This article analyzes the evolution of the land ethic re-presented by J. Baird Callicott over the last two decades under pressure from the charge of misanthropy and ecofascism. It also traces the development of Callicott?s own ethical system, and examines its most current phase both in itself and in relation to his other theoretical commitments, including his particular version of moral monism, and his communitarian critique of egalitarianism. It concludes that Callicott?s communitarianism is by itself insufficient to fund an adequate (...) environmental ethic, and that for the sake of self-consistency he should either discard his moral monism or else further revise his ethical system. (shrink)

Adaptation by means of natural selection depends on the ability of populations to maintain variation in heritable traits. According to the Modern Synthesis this variation is sustained by mutations and genetic drift. Epigenetics, evodevo, niche construction and cultural factors have more recently been shown to contribute to heritable variation, however, leading an increasing number of biologists to call for an extended view of speciation and evolution. An additional common feature across the animal kingdom is learning, defined as the ability to (...) change behavior according to novel experiences or skills. Learning constitutes an additional source for phenotypic variation, and change in behavior may induce long lasting shifts in fitness, and hence favor evolutionary novelties. Based on published studies, I demonstrate how learning about food, mate choice and habitats has contributed substantially to speciation in the canonical story of Darwin’s finches on the Galapagos Islands. Learning cannot be reduced to genetics, because it demands decisions, which requires a subject. Evolutionary novelties may hence emerge both from shifts in allelic frequencies and from shifts in learned, subject driven behavior. The existence of two principally different sources of variation also prevents the Modern Synthesis from self-referring explanations. (shrink)

The question of whether there are laws of nature in ecology has developed substantially in the last 20 years. Many have attempted to rehabilitate ecology’s lawlike status through establishing that ecology possesses laws that robustly appear across many different ecological systems. I argue that there is still something missing, which explains why so many have been skeptical of ecology’s lawlike status. Community ecology has struggled to establish what I call a General Unificatory Theory. The lack of a GUT causes problems (...) for explanation as there are no guidelines for how to integrate the lower-level mathematical and causal models into a larger theory of how ecological assemblages are formed. I turn to a promising modern attempt to provide a unified higher-level explanation in ecology, presented by ecologist Mark Vellend, and advocate for philosophical engagement with its prospects for aiding ecological explanation. (shrink)

The neutral theory of biodiversity purports that patterns in the distribution and abundance of species do not depend on adaptive differences between species (i.e. niche differentiation) but solely on random fluctuations in population size (“ecological drift”), along with dispersal and speciation. In this framework, the ultimate driver of biodiversity is speciation. However, the original neutral theory made strongly simplifying assumptions about the mechanisms of speciation, which has led to some clearly unrealistic predictions. In response, several recent studies have combined neutral (...) community models with more elaborate speciation models. These efforts have alleviated some of the problems of the earlier approaches, while confirming the general ability of neutral theory to predict empirical patterns of biodiversity. However, the models also show that the mode of speciation can have a strong impact on relative species abundances. Future work should compare these results to diversity patterns arising from non‐neutral modes of speciation, such as adaptive radiations. (shrink)

The dynamics of single populations up to ecosystems, are often described by one or a set of non-linear ordinary differential equations. In this paper we review the use of bifurcation theory to analyse these non-linear dynamical systems. Bifurcation analysis gives regimes in the parameter space with quantitatively different asymptotic dynamic behaviour of the system. In small-scale systems the underlying models for the populations and their interaction are simple Lotka-Volterra models or more elaborated models with more biological detail. The latter ones (...) are more difficult to analyse, especially when the number of populations is large. Therefore for large-scale systems the Lotka-Volterra equations are still popular despite the limited realism. Various approaches are discussed in which the different time-scale of ecological and evolutionary biological processes are considered together. (shrink)

In the first part ("Determination"), I consider different notions of determination, contrast and compare modal with non-modal accounts and then defend two a-modality theses concerning essence and supervenience. I argue, first, that essence is a a-modal notion, i.e. not usefully analysed in terms of metaphysical modality, and then, contra Kit Fine, that essential properties can be exemplified contingently. I argue, second, that supervenience is also an a-modal notion, and that it should be analysed in terms of constitution relations between properties. (...) In the second part ("A Theory of Truthmaking"), I first review the literature on ontological commitment, and argue that the notion of truthmaking is better suited to play its explanatory rôle. I then argue that we should take truthmaker theory seriously, and that we should provide actual truthmakers for all truths there are. In the last chapter of this part, I review Armstrong's truthmaker theories and argue that they are unsatisfactory. I generalise my criticism to an argument against truthmaker necessitarianism, the view that truthmakers necessarily make true the truths they are truthmakers of. In the third part ("Properties and their Kind(s)"), I discuss qualitative determination. I present and endorse the truthmaker argument for universals, and defend universalism against friends of tropes, states of affairs and facts. I then give a novel characterisation of the important class of intrinsic properties, building on Lewis' work. With a workable notion of intrinsicness at hand, I argue that relations are ontologically – but not "ideologically" – dispensable: qualitative determination in general is a matter of intrinsic structure. In the future, I plan to add two parts and to publish my thesis as a book: In the fourth part ("Exemplification"), I argue for the existence of an exemplification relation tying universals and particulars together. I derive theoretical benefits from this relation by providing adverbialist theories of modality and tense and identify exemplification with a type of parthood. In the fifth part ("Qua qua qua"), I investigate the curious and interesting ontological category of so-called "qua-objects" (Picasso-as-a-painter, for example) and argue that they exist, are not identical with transworld indidivuals or modal parts and that they provide (contingent) truthmakers for all true predications. (shrink)

