Barely a decade after the discovery of the chromosomal basis of inheritance, and the articulation of the genetical theory of population change, the gene came to be widely regarded as the fundamental unit of biological organization. This is hardly surprising. The gene concept is a powerful one; it plays a unifying role in our understanding of evolution. Darwin told us that evolution by natural selection occurs in a population when organisms survive, die and reproduce differentially on account of their heritable (...) form. This is a very schematic theory. It requires an account of the process of inheritance and also an account of the generation of phenotype. The gene concept plays a prominent role in explaining, and uniting, these phenomena. Genes are the units of inheritance; they are passed from parents to offspring in reproduction. Moreover, they are seen as units of phenotypic control. Evolutionary biologists often speak of the genome as a program for the production of an organism. Genes also became the elements of which populations are composed. Our best theory of population dynamics—inherited from Fisher, Haldane, and Wright—is a theory of changes in the relative frequencies of gene types. Genes are not just the principal causes of evolutionary change, they are also the units over which evolutionary change is defined and measured. So, at least, the orthodox reading of the Modern Synthesis theory of evolution would have us believe... (shrink)

There is ongoing controversy as to whether the genome is a representing system. Although it is widely recognised that DNA carries information, both correlating with and coding for various outcomes, neither of these implies that the genome has semantic properties like correctness or satisfaction conditions, In the Scope of Logic, Methodology, and the Philosophy of Sciences, Vol. II. Kluwer, Dordrecht, pp. 387–400). Here a modified version of teleosemantics is applied to the genome to show that it does indeed have semantic (...) properties – there is representation in the genome. The account differs in three respects from previous attempts to apply teleosemantics to genes. It emphasises the role of the consumer of representations. It rejects the standard assumption that genetic representation can be used to explain the course of an organism’s development. And it identifies the explanatory role played by representational properties of the genome. A striking consequence of this account is that other inheritance systems could also be representational. Thus, a version of the parity thesis is accepted. However, the criteria for being an inheritance system are demanding, so semantic properties are not ubiquitous. (shrink)

The current debate over extending inheritance and its evolutionary impact has focused on adding new categories of non-genetic factors to the classical transmission of DNA, and on trying to redefine inheritance. Transmitted factors have been mainly characterized by their directions of transmission and the way they store variations. In this paper, we leave aside the issue of defining inheritance. We rather try to build an evolutionary conceptual framework that allows for tracing most, if not all forms of transmission and makes (...) sense of their different tempos and modes. We discuss three key distinctions that should in particular be the targets of theoretical and empirical investigation, and try to assess the interplay among them and evolutionary dynamics. We distinguish two channels of transmission, two measurements of the temporal dynamics of transmission, respectively across and within generations, and two types of transmitted factors according to their evolutionary relevance. By implementing these three distinctions we can then map different forms of transmission over a continuous space describing the combination of their varying dynamical features. While our aim is not to provide yet another model of inheritance, putting together these distinctions and crossing them, we manage to offer an inclusive conceptual framework of transmission, grounded in empirical observation, and coherent with evolutionary theory. This interestingly opens possibilities for qualitative and quantitative analyses, and is a necessary step, we argue, in order to question the interplay between the dynamics of evolution and the dynamics of multiple forms of transmission. (shrink)

The paper outlines the contours of an organizational perspective on extended inheritance. Based on theoretical studies about biological organization and extended physiology, this perspective allows for the conception of extended biological legacies while keeping a theoretically indispensable distinction between biological systems and their environment. In this context, the line of demarcation between these systems and their surroundings is modelled on an organizational criterion and on the related conceptual distinction between organizational constraints, whose specific role is to harness flows of matter (...) and energy across generations of composite biological systems, and resources exploited by those systems. Biological legacies are restricted to persisting constitutive elements responsible for the reoccurrence of organizational constraints in a given environment. The case of symbiotic transmission is presented as a paradigmatic system illustrating the main proposed conceptual clarifications. (shrink)

The concept of biological inheritance has recently been extended so as to integrate, among other elements, parts of organisms’ environments. The literature refers to the trans-generational reconstruction of these parts in terms of environmental or ecological inheritance. This article’s main objective is to clarify the different meanings of "environmental inheritance," to underline so far unnoticed theoretical difficulties associated to this polysemous notion and to consequently argue that inheritance, even when extended, should be theoretically distinguished from trans-generational environmental stability. After disentangling (...) the different meanings of environmental inheritance, I underline that studies dealing with this concept place themselves in the wake of earlier contributions about biological environment and elaborate on the role of organisms in the determination of their relevant developmental and selective surroundings. This leads me to question the legitimacy of the category shift operated by niche inheritance proponents—from environment to inheritance—and to explain why the very concept of inherited environment shows important and so far unnoticed theoretical limitations. In this context, I assert the necessity to distinguish two related but different research programs: the construction of a finer-grained theory of environment and the elaboration of an extended theory of inheritance. (shrink)

