After considering the Individuality Thesis, and the metaphysics of species, we see that species are the most particular terminal taxa in a clade, and that they are “defined” in terms of the particular synapomorphies, or evolved characters, that are causally responsible for keeping the lineages that organisms form distinct from one another. In this way, we can remain within the generative conception of species that has been in play for over two millennia, and yet avoid the pitfalls of prior attempts to find a universal conception of species. (shrink)

A crucial question for a process view of life is how to identify a process and how to follow it through time. The genidentity view can contribute decisively to this project. It says that the identity through time of an entity X is given by a well-identified series of continuous states of affairs. Genidentity helps address the problem of diachronic identity in the living world. This chapter describes the centrality of the concept of genidentity for David Hull and proposes an (...) extension of Hull’s view to the ubiquitous phenomenon of symbiosis. Finally, using immunology as a key example, it shows that the genidentity view suggests that the main interest of a process approach is epistemological rather than ontological and that its principal claim is one of priority, namely that processes precede and define things, and not vice versa. (shrink)

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

Recent developments toward a more holistic biology do not eliminate reductionism and determinism, but they do suggest more complex forms of them, in which there are multiple, interacting influences, as there are in complex or chaotic systems. Though there is a place in biology for both systemic and atomistic modes of explanation, for those with a theological perspective the shift to complex explanations in biology is often welcome. It suggests a more subtle view of divine action in which God's purposes (...) are affected through engagement with the complex systems of creation rather than by discrete interventions. It also invites us to connect the biological interdependence with the interdependence in the nature and purposes of God, and it is consonant with a mystical vision of the unity of all things. (shrink)

In both popular and technical discussion we often find the pairs of opposed terms ‘innate/acquired,’ ‘due to genes/due to environment,’ ‘nature/nurture,’ and so forth. They appear to be used as if they all captured a genuine distinction, and the same distinction at that. A related family of opposed pairs is held to describe the difference between those who attribute a certain trait to ‘nature’ and those who attribute it to ‘nurture’: ‘nativists’ versus ‘constructivists’ is one such pair. Chomsky and his (...) followers are often described as ‘nativists’ regarding certain features of language. On a cursory examination, many of the claims of Evolutionary Psychology bear the appearance of ‘nativism.’ That is, it looks as though Evolutionary Psychologists are making claims to the effect that many features of the mind which are often thought to be due to ‘nurture’ are in fact due to ‘nature.’ However, two of the genre's leading practitioners, John Tooby and Leda Cosmides, vigorously deny that this is their position. (shrink)

A survey of innateness in cognitive science, focusing on (1) what innateness might be, and (2) whether concepts might be innate.

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)

In his theory of evolution, Darwin recognized that the conditions of life play a role in the generation of hereditary variations, as well as in their selection. However, as evolutionary theory was developed further, heredity became identified with genetics, and variation was seen in terms of combinations of randomly generated gene mutations. We argue that this view is now changing, because it is clear that a notion of hereditary variation that is based solely on randomly varying genes that are unaffected (...) by developmental conditions is an inadequate basis for evolutionary theories. Such a view not only fails to provide satisfying explanations of many evolutionary phenomena, it also makes assumptions that are not consistent with the data that are emerging from disciplines ranging from molecular biology to cultural studies. These data show that the genome is far more responsive to the environment than previously thought, and that not all transmissible variation is underlain by genetic differences. In Evolution in Four Dimensions (2005) we identify four types of inheritance (genetic, epigenetic, behavioral, and symbol-based), each of which can provide variations on which natural selection will act. Some of these variations arise in response to developmental conditions, so there are Lamarckian aspects to evolution. We argue that a better insight into evolutionary processes will result from recognizing that transmitted variations that are not based on DNA differences have played a role. This is particularly true for understanding the evolution of human behavior, where all four dimensions of heredity have been important. (shrink)

Heyes argues that human metacognitive strategies (“cognitive gadgets” or “mills”) are the products of cultural evolution based on domain‐general cognition with few simple biases. Although like Heyes, we believe that the evolution of domain‐general cognitive processes played a crucial role in the evolution of human cognition, we argue that Heyes' distinction between mills and grist is too sharp, that associative learning evolved gradually to become more complex and hierarchical, something that is not captured by the system 1/system 2 distinction, and (...) that human cognitive plasticity required the genetic accommodation of gadget‐specific processes that led to more plasticity‐enhancing human‐specific biases. (shrink)