The world’s leading environmental advisory institutions look to ecological theory and research as an objective guide for policy and resource management decision-making. In addition to various theoretical merits of doing so, it is therefore crucially important to clear up confusions about ecology’s conceptual foundations and to make plain the basic workings of inferential methods used in the science. Through discussion of key moments in the genesis of the theoretical branch of ecology, this essay elucidates a general heuristic role of teleological (...) metaphor in ecological research and defuses certain enduring confusions and misguided criticisms of current work in ecology. (shrink)

In view of the confusion about the use of the term ‘population’ in biology, an analysis has been made of the use of the concept of population in biology. Origin and development, logic and epistemology of the population-concept have been investigated for that purpose. It appears that several concepts of population coexist in biology. The term ‘population’ is used for all these concepts, so it is not amazing that confusion has arisen. This confusion is the more dangerous, since ‘population’ plays (...) an important part in the theory of evolution. A modus has been suggested to diminish this confusion by defining ‘population’ in a very wide sense of the term and by proposing a ‘deme’—terminology to specify the various special concepts of population. (shrink)

Besides mechanistic explanations of phenomena, which have been seriously investigated in the last decade, biology and ecology also include explanations that pinpoint specific mathematical properties as explanatory of the explanandum under focus. Among these structural explanations, one finds topological explanations, and recent science pervasively relies on them. This reliance is especially due to the necessity to model large sets of data with no practical possibility to track the proper activities of all the numerous entities. The paper first defines topological explanations (...) and then explains why topological explanations and mechanisms are different in principle. Then it shows that they are pervasive both in the study of networks—whose importance has been increasingly acknowledged at each level of the biological hierarchy—and in contexts where the notion of selective neutrality is crucial; this allows me to capture the difference between mechanisms and topological explanations in terms of practical modelling practices. The rest of the paper investigates how in practice mechanisms and topologies are combined. They may be articulated in theoretical structures and explanatory strategies, first through a relation of constraint, second in interlevel theories, or they may condition each other. Finally, I explore how a particular model can integrate mechanistic informations, by focusing on the recent practice of merging networks in ecology and its consequences upon multiscale modelling. (shrink)

Terminology within the biological sciences gets its import not just from semantic meaning, but also from the way it functions within the rhetorics of the various disciplinary practices. The ‘sociobiology’ of human behavior inherits three distinct rhetorics from the genetic disciplines. Sociobiologists use population genetic, biometrical genetic, and molecular genetic rhetorics, without acknowledging the conceptual and experimental constraints that are assumed by geneticists. The eclectic blending of these three rhetorics obscures important differences of context and meaning. Sociobiologists use foundational terms (...) in genetics, such as ‘gene’, ‘fitness’, ‘evolution’, ‘heritability’, ‘trait’ and ‘polygenic inheritance’, in starkly different ways from geneticists, while basing their analysis of human behavior on the implied authority of genetics. As a free‐floating ‘gene talk’ moves across different disciplinary contexts, and before different audiences, it takes the form of an over‐simplified and misleading arch‐determinism. The result is widespread application of vague, incomplete, and distorted biological theory. If most sociobiologists, do not deliberately promote biological determinism, still less a political agenda, there is ample evidence that they misconstrue the implications of the genetic language that they borrow. (shrink)

The Swiss-born medical researcher Karl Friedrich Meyer is best known as a ‘microbe hunter’ who pioneered investigations into diseases at the intersection of animal and human health in California in the 1920s and 1930s. In particular, historians have singled out Meyer’s 1931 Ludwig Hektoen Lecture in which he described the animal kingdom as a ‘reservoir of disease’ as a forerunner of ‘one medicine’ approaches to emerging zoonoses. In so doing, however, historians risk overlooking Meyer’s other intellectual contributions. Developed in a (...) series of papers from the mid-1930s onwards, these were ordered around the concept of latent infections and sought to link microbial behavior to broader bio-ecological, environmental, and social factors that impact hostpathogen interactions. In this respect Meyer—like the comparative pathologist Theobald Smith and the immunologist Frank Macfarlane Burnet—can be seen as a pioneer of modern ideas of disease ecology. However, while Burnet’s and Smith’s contributions to this scientific field have been widely acknowledged, Meyer’s have been largely ignored. Drawing on Meyer’s published writings and private correspondence, this paper aims to correct that lacuna while contributing to a reorientation of the historiography of bacteriological epidemiology. In particular I trace Meyer’s intellectual exchanges with Smith, Burnet and the animal ecologist Charles Elton, over brucellosis, psittacosis and plague—exchanges that not only showed how environmental and ecological conditions could ‘tip the balance’ in favor of parasites but which transformed Meyer thinking about resistance to infection and disease. (shrink)

