Natural unity is not an entirely innocent worldview. Various scientists and scholars have tried to use natural unity to legitimize political viewpoints that stand in stark opposition to those of environmentalists who promulgate the unity of nature for progressive social change. This book discusses how this situation has transpired by examining the recent history of the social political and intellectual processes involved. These include the social construction of the Organicism versus Mechanicism debate in ecology, the links between neo-classical economic principles (...) and the 'New Sciences', the techno-scientific background of Chaos theory and Gaja theory, and the social conservatism of ecological functionalism. (shrink)

In the six decades since the publication of Julian Huxley's Evolution: The Modern Synthesis, spectacular empirical advances in the biological sciences have been accompanied by equally significant developments within the core theoretical framework of the discipline. As a result, evolutionary theory today includes concepts and even entire new fields that were not part of the foundational structure of the Modern Synthesis. In this volume, sixteen leading evolutionary biologists and philosophers of science survey the conceptual changes that have emerged since Huxley's (...) landmark publication, not only in such traditional domains of evolutionary biology as quantitative genetics and paleontology but also in such new fields of research as genomics and EvoDevo. Most of the contributors to Evolution—The Extended Synthesis accept many of the tenets of the classical framework but want to relax some of its assumptions and introduce significant conceptual augmentations of the basic Modern Synthesis structure—just as the architects of the Modern Synthesis themselves expanded and modulated previous versions of Darwinism. This continuing revision of a theoretical edifice the foundations of which were laid in the middle of the nineteenth century—the reexamination of old ideas, proposals of new ones, and the synthesis of the most suitable—shows us how science works, and how scientists have painstakingly built a solid set of explanations for what Darwin called the "grandeur" of life. (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)

Philosophers of evolutionary biology favor the so-called etiological concept of function according to which the function of a trait is its evolutionary purpose, defined as the effect for which that trait was favored by natural selection. We term this the selected effect (SE) analysis of function. An alternative account of function was introduced by Robert Cummins in a non-evolutionary and non-purposive context. Cummins''s account has received attention but little support from philosophers of biology. This paper will show that a similar (...) non-purposive concept of function, which we term causal role (CR) function, is crucial to certain research programs in evolutionary biology, and that philosophical criticisms of Cummins''s concept are ineffective in this scientific context. Specifically, we demonstrate that CR functions are a vital and ineliminable part of research in comparative and functional anatomy, and that biological categories used by anatomists are not defined by the application of SE functional analysis. Causal role functions are non-historically defined, but may themselves be used in an historical analysis. Furthermore, we show that a philosophical insistence on the primary of SE functions places practicing biologists in an untenable position, as such functions can rarely be demonstrated (in contrast to CR functions). Biologists who study the form and function of organismal design recognize that it is virtually impossible to identify the past action of selection on any particular structure retrospectively, a requirement for recognizing SE functions. (shrink)

In this paper I defend an etiological theory of biological functions (according to which the proper function of a trait is the effect for which it was selected by natural selection) against three objections which have been influential. I argue, contrary to Millikan, that it is wrong to base our defense of the theory on a rejection of conceptual analysis, for conceptual analysis does have an important role in philosophy of science. I also argue that biology requires a normative notion (...) of a "proper function", and that a normative notion is not ahistorical. (shrink)

Many areas of science develop by discovering mechanisms and role functions. Cummins' (1975) analysis of role functions-according to which an item's role function is a capacity of that item that appears in an analytic explanation of the capacity of some containing system-captures one important sense of "function" in the biological sciences and elsewhere. Here I synthesize Cummins' account with recent work on mechanisms and causal/mechanical explanation. The synthesis produces an analysis of specifically mechanistic role functions, one that uses the characteristic (...) active, spatial, temporal, and hierarchical organization of mechanisms to add precision and content to Cummins' original suggestion. This synthesis also shows why the discovery of role functions is a scientific achievement. Discovering a role function (i) contributes to the interlevel integration of multilevel mechanisms, and (ii) provides a unique, contextual variety of causal/mechanical explanation. (shrink)

I defend the historical definition of "function" originally given in my Language, Thought and Other Biological Categories (1984a). The definition was not offered in the spirit of conceptual analysis but is more akin to a theoretical definition of "function". A major theme is that nonhistorical analyses of "function" fail to deal adequately with items that are not capable of performing their functions.