In 2011, Peterson suggested that the main reason why C.H. Waddington was essentially ignored by the framers of the modern evolutionary synthesis in the 1950s was because they were Cartesian reductionists and mathematical population geneticists while he was a Whiteheadian organicist and experimental geneticist who worked with Drosophila. This paper suggests a further reason that can only be seen now. The former defined genes and their alleles by their selectable phenotypes, essentially the Mendelian view, while Waddington defined a gene through (...) its functional role as determined by genetic analysis, a view that foresaw the modern view that a gene is a DNA sequence with some function. The former were interested in selection, while Waddington focused on variation. The differences between the two views of a gene are briefly considered in the context of systems biology. (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)

The claims are grounded in a wealth of fascinating data, particularly on primate and young child communication and social cognition, much produced by Tomasello’s own lab. But there is certainly no dearth of stimulating speculation. Tomasello’s story is rich and complex. In what follows, I focus on aspects of the three hypotheses listed above, offering some commentary as I go.

From the 1930s through the 1970s, C. H. Waddington attempted to reunite genetics, embryology, and evolution. One of the means to effect this synthesis was his model of the epigenetic landscape. This image originally recast genetic data in terms of embryological diagrams and was used to show the identity of genes and inducers and to suggest the similarities between embryological and genetic approaches to development. Later, the image became more complex and integrated gene activity and mutations. These revised epigenetic landscapes (...) presented an image of how mutations could alter developmental pathways to yield larger phenotypic changes. These diagrams became less important as the operon became used to model differential gene regulation. (shrink)

During the postwar period, the concept of instinct came to encapsulate the debate around the importance of nature versus nurture. The fact that animals show highly organized behavior early in development suggested the presence of an underlying fixity where behavior was “inbuilt” into an animal’s biology despite an individual’s experiences. This placed a discrete and exhaustive line between the innate and acquired that became a foundation for the European-dominated field of ethology. Across the Atlantic, a group of comparative psychologists led (...) by the American Museum of Natural History’s T. C. Schneirla contested this approach, proposing that the study of animal behavior should avoid abstract dichotomies with a renewed focus on developmental processes. While Schneirla’s theoretical and empirical work shaped the modern study of animal behavior, his legacy requires revisiting in an era where the nature versus nurture debate is regaining prominence. In this article, I revisit Schneirla’s approach to behavior with a focus on his paper “Interrelationships of the ‘Innate’ and the ‘Acquired’ in Instinctive Behavior” (published in M. Autuori et al. (1956) _L’instinct dans le comportement des animaux et de l’homme_; Masson, Paris, pp. 387–452) for the journal’s “Classics in Biological Theory” collection; the paper is available as supplementary material in the online version of this article. A companion article (this issue; G. M. Kohn (2024) “A Discussion on Instinct, Paris, 1954”) presents the commentary that was published with it. (shrink)

In this paper, we propose the expression cognitive twists for cognitive mechanisms that result from the coevolution of genes and learning. Evidence is available that at least some cultural learning mechanisms, such as imitation and language, have evolved genetically under the pressure produced by culture, even though they are mostly acquired through domain-general learning during development. Although the existence of these mechanisms is consistent with evolutionary theory, their importance has not been sufficiently emphasized by mind-centered accounts of human cognitive evolution, (...) namely evolutionary psychology and cultural evolutionary psychology. We provide concrete examples of cognitive twists, such as vocal imitation. Genetic changes in action-perception matching circuits suggest that human imitation and perhaps language are cognitive twists, namely plastic, learnable, yet genetically evolved cognitive mechanisms. We conclude that cognitive twists depict plausible evolutionary scenarios for the evolution of cognition in Homo sapiens. (shrink)

This article develops a media philosophical framework for addressing the intersection of epigenetics and complex dynamical systems in theoretical biology. In particular, it argues that the theoretical humanities need to think critically about the computability of epigenomic regulation, as well as speculatively about the possibility of an epigenomics beyond complexity. The fact that such a conceptual framework does not exist suggests not only a failure to engage with the mathematics of complexity, but also a failure to engage with its history. (...) Both epigenetics and the application of complex dynamical systems to biology originate in the mid-twentieth-century work of Conrad Hal Waddington. The article demonstrates how this genealogy of epigenetic complexity reveals the need for a speculative conception of epigenomic mediation beyond mathematical complexity. Using Whitehead’s philosophy as a conceptual anchor, the “epimedial landscape” serves as a first pass at a media philosophical revision of the complex dynamical systems view of the epigenetic landscape. (shrink)