Reflecting on his scientific career toward the end of his life, the French-educated medical researcher René Dubos presented his flowering as an ecological thinker as a story of linear progression—the inevitable product of the intellectual seeds planted in his youth. But how much store should we set by Dubos’s account of his ecological journey? Resisting retrospective biographical readings, this paper seeks to relate the development of Dubos’s ecological ideas to his experimental practices and his career as a laboratory researcher. In (...) particular, I focus on Dubos’s studies of tuberculosis at the Rockefeller Institute in the period 1944–1956—studies which began with an inquiry into the tubercle bacillus and the physiochemical determinants of virulence, but which soon encompassed a wider investigation of the influence of environmental forces and host–parasite interactions on susceptibility and resistance to infection in animal models. At the same time, through a close reading of Dubos’s scientific papers and correspondence, I show how he both drew on and distinguished his ecological ideas from those of other medical researchers such as Theobald Smith, Frank Macfarlane Burnet, and Frank Fenner. However, whereas Burnet and Fenner tended to view ecological interactions at the level of populations, Dubos focused on the interface of hosts and parasites in the physiological environments of individuals. The result was that although Dubos never fully engaged with the science of ecology, he was able to incorporate ecological ideas into his thought and practices, and relate them to his holistic views on health and the natural harmony of man and his environment. (shrink)

The dualistic structures permeating western culture emphasize radical discontinuity between humans and nonhumans, but receptive attention to nonhuman others discloses both continuity and difference prevailing between other forms of life and our own. Recognizing that agency and subjectivity abound within nature alerts us to our potential for dominating and oppressing nonhuman others, as individuals and as groups. Reciprocally, seeing ourselves as biological beings may facilitate reconstructing our social reality to undo such destructive relationships.

Ecology has often been characterized as an immature scientific discipline. This paper explores some of the sources of this alleged immaturity. I argue that the perception of immaturity results primarily from the fact that historically ecologists have based their work upon two very different approaches to research.

In summary, many extant definitions of the niche concept are based on the geometric metaphor which represents the niche as an object embedded in a geometric space. There are several difficulties with this approach; the activities of organisms are not fully described, certain attributes of the functional aspect of the niche are not represented, the life cycles of organisms are not described, and the heuristic value of the concept diminishes with increasing dimensionality.An alternative and complementary approach to the niche is (...) the linguistic metaphor. This approach is based on the explanatory objective to explain the assembly of the ecosystems of successional seres from species pools and abiotic environmental factors. The linguistic metaphor suggests that a theory of ecosystem assembly is a generative grammar that associates particular species with environmental features. The niche of a particular species is defined in terms of the rules of a generative-transformational grammar used to derive an hierarchically organized description of that species. This approach also has conceptual difficulties: (1) empirically testing competence theories and distinguishing possible and impossible objects (e.g. species and ecosystems) are difficult, (2) evolutionary processes are difficult to incorporate, (3) the mathematical tools associated with grammars do not have the range of applicability of the mathematical tools associated with the geometric metaphor, and (4) the productions (hierarchical trees) of a grammar are not easily visualized.The geometric and linguistic metaphors of the niche should be viewed as complementary pictures of the same object. The geometric representation is most adept at describing community dynamics and stability, niche overlap, and organism response to abiotic environments. The linguistic metaphor represents well organism activities, niche dynamics, and ecosystem assembly. (shrink)

In many experimental sciences, like particle physics or molecular biology, the proper place for establishing facts is the laboratory. In the sciences of population biology, however, the laboratory is often seen as a poor approximation of what occurs in nature. Results obtained in the field are usually more convincing. This raises special problems: it is much more difficult to obtain stable, repeatable results in the field, where environmental conditions vary out of the experimenter’s control, than in the laboratory. We examine (...) here how this problem affected an influential experimental research tradition in community ecology, the study of the ecology of the rocky seashores. In the 1960s, a handful of North-American ecologists, most notably Joseph Connell, Robert Paine and Paul Dayton, made the rocky seashores a model study system for experimenting in the field. Their experiments were deceptively simple: they removed species living on the seashore and described the resulting effects on the local ecology. These experiments exerted a deep influence on community ecology. They provided evidence for speculative developments concerning the theory of interspecific competition, the factors responsible for species richness and the ecology of food webs. They also stimulated novel conceptual developments. In particular, Paine developed the predation hypothesis, which states that the presence of predators can favour species richness, before introducing the keystone species concept, according to which some species exert disproportionate effects on ecological systems. More broadly, these experiments gave support to a methodological trend in favour of field experimentation. Only controlled perturbations in the field, it seemed, provided a reliable method to get insights into the structure of ecological communities. However, as experiments were continued in time and repeated in different sites, divergent results appeared. We analyse here how intertidal researchers coped with the variability of environmental conditions and tried to stabilize their results. In the process, they reconsidered not only their early conclusions, but also the exclusive status given to field experiments. Expanding on this case study, we discuss some significant differences between laboratory and field experiments. (shrink)

In this paper I contrast two causal explanations of the outcome of a set of laboratory experiments in population ecology conducted by Thomas Park in the 1940s and 1950s. These experiments shed light on the problem of adducing evidence for the operation of competition (see Lloyd 1987 for a recent philosophical discussion) and are central to the group selection controversy because they form the empirical base for experimental studies of group selection conducted over the last 12 years (see Griesemer and (...) Wade 1988 for an analysis).The experiments investigated competition in laboratory populations of two flour beetle species, Tribolium confusum and Tribolium castaneum, but the results were more complex and interesting than expected; in addition to demonstrating competition in a laboratory system, they exhibited what Park called “competitive indeterminacy.” This phenomenon was of great interest to statisticians, who devised stochastic models to describe its demographic basis (e.g., M.S. Bartlett, P.H. Leslie, and J. Neyman; see Mertz 1972 for references). (shrink)