This book is a critical survey of and guidebook to the literature on biological functions. It ties in with current debates and developments, and at the same time, it looks back on the state of discourse in naturalized teleology prior to the 1970s. It also presents three significant new proposals. First, it describes the generalized selected effects theory, which is one version of the selected effects theory, maintaining that the function of a trait consists in the activity that led to (...) its differential persistence or reproduction in a population, and not merely its differential reproduction. Secondly, it advances “within-discipline pluralism” (as opposed to between-discipline pluralism) a new form of function pluralism, which emphasizes the coexistence of function concepts within diverse biological sub-disciplines. Lastly, it provides a critical assessment of recent alternatives to the selected effects theory of function, namely, the weak etiological theory and the systems-theoretic theory. The book argues that, to the extent that functions purport to offer causal explanations for the existence of a trait, there are no viable alternatives to the selected effects view. -/- The debate about biological functions is still as relevant and important to biology and philosophy as it ever was. Recent controversies surrounding the ENCODE Project Consortium in genetics, the nature of psychiatric classification, and the value of ecological restoration, all point to the continuing relevance to biology of philosophical discussion about the nature of functions. In philosophy, ongoing debates about the nature of biological information, intentionality, health and disease, mechanism, and even biological trait classification, are closely related to debates about biological functions. (shrink)

This book is a sustained examination of issues in the philosophy of ecology that have been a source of controversy since the emergence of ecology as an explicit scientific discipline. The controversies revolve around the idea of a balance of nature, the possibility of general ecological knowledge and the role of model-building in ecology. The Science of the Struggle for Existence is also a detailed treatment of these issues that incorporates both a comprehensive investigation of the relevant ecological literature and (...) the development of an explicit theoretical framework in the philosophy of science. It addresses issues in the philosophy of ecology that are of particular importance for the deployment of ecology in the solution of environmental problems. It will have a cross-disciplinary appeal and will interest students and professionals in science, the philosophy of science, and environmental studies as well as policy-makers. (shrink)

This book is a personal history and apology, written by one of this century's most distinguished small mammal ecologists, for a life in science spent working on problems for which no final dramatic conclusion was reached. Included along the way are some important anecdotes and history about Charles Elton and the pioneering work at the Bureau of Animal Population at Oxford University, from which most of modern population ecology has grown, and insigts on the philosophy and practice of science.

What Is Biodiversity? is a theoretical and conceptual exploration of the biological world and how diversity is valued. Maclaurin and Sterelny explore not only the origins of the concept of biodiversity, but also how that concept has been shaped by ecology and more recently by conservation biology. They explain the different types of biodiversity important in evolutionary theory, developmental biology, ecology, morphology and taxonomy and conclude that biological heritage is rich in not just one biodiversity but many. Maclaurin and Sterelny (...) also explore the case for the conservation of these biodiversities using option value theory, a tool borrowed from economics. (shrink)

An examination of difficulties in three standard accounts of functions leads to the suggestion that sentences of the form "the function of x is to do y" are used to make a variety of different claims, all of which involve a means-end relationship and the idea of design, or use, or benefit. The analysis proposed enables us to see what is right and also wrong with accounts that analyze the meaning of function statements in terms of good consequences, goals, and (...) etiological explanation. It also enables us to show that function sentences can be used in providing various types of explanations, including, in certain cases, noncausal explanations of the presence of the item with the function. (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 2001 book offers an examination of functional explanation as it is used in biology and the social sciences, and focuses on the kinds of philosophical presuppositions that such explanations carry with them. It tackles such questions as: why are some things explained functionally while others are not? What do the functional explanations tell us about how these objects are conceptualized? What do we commit ourselves to when we give and take functional explanations in the life sciences and the social (...) sciences? McLaughlin gives a critical review of the debate on functional explanation in the philosophy of science. He discusses the history of the philosophical question of teleology, and provides a comprehensive review of the post-war literature on functional explanation. What Functions Explain provides a sophisticated and detailed Aristotelian analysis of our concept of natural functions, and offers a positive contribution to the ongoing debate on the topic. (shrink)

This paper reviews the debate on the notion of biological function and on functional explanation as this takes place in philosophy. It describes the different perspectives, issues, intuitions, theories and arguments that have emerged. The author shows that the debate has been too heavily influenced by the concerns of a naturalistic philosophy of mind and argues that in order to improve our understanding of biology the attention should be shifted from the study of intuitions to the study of the actual (...) practice of biological inquiry. (shrink)

I argue that there are at least four different ways in which the term ‘function’ is used in connection with the study of living organisms, namely: function as activity, function as biological role, function as biological advantage, and function as selected effect. Notion refers to what an item does by itself; refers to the contribution of an item or activity to a complex activity or capacity of an organism; refers to the value for the organism of an item having a (...) certain character rather than another; refers to the way in which a trait acquired and has maintained its current share in the population. The recognition of a separate notion of function as biological advantage solves the problem of the indeterminate reference situation that has been raised against a counterfactual analysis of function, and emphasizes the importance of counterfactual comparison in the explanatory practice of organismal biology. This reveals a neglected problem in the philosophy of biology, namely that of accounting for the insights provided by counterfactual comparison. (shrink)