In this paper, I argue that the increasing data about non-genetic inheritance requires the construction of a new conceptual framework that should complement the inclusive approaches already discussed in the literature. More precisely, I hold that this framework should be epistemologically relevant for evolutionary biologists in capturing the limits of extended inheritance and in reassessing the boundaries of biological systems that transmit traits to their offspring. I outline the first elements of an organizational account of extended inheritance. In this account, (...) the category of inherited factors is neither restricted to genes nor extended to stable resources related to trans-generational similarities. Instead, it includes persisting constitutive elements appearing as difference makers for heterogeneous organizational constraints, namely for heterogeneous constitutive parts whose specific role is to harness flows of matter and energy across generations of clearly delimited extended organized systems. This both inclusive and restrictive framework opens an additional way to apprehend how extended inheritance may affect evolutionary trajectories. (shrink)

The concept of biological inheritance has recently been extended so as to integrate, among other elements, parts of organisms’ environments. The literature refers to the trans-generational reconstruction of these parts in terms of environmental or ecological inheritance. This article’s main objective is to clarify the different meanings of "environmental inheritance," to underline so far unnoticed theoretical difficulties associated to this polysemous notion and to consequently argue that inheritance, even when extended, should be theoretically distinguished from trans-generational environmental stability. After disentangling (...) the different meanings of environmental inheritance, I underline that studies dealing with this concept place themselves in the wake of earlier contributions about biological environment and elaborate on the role of organisms in the determination of their relevant developmental and selective surroundings. This leads me to question the legitimacy of the category shift operated by niche inheritance proponents—from environment to inheritance—and to explain why the very concept of inherited environment shows important and so far unnoticed theoretical limitations. In this context, I assert the necessity to distinguish two related but different research programs: the construction of a finer-grained theory of environment and the elaboration of an extended theory of inheritance. (shrink)

In this paper I use a case study—the discovery of the chaperon function exerted by proteins in the various steps of the hereditary process—to re-discuss the question whether the nucleic acids are the sole repositories of relevant information as assumed in the information theory of heredity. The evidence I here present of a crucial role for molecular chaperones in the folding of nascent proteins, as well as in DNA duplication, RNA folding and gene control, suggests that the family of proteins (...) acting as molecular chaperones provides information that is complementary to that stored in the nucleic acids, and equally important. A re-evaluation of the role of proteins in the hereditary process is in order away from the gene-centric approach of the information theory of heredity, to which neo-Darwinian evolutionists adhere. (shrink)

We develop a conceptual framework that connects biological heredity and organization. We refer to heredity as the cross-generation conservation of functional elements, defined as constraints subject to organizational closure. While hereditary objects are functional constituents of biological systems, any other entity that is stable across generations—and possibly involved in the recurrence of phenotypes—belongs to their environment. The central outcome of the organizational perspective consists in extending the scope of heredity beyond the genetic domain without merging it with the broad category (...) of cross-generation stability. After discussing some implications, we conclude with a reflection on the relationship between stability and variation. 1Introduction2From Extended Heredity to Cross-generation Stability 2.1Extending the scope of heredity: A brief state of the art2.2Rethinking heredity: Conceptual challenges3Biological Heredity in Light of Organization 3.1Biological organization within organisms and beyond3.2Extending organization in time3.3What is biological heredity?4Implications and Objections 4.1Heredity as a specific kind of cross-generation stability4.2Heredity at various levels of description4.3Non-functional and dysfunctional objects5Conclusions: From Conservation to Variation. (shrink)

In recent years, a proliferation of books about empathy, cooperation and pro-social behaviours has significantly influenced the discourse of the life-sciences and reversed consolidated views of nature as a place only for competition and aggression. In this article I describe the recent contribution of three disciplines – moral psychology, primatology and the neuroscience of morality – to the present transformation of biology and evolution into direct sources of moral phenomena, a process here named the ‘moralization of biology’. I conclude by (...) addressing the ambivalent status of this constellation of authors, for whom today ‘morality comes naturally’: I explore both the attractiveness of their message, and the problematic epistemological assumptions of their research programmes in the light of new discoveries in developmental and molecular biology. (shrink)

In recent years, a proliferation of books about empathy, cooperation and pro-social behaviours (Brooks, 2011a) has significantly influenced the discourse of the life-sciences and reversed consolidated views of nature as a place only for competition and aggression. In this article I describe the recent contribution of three disciplines – moral psychology (Jonathan Haidt), primatology (Frans de Waal) and the neuroscience of morality – to the present transformation of biology and evolution into direct sources of moral phenomena, a process here named (...) the ‘moralization of biology’. I conclude by addressing the ambivalent status of this constellation of authors, for whom today ‘morality comes naturally’: I explore both the attractiveness of their message, and the problematic epistemological assumptions of their research programmes in the light of new discoveries in developmental and molecular biology. (shrink)