Two aspects of cognitive coupling, as brought forward in the Extended Mind Hypothesis, are discussed in this paper: how shall the functional coupling between the organism and some entity in his environment be spelled out in detail? What are the paradigmatic external entities to enter into that coupling? These two related questions are best answered in the light of an aetiological variety of functionalist argument that adds historical depth to the “active externalism” promoted by Clark and Chalmers and helps to (...) counter some of the core criticisms levelled against this view. Under additional reference to conceptual parallels between the Extended Mind Hypothesis and a set of heterodox theories in biology—environmental constructivism, niche construction, developmental systems theory—an argument for the grounding of environmentally extended cognitive traits in evolved biological functions is developed. In a spirit that seeks to integrate extended functionalism with views from cognitive integration and complementarity, it is argued that instances of environmental coupling should be understood as being constitutive to cognitive functions in either of two distinct ways. It is further argued that the historically and systematically prior environmental counterparts in that coupling are features of the natural environment. Language and linguistically imbued artefacts are likely to have descended from more basic relations that have an extension over the environment. (shrink)

I argue that a phenotypic trait can be an adaptation to a particular environmental condition, as against others, only if the environmental condition and the phenotype interactively cause fitness. Models of interactive environmental causes of fitness generally require that environments be individuated by explicit representation rather than by measures of environmental quality, although the latter understanding of ‘environment’ is more prominent in the philosophy of biology. Hence, talk of adaptations to some but not other environmental conditions relies on conceptions of (...) ‘environment’ importantly different from that commonly presupposed in philosophy of biology. (shrink)

We study the case of two sibling species ofHippolais(Aves). Very little differences can be observed in the morphology of both species. The breeding area of these species are complementary. Roughly, one species breeds North and East of Europe (Hippolais icterina) while the other breeds South and West of Europe (Hippolais polyglotta). There exitst a narrow zone of sympatry passing through Burgundy. Since several years, it has been observed that this area of sympatry was moving in the North-East direction at a (...) European scale. This means that progressivelyH. icterina is declining and is replaced byH. polyglotta. Some assumptions can be made in order to explain this evolution, for instance competition or predation. Series of observations concerning the diets of nestlings of both species have been realized. These observations show some differences in the diet compositions. The breeding success of the two species has been studied. Numerical simulations of a competition model taking into account the observed differences between the food types eaten by the two species are presented. These simulations do not explain the regression ofH. icterina. Then, we present numerical simulations of a predation model with one predator attacking the nestlings of both species. These simulations show that with time one of the two preys must extinct. Predation rather than competition seems to be the right explanation. (shrink)

This paper analyzes community ecologist Charles Elton’s ideas on animal communities, and situates them with respect to the classical opposition between organicist–holistic and individualistic–reductionist ecological views drawn by many historians of ecology. It is argued that Elton espoused a moderate ecological holism, which drew a middle way between the stricter ecological holism advocated by organicist ecologists and the merely aggregationist views advocated by some of their opponents. It is also argued that Elton’s moderate ecological holism resonated with his preference for (...) analogies between ecological communities and human societies over more common ones between communities and individual organisms. I discuss, on the one hand, how the functionalist-interactionist approach to community ecology introduced by Elton entailed a view of ecological communities as more or less self-maintaining functionally organized wholes, and how his ideas on this matter were incorporated into their views by organicist ecologists Frederic Clements, Victor Shelford, and Warder C. Allee et al. On the other hand, I identify some important divergences between Elton’s ecological ideas and those of organicist ecologists. Specifically, I show how Elton’s ideas on species distribution, animal migrations, and ecological succession entailed a view of animal communities as exhibiting a weaker degree of part-whole integration than that attributed to them by Clements and Shelford; and how Elton’s mixed stance on the balance of nature idea and his associated views on community stability attributed to communities a weaker form of self-regulation than that attributed to them by Allee et al. (shrink)

This paper reinforces the current consensus against the applicability of the selected effect theory of function in ecology. It does so by presenting an argument which, in contrast with the usual argument invoked in support of this consensus, is not based on claims about whether ecosystems are customary units of natural selection. Instead, the argument developed here is based on observations about the use of the function concept in functional ecology, and more specifically, research into the relationship between biodiversity and (...) ecosystem functioning. It is argued that a selected effect account of ecological functions is made implausible by the fact that it would conflict with important aspects of the understanding of function and ecosystem functional organization which underpins functional ecology’s research program. Specifically, it would conflict with (1) Functional ecology’s adoption of a context-based understanding of function and its aim to study the functional equivalence between phylogenetically-divergent organisms; (2) Functional ecology’s attribution to ecosystems of a lower degree of part-whole integration than the one found in paradigm individual organisms; and (3) Functional ecology’s adoption of a physiological or metabolic perspective on ecosystems rather than an evolutionary one. (shrink)