With the rejection of group selectionist derivations of ecological phenomena so incisively given by George Williams in 1966,43 Nicholson's long-ignored messages met with acceptance. Species benefit became, explicitly, incidental. But the reorientation was not just about a point of ecological theory. It was more fundamentally about theoretical style, the element shared by Wynne-Edwards' work and the newer, evolutionary ecology. That current approach is well expressed in an already classic paper by the British plant ecologist John Harper: Ultimately all the discoveries (...) of descriptive, production or population ecology must find their meaning in evolutionary phenomena... Evolutionary thinking concentrates attention on the behaviour of the individual and his descendants. If nothing in biology has meaning except in the light of evolution and if evolution is about individuals and their descendants — i.e. fitness — we should not expect to reach any depth of understanding from studies that are based at the level of the superindividual... What we see as the organised behaviour of systems is the result of the fate of individuals.44The emphasis in this passage is on style of thinking more than on study matter. What is so different from earlier ecologists is the rejection of the superorganism and its associated baggage of selfimposed constraints in an end-oriented process, such as balance of nature or species-level adaptation.The hallmark of post-1959 evolutionary ecology is the recognition, the making explicit of the question that the selective, genetical process of fitness (fates of genes) has to be separated from the observations of different levels of adaptiveness. An easy conflation of cause and result is no longer satisfying; now attempts to disentangle the complex strands tying population phenomena to the different levels of selection are providing the excitement of a new research program. In this program evolutionary theory is used to define the appropriate questions, as exemplified by the organization of the textbook Evolutionary Ecology.45 Our discussion of the invention of a new approach, with a characteristic style and set of interests, must also be about how the practitioners of ecology have perceived themselves. The early attempts of Poulton and Nicholson were not taken up by the vigorously growing science of ecology, which developed first - and intentionally — along a narrower, hierarchically restrained set of questions about population. The depth of Poulton's and Nicholson's submergence is indicated by the fact that even Wynne-Edwards did not argue in the manner of one rescuing a minor tradition. Rather, he thought he had a new approach, that of placing evolution in the foundations of ecology. Although this approach was not exactly original, we should be cautious in grouping all evolutionary concerns within ecology in one intellectual tradition. The particular style of selectionist deduction invigorated by Wynne-Edwards was successful only in the volatile situation of a rapidly growing discipline, full of self-critical examination and new rigor. Whereas in 1930 evolutionary concerns did not provide that rigor, by 1960 they could - and did. (shrink)

I argue that there are at least four different ways in which the term ‘function’ is used in connection with the study of living organisms, namely: function as activity, function as biological role, function as biological advantage, and function as selected effect. Notion refers to what an item does by itself; refers to the contribution of an item or activity to a complex activity or capacity of an organism; refers to the value for the organism of an item having a (...) certain character rather than another; refers to the way in which a trait acquired and has maintained its current share in the population. The recognition of a separate notion of function as biological advantage solves the problem of the indeterminate reference situation that has been raised against a counterfactual analysis of function, and emphasizes the importance of counterfactual comparison in the explanatory practice of organismal biology. This reveals a neglected problem in the philosophy of biology, namely that of accounting for the insights provided by counterfactual comparison. (shrink)

According to a traditional view, scientific laws and theories constitute algorithmic compressions of empirical data sets collected from observations and measurements. This article defends the thesis that, to the contrary, empirical data sets are algorithmically incompressible. The reason is that individual data points are determined partly by perturbations, or causal factors that cannot be reduced to any pattern. If empirical data sets are incompressible, then they exhibit maximal algorithmic complexity, maximal entropy and zero redundancy. They are therefore maximally efficient carriers (...) of information about the world. Since, on algorithmic information theory, a string is algorithmically random just if it is incompressible, the thesis entails that empirical data sets consist of algorithmically random strings of digits. Rather than constituting compressions of empirical data, scientific laws and theories pick out patterns that data sets exhibit with a certain noise. (shrink)

Function and teleology can be naturalized either by reference to systems with a particular type of organization or by reference to a particular kind of history. As functions are generally ascribed to states or traits according to their current role and regardless of their origin, etiological accounts are inappropriate. Here, I offer a systems-theoretical interpretation as a new version of an organizational account of functionality, which is more comprehensive than traditional cybernetic views and provides explicit criteria for empirically testable function (...) ascriptions. I propose, that functional states, traits or items are those components of a complex system, which are under certain circumstances necessary for their self-re-production. I show, how this notion can be applied in intra- and trans-generational function ascriptions in biology, how it can deal with the problems of multifunctionality and functional equivalents, and how it relates to concepts like fitness and adaptation. Finally, I argue that most intentional explanations can be treated as functional explanations. (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)

I examine the use of the term function in Aldo Leopold’s land ethic, invoked as (1) the healthy functioning of the land community, which is dependent on (2) the maintenance of the characteristic functions of populations that are parts of the land community. The latter can be understood as referring to interactions between species that are the products of coevolution (such as parasite-host, predator-prey) and, thus, in terms of the “selected effect” account of function. The performance of these functions under (...) certain conditions maintain what Leopold took to be the healthy functioning of a land community. (shrink)

