Organisms' environments are thought to play a fundamental role in determining their fitness and hence in natural selection. Existing intuitive conceptions of environment are sufficient for biological practice. I argue, however, that attempts to produce a general characterization of fitness and natural selection are incomplete without the help of general conceptions of what conditions are included in the environment. Thus there is a "problem of the reference environment"—more particularly, problems of specifying principles which pick out those environmental conditions which determine (...) fitness. I distinguish various reference environment problems and propose solutions to some of them. While there has been a limited amount of work on problems concerning what I call "subenvironments", there appears to be no earlier work on problems of what I call the "whole environment". The first solution I propose for a whole environment problem specifies the overall environment for natural selection on a set of biological types present in a population over a specified period of time. The second specifies an environment relevant to extinction of types in a population; this kind of environment is especially relevant to certain kinds of long-term evolution. (shrink)

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)

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

In this paper, we develop an organizational account that defines biological functions as causal relations subject to closure in living systems, interpreted as the most typical example of organizationally closed and differentiated self-maintaining systems. We argue that this account adequately grounds the teleological and normative dimensions of functions in the current organization of a system, insofar as it provides an explanation for the existence of the function bearer and, at the same time, identifies in a non-arbitrary way the norms that (...) functions are supposed to obey. Accordingly, we suggest that the organizational account combines the etiological and dispositional perspectives in an integrated theoretical framework. IntroductionDispositional ApproachesEtiological TheoriesBiological Self-maintenance Closure, teleology, and normativityOrganizational differentiationFunctions C1: Contributing to the maintenance of the organization C2: Producing the functional trait Implications and Objections Functional versus useful Dysfunctions, side effects, and accidental contributionsProper functions and selected effectsReproductionRelation with other ‘unitarian’ approachesConclusions. (shrink)

Phenotype, whether conventional or extended, is defined as a reflectionof an underlying genotype. Adaptation and the natural selection thatfollows from it depends upon a progressively harmonious fit betweenphenotype and environment. There is in Richard Dawkins' notion ofthe extended phenotype a paradox that seems to undercut conventionalviews of adaptation, natural selection and adaptation. In a nutshell, ifthe phenotype includes an organism's environment, how then can theorganism adapt to itself? The paradox is resolvable through aphysiological, as opposed to a genetic, theory of (...) natural selection andadaptation. (shrink)

The word ‘environment’ has a history. Before the mid-nineteenth century, the idea of a singular, abstract entity—the organism—interacting with another singular, abstract entity—the environment—was virtually unknown. In this paper I trace how the idea of a plurality of external conditions or circumstances was replaced by the idea of a singular environment. The central figure behind this shift, at least in Anglo-American intellectual life, was the philosopher Herbert Spencer. I examine Spencer’s work from 1840 to 1855, demonstrating that he was exposed (...) to a variety of discussions of the ‘force of circumstances’ in this period, and was decisively influenced by the ideas of Auguste Comte in the years preceding the publication of Principles of psychology (1855). It is this latter work that popularized the word ‘environment’ and the corresponding idea of organism–environment interaction—an idea with important metaphysical and methodological implications. Spencer introduced into the English-speaking world one of our most enduring dichotomies: organism and environment. (shrink)

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)

Abstract Gene-selectionists define fundamental terms in non-standard ways. Genes are determinants of difference. Phenotypes are defined as a gene’s effects relative to some alternative whereas the environment is defined as all parts of the world that are shared by the alternatives being compared. Environments choose among phenotypes and thereby choose among genes. By this process, successful gene sequences become stores of information about what works in the environment. The strategic gene is defined as a set of gene tokens that combines (...) ‘actor’ tokens responsible for an effect with ‘recipient’ tokens whose replication is thereby enhanced. This set of tokens can extend across the boundaries of individual organisms, or other levels of selection, as these are traditionally defined. Content Type Journal Article Pages 1-19 DOI 10.1007/s10539-012-9315-5 Authors David Haig, Department of Organismic and Evolutionary Biology, Harvard University, 26 Oxford Street, Cambridge, MA 02138, USA Journal Biology and Philosophy Online ISSN 1572-8404 Print ISSN 0169-3867. (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)

Elsewhere, I defend the “causal interactionist population concept” (CIPC). Here I further defend the CIPC by showing how it clarifies another concept that biologists grapple with, namely, environment. Should we understand selection as ranging only over homogeneous environments or, alternatively, as ranging over any habitat area we choose to study? I argue instead that the boundaries of the population dictate the range of the environment, whether homogeneous or heterogeneous, over which selection operates. Thus, understanding the concept of population helps us (...) to understand concepts of selective environment, exemplifying the importance of the CIPC to other concepts and debates. (shrink)

The standard picture of evolution, is externalist: a causal arrow runs from environment to organism, and that arrow explains why organisms are as they are (Godfrey-Smith 1996). Natural selection allows a lineage to accommodate itself to the specifics of its environment. As the interior of Australia became hotter and drier, phenotypes changed in many lineages of plants and animals, so that those organisms came to suit the new conditions under which they lived. Odling-Smee, Laland and Feldman, building on the work (...) of Richard Lewontin, have shown that while sometimes appropriate, this is an inadequate conception of the relationship between organisms and the environments in which they live. Over time organisms alter their environment as well as being altered by their environments (Lewontin 1982; Lewontin 1983; Lewontin 1985). For example, animals modulate the effects of their physical and biological environment by building shelters: the beaver’s dam and lodge system, and termite mounds are two famous cases of animal structures, but they are few of many. There are many thousands of animals which make nests, burrows and other shelters. Likewise, animals make tools that give them access to resources from which they would otherwise be excluded: thus the Galapagos woodpecker finch uses a cactus needle to extract insects from crevasses in bark — insects that they would otherwise be unable to catch (Tebbich, Taborsky et al. 2001). Tool making is not as common as shelter-making, but it is common. For example many animals make traps: there are many species of pit-making antlions. Thus in part organisms make the world in which they live. They partially construct their own niches. Odling-Smee, Laland and Feldman argue that this has five major and under-appreciated consequences for biological theory. (shrink)

Since the discovery of the double helical structure of DNA, the standard account of the inheritance of features has been in terms of DNA-copying and DNA-transmission. This theory is just a version of the old theory according to which the inheritance of features is explained by the transfer at conception of some developmentally privileged material from parents to offspring. This paper does the following things: (1) it explains what the inheritance of features is; (2) it explains how the DNA-centric theory (...) emerged; (3) it clarifies the relation between the DNA-centric theory and the ‘unfolding’ theory of development; (4) it argues that (given what we now know about developmental processes and genetic activity) the DNA-centric theory should be abandoned in favour of a pluralistic (but not holistic) theory of the inheritance of features. According to this pluralistic theory, the reliable reoccurrence of phenotypes must be explained by appealing not only to processes responsible for the reliable reoccurrence of genetic developmental factors but also to processes responsible for the reliable reoccurrence (or persistence) of nongenetic developmental factors. (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)

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