This article offers an analysis of ecologist Charles Elton’s “functional” concept of the niche and of the notion of function implicitly associated with it. It does so in part by situating Elton’s niche concept within the broader context of the “functionalist-interactionist” approach to ecology he introduced, and in relation to his views on the relationship between ecology and evolution. This involves criticizing the common claim that Elton’s idea of species as fulfilling functional roles within ecological communities committed him to an (...) idea of communities as units of selection. While such a claim implicitly attributes to Elton an understanding of function along the lines of the selected-effects theory of function advocated by many biologists and philosophers of biology, Elton’s use of the niche concept, I maintain, involves an understanding of function more along the lines of alternative nonselectionist theories such as the causal-role, goal-contribution, and organizational theories. I also briefly discuss how ecologists after Elton also tend to have typically adopted a nonselectionist understanding of the function concept, similar to his. (shrink)

We argue that ecology in general and biodiversity and ecosystem function research in particular need an understanding of functions which is both ahistorical and evolutionarily grounded. A natural candidate in this context is Bigelow and Pargetter’s evolutionary forward-looking account which, like the causal role account, assigns functions to parts of integrated systems regardless of their past history, but supplements this with an evolutionary dimension that relates functions to their bearers’ ability to thrive and perpetuate themselves. While Bigelow and Pargetter’s account (...) focused on functional organization at the level of organisms, we argue that such an account can be extended to functional organization at the community and ecosystem levels in a way that broadens the scope of the reconciliation between ecosystem ecology and evolutionary biology envisioned by many BEF researchers. By linking an evolutionary forward-looking account of functions to the persistence-based understanding of evolution defended by Bouchard and others, and to the theoretical research on complex adaptive systems, we argue that ecosystems, by forming more or less resilient assemblages, can evolve even while they do not reproduce and form lineages. We thus propose a Persistence Enhancing Propensity account of role functions in ecology to account for this overlap of evolutionary and ecological processes. (shrink)

This paper revisits the debate over whether the study of facilitation requires ecologists to revise their understanding of the relationship between realized and fundamental niches as conceptualized by Hutchinson. Following Rodriguez-Cabal et al., I argue against Bruno et al.’s claim that facilitation can make a species’ realized niche larger than its fundamental niche. However, I also maintain that the abstract Hutchinsonian conceptualization of the niche makes a whole range of facilitative interactions—which I propose to call ameliorative facilitation—invisible to niche-based approaches (...) to the study of ecological communities. I propose a way to incorporate ameliorative facilitation into such approaches. My proposal involves supplementing the Hutchinsonian realized/fundamental dyad with a third concept: the potential niche. This concept was introduced by ecologists studying the effects of environmental change on species distributions, but I show how it could also be fruitfully used in facilitation studies. I argue that this proposed solution is more appealing than Stachowicz’s suggestion that Hutchinson’s realized/fundamental contrast be applied to a spatial-geographical, as opposed to an abstract-conceptual, notion of the niche. (shrink)

Value claims about ecological entities, their functionality, and properties take center stage in so-called “ecological” ethical and aesthetic theories. For example, the claim that the biodiversity in an old-growth forest imbues it with “value in and for itself” is an explicit value claim about an ecological property. And the claim that one can study “the aesthetics of nature, including natural objects...such as ecosystems” presupposes that natural instances of a type of ecological entity exist and can be regarded as more or (...) less aesthetically valuable. My discussion below will bear wide implications for how claims like... (shrink)

This paper is a critical assessment of the epistemological impact of the systematic quantification of nature with the accumulation of big datasets on the practice and orientation of ecological science. We examine the contents of big databases and argue that it is not just accumulated information; records are translated into digital data in a process that changes their meanings. In order to better understand what is at stake in the ‘datafication’ process, we explore the context for the emergence and quantification (...) of biodiversity in the 1980s, along with the concept of the global environment. In tracing the origin and development of the global biodiversity information facility we describe big data biodiversity projects as a techno-political construction dedicated to monitoring a new object: the global diversity. We argue that, biodiversity big data became a powerful driver behind the invention of the concept of the global environment, and a way to embed ecological science in the political agenda. (shrink)

A theoretical framework is provided to explore teleonomy as a problem of self-causation, distinct from upward, downward and reticulate causation. Causality theories in biology are often formulated within hierarchy theories, where causation is conceptualized as running up or down the rungs of a ladder-like hierarchy or, more recently, as moving between multiple hierarchies. Research on the genealogy of cosmologies demonstrates that in addition to hierarchy theories, causality theories also depend upon ideas of time. This paper explores the roots and impact (...) of both time and hierarchy thinking on causal reasoning in the evolutionary sciences. Within evolutionary biology, the Neodarwinian synthesis adheres to a linear notion of time associated with linear hierarchies that portray upward causation. Eco-evo-devo schools recognize the importance of downward causation and consequently receive resistance from the standard view because downward causation is sometimes understood as backward causation, considered impossible by adherents of a linear time model. In contrast, downward causation works with a spatial or presential time notion. Hybridization, lateral gene transfer, infective heredity, symbiosis and symbiogenesis require recognition of reticulate causation occurring in both space and time, or spacetime, between distinct and interacting ontological hierarchies. Teleonomy is distinct from these types of causation because it invokes the problem of self-causation. By asking how the focal level in a hierarchy can persist through time, self-causation raises philosophical concerns on the nature of duration, identity and individuality. (shrink)