Although there are many historical and philosophical analyses of evolutionary developmental biology (EvoDevo), its development in the 1980s, when many individual or collective attempts to synthesize evolution and development were made, has not been examined in detail. This article focuses on some interdisciplinary studies during the 1980s and argues that they had important characteristics that previous historical and philosophical work has not recognized. First, we clarify how each set of studies from the 1980s integrated the results or approaches from different (...) biological fields, such as paleontology, developmental genetics, comparative morphology, experimental embryology, theoretical developmental biology, and population genetics. Second, after close examination we show that the interdisciplinary studies during the 1980s adopted different and conflicting views of genes, such as developmental-genetic, epigenetic, or population-genetic ones. We conclude that EvoDevo in the 1980s was a motley aggregation of various kinds of local integration. Finally, we discuss the implications of our analysis by comparing these early EvoDevo studies with those of the Modern Synthesis and with the present state of EvoDevo. (shrink)

C.H. Waddington is today remembered chiefly as a Drosophila developmental geneticist who developed the concepts of “canalization” and “the epigenetic landscape.” In his lifetime, however, he was widely perceived primarily as a critic of Neo-Darwinian evolutionary theory. His criticisms of Neo-Darwinian evolutionary theory were focused on what he saw as unrealistic, “atomistic” models of both gene selection and trait evolution. In particular, he felt that the Neo-Darwinians badly neglected the phenomenon of extensive gene interactions and that the “randomness” of mutational (...) effects, posited in the theory, was a false postulate. This last criticism dealt with the phenomenon known today as “developmental constraints.” Although population genetics itself has evolved considerably from its form at mid-20th century, much of Waddington’s critique, it is argued here, retains cogency. Yet, he did not attempt to develop a full-fledged alternative theory himself. Perhaps most surprisingly, in retrospect, is that he did not try to marry his work on gene interactions in development with his evolutionary interests, to create a theory of “genetic networks” and their evolution. Whether evolutionary genetics today will incorporate “network thinking” as a central element or whether there will be a general retreat to the more atomistic approach offered by genomics, remains an open question. (shrink)

Recently a growing number of biologists have begun to consider the causal role that processes of embryonic development may play in evolution. This constitutes a reunion of these phenomena which had been linked in the nineteenth century through Haeckel's biogenetic law. This reunion may result in a new subdiscipline of biology, if there is a set of unique concepts and methods which tie the various research approaches together. Such concepts as bauplan, canalization, and developmental constraint, may serve in such a (...) capacity. The methods employed must combine comparative and experimental analyses, with special attention paid to the range of variation in developmental events within each taxon. These concepts and methods are also applicable to the problem of how development evolved, thus the evolution of development may also be considered part of the reunion. The reunion is discussed in terms of the potential participation of various schools of thought found in the current literature. (shrink)

In this paper, I develop a naturalistic conception of evolutionary progress. I argue that the Waddingtonian notion of adaptability can be embedded meaningfully into a framework which views living things as nonequilibrium structures. This thermodynamic interpretation places great emphasis on the dynamics of environmental change, whereas the classical conceptions are based on equilibrium conceptions of the evolutionary process. What improves in evolution is the ability of living things to stay alive in increasingly heterogeneous environments.

The concept of innateness is a part of folk wisdom but is also used by biologists and cognitive scientists. This concept has a legitimate role to play in science only if the colloquial usage relates to a coherent body of evidence. We examine many different candidates for the post of scientific successor of the folk concept of innateness. We argue that none of these candidates is entirely satisfactory. Some of the candidates are more interesting and useful than others, but the (...) interesting candidates are not equivalent to each other and the empirical and evidential relations between them are far from clear. Researchers have treated the various scientific notions that capture some aspect of the folk concept of innateness as equivalent to each other or at least as tracking properties that are strongly correlated with each other. But whether these correlations exist is an empirical issue. This empirical issue has not been thoroughly investigated because in the attempt to create a bridge between the folk view and their theories, researchers have often assumed that the properties must somehow cluster. Rather than making further attempts to import the folk concept of innateness into the sciences, efforts should now be made to focus on the empirical questions raised by the debates and pave the way to a better way of studying the development of living organisms. Such empirical questions must be answered before it can be decided whether a good scientific successor – in the form of a concept that refers to a collection of biologically significant properties that tend to co-occur – can be identified or whether the concept of innateness deserves no place in science. (shrink)