Gary Marcus has written a very interesting book about mental development from a nativist perspective. For the general readership at which the book is largely aimed, it will be interesting because of its many informative examples of the development of cognitive structures and because of its illuminating explanations of ways in which genes can contribute to these developmental processes. However, the book is also interesting from a theoretical point of view. Marcus tries to make nativism compatible with the central arguments (...) that anti-nativists use to attack nativism and with many recent discoveries about genetic activity and brain development. In so doing, he reconfigures the nativist position to a considerable extent. (shrink)

(2) There is significant cultural variation in the way people reason, categorize, and react to various aspects of the world. A proper understanding of such variation has implications for theories about human nature – and cognitive architecture – and its malleability. In turn, these theories have implications for theories about the status and generalisability of psychological explanations (see Nisbett 2003), for theories about the extent to which social engineering and social reform is possible (see Singer 2000), etc.

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)

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

The presence of various mechanisms of non-genetic inheritance is one of the main problems for current evolutionary theory according to several critics. Sufficient empirical and conceptual reasons exist to take this claim seriously, but there is little consensus on the implications of multiple inheritance systems for evolutionary processes. Here we use the Price Equation as a starting point for a discussion of the differences between four recently proposed categories of inheritance systems; genetic, epigenetic, behavioral and symbolic. Specifically, we address how (...) the components of the Price Equation encompass different non-genetic systems of inheritance in an attempt to clarify how the different systems are conceptually related. We conclude that the four classes of inheritance systems do not form distinct clusters with respect to their effect on the rate and direction of phenotypic change from one generation to the next in the absence or presence of selection. Instead, our analyses suggest that different inheritance systems can share features that are conceptually very similar, but that their implications for adaptive evolution nevertheless differ substantially as a result of differences in their ability to couple selection and inheritance. (shrink)

For a population to undergo evolution by natural selection, it is assumed that the constituents of the population form parent-offspring lineages, that is, that they must reproduce. I challenge this assumption by dividing the notion of reproduction into two subprocesses, that is, multiplication and inheritance, that produce parent-offspring lineages between the parts of a population, and I show that their population-level roles, generation and memory, respectively, can be effected by processes that do not rely on such local-level lineages. I further (...) argue that these two population-level processes, not local parent-offspring lineages, are necessary conditions for a population to undergo Darwinian evolution. (shrink)

Two problems related to the biological identity of living beings are faced: the who-problem (which are the biological properties making that living being unique and different from the others?); the persistence-problem (what does it take for a living being to persist from a time to another?). They are discussed inside a molecular biology framework, which shows how epigenetics can be a good ground to provide plausible answers. That is, we propose an empirical solution to the who-problem and to the persistence-problem (...) on the basis of the new perspectives opened by a molecular understanding of epigenetic processes. In particular, concerning the former, we argue that any living being is the result of the epigenetic processes that have regulated the expression of its genome; concerning the latter, we defend the idea that the criterion for the persistence of its identity is to be indicated in the continuity of those epigenetic processes. We also counteract possible objections, in particular (1) whether our approach has something to say at a metaphysical level; (2) how it could account for the passage from the two phenotypes of the parental gametes to the single phenotype of the zygote; (3) how it could account for the identity of derivatives of one living being that continue to live disjoined from that original living being; (4) how it could account for higher mental functions. (shrink)

In this paper, I will conduct three interrelated analyses. First, I will develop an analysis of various concepts in the history of biology that used to refer to individual-level phenomena but were then reinterpreted by the Modern Synthesis in terms of populations. Second, I argue that a similar situation can be found in contemporary biological theory. While different approaches reflect on the causal role of developing organisms in evolution, proponents of the Modern Synthesis avoid any substantial change by reinterpreting and (...) explaining individual-level phenomena from a population perspective. Finally, I will approach this debate by advocating the statistical reading of natural selection, which holds that explanations by natural selection are statistical. I will argue that the above historical conceptual reinterpretations belong to a new explanatory strategy developed by the Modern Synthesis based on population thinking. The reinterpretation of concepts at the individual level is part of the explanatory framework of the Modern Synthesis and the empty role of development within this framework. Moreover, the statistical perspective adopted here allows for the integration of two explanatory models: population-statistical and individual-causal. Finally, I will argue that this pluralistic framework can help to define the explanatory scope of the different biological approaches in order to achieve a coherent integration of development into evolution without rejecting population thinking. (shrink)