Holism and reductionism are usually seen as opposite and mutually exclusive approaches to nature. Recently, some have come to see them as complementary rather than mutually exclusive. In this book I have argued that, even stronger, they should be seen as mutually dependent and co-operating research programmes. I have discussed holism and reductionism in biology in general and in ecology in particular. After an introductory chapter I have provided an overview of holistic and reductionistic positions in biology, and of the (...) reduction problems to which they relate. I have argued that it is extremely important to distinguish between ontological, epistemological and methodological aspects of these problems. The overview has shown that there are actually several approaches to reduction problems in biology, which can be characterized as more or less radically or moderately holistic or reductionistic. It has shown also that there are three major ’contradistinctions’ between holism and reductionism in biology, to wit the doctrine of emergence versus the reduction thesis; functional explanations versus causal explanations; and phenomenology versus mechanicism. In later chapters I have shown that these ’contradistinctions’ are not really contradistinctions at all. In the next chapters I have discussed the terms ’reduce’ and ’reduction’ in science. In chapter 3 I have discussed the reduction of laws and theories. I have shown that there are many different types of reduction depending on the auxiliary hypotheses that are being used in addition to the reducing theory: approximation-, aggregation, correlation and/or identification-hypotheses. The distinctions between these types has later, in chapter 5, proved to be of great importance in defining the concept of ’emergence’. The major conclusion of chapter 3 was that reduction is an epistemological issue, pertaining to logical relations between statements or systems of statements. It should not be confused, therefore, with ’ontological reduction’, certainly not in one of its ordinary senses, such as diminishing, devaluating or the like. Scientific reductions are, with the exception of instrumentalistic reductions, kinds of explanation, and, moreover, non-eliminative kinds of explanation. This means that a reduced law or theory is not eliminated by the reducing theory but rather consolidated or even reinforced. The ontology of the reduced law or theory is thereby also being consolidated. In chapter 4 I have argued that the same holds for reductions of concepts. Concept reductions are supposed to be accomplished by means of so-called ontological identity relations, which are in turn supposed to connect terms in the law or theory to be reduced, which do not occur in the reducing theory, with theoretical terms of the reducing theory. Because they are called ontological identity relations, several philosophers have thought that concept reductions imply some form of ontological reduction. And because they often occur in the context of micro-reductions, some have even thought that concept reductions are themselves micro-reductions. As a result, the idea has arisen that ontological identity relations are connections between macro-concepts and micro-concepts. However, concept reductions by means of ontological identity relations cannot be micro-reductions, because micro-reduction is, by definition, reduction with an aggregation step. Moreover, ontological identity relations are relations between different representations of and the same type of thing or attribute. That is, they express different epistemological sides of the same ontological coin. Therefore, concept reductions cannot be considered ontological reductions either, certainly not in the ordinary sense of diminishing, devaluating or the like. Thus, concept reduction is, like law or theory reduction, an epistemological issue: it pertains to relations between, in this case, different concepts or representations of one and the same type of thing or attribute. ’Ontological reduction’, then, may, in the context of scientific reductions, actually be considered a contradiction of terms. In chapter 5 I have linked this conclusion to the emergence thesis and to the alleged contradistinction between this thesis and the reduction thesis. I have shown that the doctrine of emergence actually consists of two separate claims, namely an ontological thesis stating that a whole has emergent properties which the component parts do not possess, neither separately nor in other partial combinations; and an epistemological thesis stating that emergent properties of wholes are ’in principle’ irreducible. I have argued that the first claim may be regarded as a valid, universal thesis about relations between properties of wholes and properties of parts, but that the second claim is untenable. I have discussed many emergent properties of wholes, biological as well as physico-chemical, that have proved to be explainable in terms of micro-theories about the component parts and auxiliary hypotheses. Once again, however, it is important to realize that reduction is an epistemological issue, whereas emergence, in the sense of thesis, is an ontological one. It is not ’wholes’ or ’emergent properties of wholes’ that are being reduced but statements about them. Reduction is a kind of explanation, not of explaining-away. Thus, it leaves the ontology of whatever is reduced fully intact. I have developed a new definition of the term ’emergence’ which expresses emergence in terms of the auxiliary hypotheses that may be used in reductions, in particular aggregation-, correlation- and identification-hypotheses. This has even lead to two very remarkable conclusions. First, emergence may be considered the opposite of ontological identity. And second, if there were no emergence, there wouldn’t be any reductions either. to reduce.) In this respect, therefore, there is absolutely no contradistinction between holism and reductionism. On these grounds I have introduced my thesis that holism and reductionism should rather be seen as mutually dependent, and hence co-operating, research programmes than as conflicting views of nature or of relations between sciences. Holistic programmes play an important role in science as guide programmes for reductionistic programmes by discovering or developing macro-laws or -theories about phenomena at the level of wholes, which they themselves, however, cannot explain. For these explanations they depend on the fruits of reductionistic programmes. If the latter succeed in providing the explanations they act as supply programmes for the holistic guide programmes. Reductionistic programmes depend in turn on holistic programmes for providing the macro-laws or -theories that call for these explanations. In chapter 6 I have discussed an example of this mutual dependency in the form of the reduction of the Bohr-effect in animal physiology. I have shown that this law has been reduced to the theory of allostery, applied to hemoglobin molecules in red blood cells, and that this application of the theory of allostery has been reduced to the theory of chemical bonding. I have also shown that at least six research programmes were involved in these reductions, and that the relations between these programmes can be characterized very well in terms of the model of holistic guide programmes and reductionistic supply programmes. The only, but highly significant, qualification appearing from example was that the terms ’holistic’ and ’reductionistic’ are extremely relative and should always be related to a certain given level of organization. In chapter 7 I have resolved the remaining contradistinction between holism and reductionism in biology, viz. the need for functional explanations and the demand that explanations be causal. I have shown that