Functional language is ubiquitous in ecology, mainly in the researches about biodiversity and ecosystem function. However, it has not been adequately investigated by ecologists or philosophers of ecology. In the contemporary philosophy of ecology we can recognize a kind of implicit consensus about this issue: while the etiological approaches cannot offer a good concept of function in ecology, Cummins’ systemic approach can. Here we propose to go beyond this implicit consensus, because we think these approaches are not adequate for ecology. (...) We argue that a sound epistemological framework to function in ecology is to be found in organizational approaches. In this line, we define function in ecology as a precise effect of a given constraint on the ecosystem flow of matter and energy performed by a given item of biodiversity, within a closure of constraints. We elaborate on this definition by developing a case study of a bromeliad ecosystem. (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 concept of an ecological niche (econiche) has been used in a variety of ways, some of which are incompatible with a relational or functional interpretation of the term. This essay seeks to standardize usage by limiting the concept to functional relations between organisms and their surroundings, and to revise the concept to include epistemic relations. For most organisms, epistemics are a vital aspect of their functional relationships to their surroundings and, hence, a major determinant of their econiche. Rejecting the (...) traditional dualism of organism and environment, an econiche is defined as the reciprocal (dual) of a functionally specified class of organisms (FSTU). From this perspective, an econiche necessarily implies a certain type of organism, and a class of functionally similar organisms implies a special econiche.The econiche concept is also discussed in relation to other ecological terms that reflect the distributional patterns of organisms, such as habitat, and the concept of an empty niche is criticized. (shrink)

This thesis investigates the equilibrium concept in population ecology. Equilibrium is a state in which something is in balance. What is in balance and why? This two-part question has concerned ecologists for decades and is still debated today. One influential account postulates density-dependent mechanisms in the population as leading to equilibrium densities. This account suggests a form of self-regulations of populations. However the existence of density-dependence and even the importance of regulation are hotly contested issues in ecology. ;Focusing on the (...) debate about equilibrium, I follow a form of naturalized epistemology, in which knowledge is to be understood by looking into the workings of science itself. I examine, in particular, the scientists' reasons to hold on to the concept of equilibrium. I describe these reasons as justifying the use of the concept. To understand the debate about equilibrium we need to answer the question: "How is equilibrium justified?" ;I show that during the debate the justification of equilibrium has changed due to a change in reference of 'equilibrium'. With 'equilibrium' being defined as the outcome of density-dependent processes, equilibrium has come to be understood as a statistical property of a data set. This change in how the application of the concept is justified is a change that has itself been negotiated by scientists in debating their claims about equilibrium. I investigate this negotiation process and argue that it cannot be explained away by referring to the use of models to understand equilibrium. On the contrary, thinking of theories as models makes the investigation of negotiations even more important. ;The negotiation of equilibrium is a normal scientific process. What is negotiated is the relation between evidence and theory, and I maintain that this relation has to be understood in historical terms. The justification of the equilibrium concept is based on evidence, but also on interests, and scientific practice. Although this historical perspective does not explain all aspects of how the concept is justified, it offers an understanding of the concept that is critical to evaluating its use in population ecology. (shrink)

In the literature the term “natural balance” occurs frequently and is used for highly divergent collections of facts and for results arrived at by different methods. In this paper it is attempted to give a review of the many possible meanings of “balance in nature”, and to evaluate the application of the term in the scientific literature.To achieve this twofold objective it seemed useful to start by giving as objective as possible a description of the “balance situation” in natural communities, (...) in analogy with the original physical concept and according to strict scientific criteria. On the basis of this working standard, part of the ecological literature is discussed with respect to statements and presumptions concerning “balance”. The extensive literature is classified according to several groups, on the one hand on the basis of the nature of the views , on the other hand according to the importance that certain authors attach to the theory of a given author. For each of these groups, an evaluation of the content of the term “balance” is given. It was found that the reasoning is not always sound and that the nature of the data on which the concept is based varies widely, which also holds for the method of reasoning applied in the conclusions drawn. In none of the publications arguments were found supporting the existence of a natural “balance situation” as formulated in the standard description. Consequently, it seemed justified to conclude that it is not correct to apply a concept “natural balance”, defined purely scientifically, to natural situations, either because “balance” in the indicated sense is simply not present or because the situation under consideration is not a natural one . The concept “natural balance” has had only heuristic value for scientific research. In other contexts it has a different value, because there criteria other than scientific or methodological ones hold for the content of a concept. It is therefore not objectionable to use the term “natural balance” in common parlance, especially since this usage makes it possible to convey to the layman a great deal in few words. (shrink)