The aim of this contribution is to investigate certain selected parts of the extended evolutionary synthesis which all have a common denominator, namely evolution by meaning attribution. We start by arguing that living organisms can manipulate and interpret their genetic script via epigenetic modifications in a semiotic manner, that is, by meaning attribution. Genes do not build living beings to be transmitted to future generations. Genes have been shaped by evolution as a memory medium that is transmitted from one generation (...) to the next, but the actual reading of such scripts is modified by momentary contexts. Secondly, we show that phenotypic evolution variously re-uses already existing homologies which in evolving systems acquire a new meaning. We also suggest that the ways in which organisms perceive their environment and other living beings is an important but still largely neglected evolutionary force. Variations in perception influence the direction and intensity of sexual selection and some behaviourally mediated regimes of natural selection. Thirdly, we point out that especially if we want to study their evolution, living beings should not be considered in isolation but in their mutual coexistence, in their historically established being together. Recent attempts to view living beings as constructors of niches and holobionts seem compatible with the classical Umwelt theory. This approach seems capable of accounting for both competitiveness and cooperation, thus making the overall image of evolution more comprehensive. And finally, we argue that if we want to expand our understanding of biological evolution, in addition to variation, selection, and inheritance we also need to take into account processes which participate in meaning attribution. (shrink)

The “Baldwin effect” occurs when natural selection supplements adaptive phenotypic changes acquired during an organism’s lifetime. This effect, first described in 1896, is now usually considered to be rare and insignificant, and is not part of the explanatory repertoire of most evolutionary scientists. Arguments against it are examined here and found to be inadequate, either because they misunderstand the effect, or confuse it with Lamarckism, or fail to perceive it even while describing it. The article proposes examples showing that the (...) Baldwin effect is far more prevalent and powerful than even Baldwin’s few supporters realize. Evolved features of the human mind are described that make this effect unusually difficult to perceive. Some of these mental features are themselves Baldwin effects. Natural selection’s supporters, and its skeptics, appear to disregard the Baldwin effect for different reasons. The article proposes that acceptance of the Baldwin effect can significantly improve the study of human nature, as practiced by both sides in the nature-nurture controversy, and can even provide a bridge to end this controversy. (shrink)

The traditional practice of establishing morphological types and investigating morphological organization has found new support from evolutionary developmental biology (evo-devo), especially with respect to the notion of body plans. Despite recurring claims that typology is at odds with evolutionary thinking, evo-devo offers mechanistic explanations of the evolutionary origin, transformation, and evolvability of morphological organization. In parallel, philosophers have developed non-essentialist conceptions of natural kinds that permit kinds to exhibit variation and undergo change. This not only facilitates a construal of species (...) and higher taxa as natural kinds, but also broadens our perspective on the diversity of kinds found in biology. There are many different natural kinds relevant to the investigative and explanatory aims of evo-devo, including homologues and developmental modules. (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)

We claim that much of the confusion associated with the "tautology problem" about survival of the fittest is due to the mistake of attributing fitness to individuals instead of to types. We argue further that the problem itself cannot be solved merely by taking fitness as the aggregate cause of reproductive success. We suggest that a satisfying explanation must center not on logical analysis of the concept of general adaptedness but on the empirical analysis of single adapted traits and their (...) causal relationship to changes in allele frequencies. (shrink)

A discussion is going on in cognitive science about the use of representations to explain how intelligent behavior is generated. In the traditional view, an organism is thought to incorporate representations. These provide an internal model that is used by the organism to instruct the motor apparatus so that the adaptive and anticipatory characteristics of behavior come about. So-called interactionists claim that this representational specification of behavior raises more problems than it solves. In their view, the notion of internal representational (...) models is to be dispensed with. Instead, behavior is to be explained as the intricate interaction between an embodied organism and the specific make up of an environment. The problem with a non-representational interactive account is that it has severe difficulties with anticipatory, future oriented behavior. The present paper extends the interactionist conceptual framework by drawing on ideas derived from the study of morphogenesis. This extended interactionist framework is based on an analysis of anticipatory behavior as a process which involves multiple spatio-temporal scales of neural, bodily and environmental dynamics. This extended conceptual framework provides the outlines for an explanation of anticipatory behavior without involving a representational specification of future goal states. (shrink)