functional explanations are perfectly legitimate explanations which in a sense can be reconstrued as causal explanations. I have argued that functional explanations are indispensable components of more comprehensive, causal-evolutionary explanations, because the latter appeal to the adaptive value, and hence the function, of the property to be explained. In that sense functions can be regarded as adaptations, and functional explanations can be regarded as a sort of ’short-hand’ for evolutionary explanations. Finally, I have shown that in the context of functional explanations, too, co-operation of holistic and reductionistic research programmes occurs. In part 2 I have applied my thesis to ecology. I have shown that in ecology too, holismreductionism disputes notwithstanding, co-operation of holistic and reductionistic research programmes occurs, or, in so far as this appears to be not the case, that it should be strived after. In chapter 8 I have given an overview of reduction problems in ecology and of the various approaches to these problems. I have noticed that in ecology, too, there are several, radical, moderate and anti-reductionistic research strategies, with respect to the levels of both ecosystems, communities and populations. I have also noticed, however, that concrete solutions to reduction problems in ecology are frustrated by what is called the intellectual immaturity or the anomalous status of ecology. This refers to the almost complete lack of general laws and theories in ecology, at least at the higher levels of communities and ecosystems. Of a number of possible causes I have lifted out two, which lend themselves to philosophical analysis and clarification. The first one is the ambiguity of a major part of the ecological vocabulary. This was the subject of chapters 9 and 10. The second is the inhibitory effect which holism-reductionism disputes have on the growth of knowledge. This was the subject of chapter 13. In chapter 9 I have discussed the concept of an community. It appeared that the term ’community’ is used for several different entities at various levels of organization. This ambiguity alone seems to be sufficient for the lack of ’general’ laws and theories about ’communities’. My purpose in chapter 9 was to try and contribute to a solution of this problem. In doing so I have argued, first, that it seems wise to use the term ’community’ only for groups of species belonging to a single taxonomic or phylogenetic group, and to use the term ’biocoenosis’ for the higher level of organization, defined as the biotic component of an ecosystem. Next I have argued that, although species within communities may of course interact with each other, interaction is itself not a necessary or sufficient condition for community membership. Finally, I have hit upon what is known as the most notorious problem in community ecology, to wit the boundary problem, as well as the problem of heterogeneity. I have found that the cause of these problems lies in the fact that communities are almost invariably being seen as groups of populations occurring together in space and time, whereas the empirical fact is rather that populations of different species mostly do not occur together in the same space, and, moreover, that the species composition of communities changes continuously. This has lead to the suggestion that a community had better be defined as the group of individuals of two or more species occurring in the area of intersection of populations of these species. It is only in such intersection areas that one can really talk of the co-occurrence of species. I have shown the empirical adequacy of this definition by discussing various salt marsh plant communities on the Dutch island of Schiermonnikoog. In chapter 10 I have clarified two other important, yet highly controversial, concepts in ecology, to wit habitat and niche. It appeared that there are at least four different habitat concepts in ecology, and as many niche concepts, the additional complication being that two of these habitat concepts correspond to two of these niche concepts, whence the distinction between habitat and niche is blurred. In addition, different habitat concepts appeared to correspond to different environment concepts and also to different biotope concepts. My purpose in chapter 10 was to disentangle all these concepts from one another and to supply each of them with a suitable term. I have done so by keeping a close eye on commonly accepted notions about habitat differentiation and niche differentiation, two ’principles’ allowing one to explain the coexistence or non-coexistence of species in communities. The results can be found in boxes 5 and 7 in chapter 10. In chapter 11 I was finally able to substantiate my claim that in ecology, too, co-operation of holistic and reductionistic research programmes occurs. I have discussed the reduction of the classical competition model of Lotka and Volterra to modern niche theory. The Lotka/Volterra model is a phenomenological model: it describes the possible effects of competition between two species. Modern niche theory, on the other hand, is a mechanistic theory, in which both the objects of competition and a mechanism are being specified. I have shown that the reduction was established with the help of a relatively simple identification hypothesis and two relatively simple aggregation hypotheses, making it yet another example of heterogeneous microreduction in biology. In the reduction, the Lotka/Volterra model acted as a holistic guide programme and modern niche theory acted as a reductionistic supply programme. In chapter 12 I have discussed another example of co-operation in the form of the approximative reduction of MacArthur and Wilson’s equilibrium theory of island biogeography. I have argued that this can be seen as a holistic model, which has been of great unifying and especially heuristic value, but which in time has been found to be a bit too simple and in some respects had to be corrected. Therefore, it can also be seen as an idealization in the sense of the model of idealization and concretization, which subsequent research programmes have concretized. Because these concretizations have a reductive character, we can see in this structure of idealization and concretization yet another example of the co-operation of holistic and reductionistic research programmes in ecology. A philosophically interesting side-conclusion was that the model of idealization and concretization thus appears to apply also to relations between research programmes, and not only to programme-internal developments. In chapter 13 I have discussed an example of the inhibitory effect that holism-reductionism disputes may have on the growth of knowledge. The example concerns a controversy in island biogeography over the role of interspecific competition in structuring communities. This controversy has lasted for about ten years, appeares to have died out rather than to have been resolved, and has produced nothing new in the field of explanations of community structure. I have shown that the major factors underlying the controversy were differing, in casu holistic and reductionistic, views of communities. Though this presented a problem for my thesis, I have argued that, ultimately, the controversy can be resolved, and in a sense has been resolved, in a way that corroborates my thesis. In the epilogue, finally, I have mentioned some remaining problems concerning the adequacy of the present reduction model in dealing with reduction in ecology. These problems pertain to the question of specificty of ecological laws and theories, the likelihood that reductions in ecology generally occur on the basis of several reducing micro-theories, and the possible relationship between the former two points. (shrink)

The classical holism-reductionism debate, which has been of major importance to the development of ecological theory and methodology, is an epistemological patchwork. At any moment, there is a risk of it slipping into an incoherent, chaotic Tower of Babel. Yet philosophy, like the sciences, requires that words and their correlative concepts be used rigorously and univocally. The prevalent use of everyday language in the holism-reductionism issue may give a false impression regarding its underlying clarity and coherence. In reality, the conceptual (...) categories underlying the debate have yet to be accurately defined and consistently used. There is a need to map out a clear conceptual, logical and epistemological framework. To this end, we propose a minimalist epistemological foundation. The issue is easier to grasp if we keep in mind that holism generally represents the ontological background of emergentism, but does not necessarily coincide with it. We therefore speak in very loose terms of the “holism-reductionism” debate, although it would really be better characterised by the terms emergentism and reductionism. The confrontation between these antagonistic paradigms unfolds at various semantic and operational levels. In definitional terms, there is not just emergentism and reductionism, but various kinds of emergentisms and reductionisms. (shrink)

This dissertation concerns the active role of the organism in evolutionary theory. In particular, it concerns how our conception of the relationship between organism and environment, and the nature of natural selection, influences the causal and explanatory role of organismic activity and behavior in evolutionary explanations. The overarching aim is to argue that the behaviors and activities of organisms can serve both as the explananda (that which is explained) and the explanantia (that which explains) in evolutionary explanations. I attempt to (...) achieve this aim by offering three central arguments. First, that the organism-environment relation is the ontologically basic unit of biology. A common way of conceiving the relationship between organism and environment is as a duality—as two causally independent systems that overlap through interaction. I think this is a mistake. There cannot be organisms without environments and vice versa. They are codependent and codetermined. Second, that natural selection is an ecological process. By this I mean that natural selection acts primarily on organism-environment interactions. It is only in virtue of organisms interacting with their environments that there can be fitness differences amongst individual organisms in a population. The ecological approach to natural selection also entails that the nature of the system(s) of inheritance involved in the reoccurrence of favorable organism-environment interactions is, in principle, immaterial to the process of selection. It only matters for selection that organism-environment interactions actually do reoccur in subsequent generations. Third, that niche construction theory—the most well-developed theoretical and conceptual framework for studying how the activities of organisms influences evolutionary dynamics—can be seen as fully a compatible and integrated part of evolutionary theory. Together, these three arguments constitute an overarching argument. Namely that we both can, and should, take organismic activity and behavior to be crucial explanatory elements in evolutionary theory. Throughout the dissertation I also show how these three arguments have consequences for other debates in the philosophy of evolutionary theory, such as the nature of evolutionary processes, the structure of selection-based explanations, the validity and utility of the proximate-ultimate distinction, and the nature of teleology in evolutionary systems. (shrink)

This chapter argues that the common claim that the ascription of ecological functions to organisms in functional ecology raises issues about levels of natural selection is ill-founded. This claim, I maintain, mistakenly assumes that the function concept as understood in functional ecology aligns with the selected effect theory of function advocated by many philosophers of biology (sometimes called “The Standard Line” on functions). After exploring the implications of Wilson and Sober’s defence of multilevel selection for the prospects of defending a (...) selected effect account of ecological functions, I identify three main ways in which functional ecology’s understanding of the function concept diverges from the selected effect theory. Specifically, I argue (1) that functional ecology conceives ecological functions as context-based rather than history-based properties of organisms; (2) that it attributes to the ecological function concept the aim of explaining ecosystem processes rather than with that of explaining the presence of organisms within ecosystems; and (3) that it conceives the ecological functions of organisms as use and service functions rather than design functions. I then discuss the extent to which the recently proposed causal role and organizational accounts of ecological functions better accord with the purposes for which the ecological concept is used in functional ecology. (shrink)

Evolutionary, ecological and ethical studies are, at the same time, specific scientific disciplines and, from an historical point of view, structurally linked domains of research. In a context of environmental crisis, the need is increasingly emerging for a connecting epistemological framework able to express a common or convergent tendency of thought and practice aimed at building, among other things, an environmental policy management respectful of the planet’s biodiversity and its evolutionary potential.

This is an informal presentation of the theory of niches understood as ecological contexts. The first part sets out the basic conceptual background. The second part outlines the main principles of the theory and addresses the question of how the theory can be extended to aid our thinking in relation to the special types of causal integrity that characterize niches and niched entities.