It is now widely accepted that microorganisms play many important roles in the lives of plants and animals. Every macroorganism has been shaped in some way by microorganisms. The recognition of the ubiquity and importance of microorganisms has led some to argue for a revolution in how we understand biological individuality and the primary units of natural selection. The term “holobiont” was introduced as a name for the biological unit made up by a host and all of its associated microorganisms, (...) and much of this new debate about biological individuality has focused on whether holobionts are integrated individuals or communities. In this paper, I show how parts of the holobiont can span both characterizations. I argue that most holobionts share more affinities with communities than they do with organisms, and that, except for maybe in rare cases, holobionts do not meet the criteria for being organisms, evolutionary individuals, or units of selection. (shrink)

Environmental philosophy is a hybrid discipline drawing extensively from epistemology, ethics, and philosophy of science and analyzing disciplines such as conservation biology, restoration ecology, sustainability studies, and political ecology. The book being discussed both provides an overview of environmental philosophy and develops an anthropocentric framework for it. That framework treats natural values as deep cultural values. Tradeoffs between natural values are analyzed using decision theory to the extent possible, leaving many interesting question for philosophical deliberation. This framework is supposed to (...) be applicable in practical contexts. (shrink)

David Steen and co-authors in this journal offer a philosophical argument to support an “Evolutionary Community Concept” to identify what they call “evolutionary communities.” They describe these as “unique collections of species that interact and have co-evolved in a given geographic area” and that include “co-evolved dependencies between different parts of a community.” Steen et al. refer to the coevolution of assemblages, collections, communities, dependencies, interspecific and abiotic interactions, and traits, but they do not define “co-evolution” or provide an example (...) in which co-evolution has been demonstrated. There may be as many ways to explain interactions among species as these interactions themselves; they may result, for example, from ecological fitting and phenotypic plasticity. In this paper, I argue that standard Darwinian Theory explains intraspecific microevolution or descent with modification within an interbreeding and potentially continuous population. The power, mechanism, or force that causes interspecific co-evolution, especially at the community scale, requires further explanation. (shrink)

Conservation biologists and other environmentalists confront five obstacles in building support for regulatory policies that seek to exclude or remove introduced plants and other non-native species that threaten to harm natural areas or the natural environment. First, the concept of “harm to the natural environment” is nebulous and undefined. Second, ecologists cannot predict how introduced species will behave in natural ecosystems. If biologists cannot define “harm” or predict the behavior of introduced species, they must target all non-native species as potentially (...) “harmful”. an impossibly large regulatory task. Third, loss of species richness may constitute harm to an environment, but introduced organisms typically, generally, and significantly add to species richness in ecosystems. If species richness correlates with desirable ecosystem properties, moreover, such as stability and productivity, as some ecologists believe, then introduced organisms, by increasing species richness, would support those desirable properties. Fourth, one may plausibly argue that extinction constitutes environmental harm, but there is no evidence that non-native species, especially plants, are significant causes of extinction, except for predators in certain lakes and other small island-like environments. Fifth, while aesthetic, ethical, and spiritual values may provide a legitimate basis for invasive species policy, biologists often cite concepts such as “biodiversity” and ecosystem “health” or “integrity” to provide a scientific justification. To assert that non-native species threaten biodiversity or undermine ecosystem health, however, may be to draw conceptual entailments or consequences from definitions of “biodiversity” and “integrity” that arbitrarily exclude non-native species or make the presence of exotic species a per se indicator of decline. (shrink)

The organizational account of biological functions interprets functions as contributions of a trait to the maintenance of the organization that, in turn, maintains the trait. As has been recently argued, however, the account seems unable to provide a unified grounding for both intra- and cross-generation functions, since the latter do not contribute to the maintenance of the same organization which produces them. To face this ‘ontological problem’, a splitting account has been proposed, according to which the two kinds of functions (...) require distinct organizational definitions. In this article, we propose a solution for the ontological problem, by arguing that intra- and cross-generation functions can be said to contribute in the same way to the maintenance of the biological organization, characterized in terms of organizational self-maintenance. As a consequence, we suggest maintaining a unified organizational account of biological functions. (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)

This paper argues that biological organisation can be legitimately conceived of as an intrinsically teleological causal regime. The core of the argument consists in establishing a connection between organisation and teleology through the concept of self-determination: biological organisation determines itself in the sense that the effects of its activity contribute to determine its own conditions of existence. We suggest that not any kind of circular regime realises self-determination, which should be specifically understood as self-constraint: in biological systems, in particular, self-constraint (...) takes the form of closure, i.e. a network of mutually dependent constitutive constraints. We then explore the occurrence of intrinsic teleology in the biological domain and beyond. On the one hand, the organisational account might possibly concede that supra-organismal biological systems could realise closure, and hence be teleological. On the other hand, the realisation of closure beyond the biological realm appears to be highly unlikely. In turn, the occurrence of simpler forms of self-determination remains a controversial issue, in particular with respect to the case of self-organising dissipative systems. (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)

While holist views such as ecocentrism have considerable intuitive appeal, arguing for the moral considerability of ecological wholes such as ecosystems has turned out to be a very difficult task. In the environmental ethics literature, individualist biocentrists have persuasively argued that individual organisms—but not ecological wholes—are properly regarded as having a good of their own . In this paper, I revisit those arguments and contend that they are fatally flawed. The paper proceeds in five parts. First, I consider some problems (...) brought about by climate change for environmental conservation strategies and argue that these problems give us good pragmatic reasons to want a better account of the welfare of ecological wholes. Second, I describe the theoretical assumptions from normative ethics that form the background of the arguments against holism. Third, I review the arguments given by individualist biocentrists in favour of individualism over holism. Fourth, I review recent work in the philosophy of biology on the units of selection problem, work in medicine on the human biome, and work in evolutionary biology on epigenetics and endogenous viral elements. I show how these developments undermine both the individualist arguments described above as well as the distinction between individuals and wholes as it has been understood by individualists. Finally, I consider five possible theoretical responses to these problems. (shrink)

Ecological communities are seldom, if ever, biological individuals. They lack causal boundaries as the populations that constitute communities are not congruent and rarely have persistent functional roles regulating the communities’ higher-level properties. Instead we should represent ecological communities indexically, by identifying ecological communities via the network of weak causal interactions between populations that unfurl from a starting set of populations. This precisification of ecological communities helps identify how community properties remain invariant, and why they have robust characteristics. This respects the (...) diversity and aggregational nature of these complex systems while still vindicating them as units worthy of investigation. (shrink)

Biodiversity is a key concept in the biological sciences. While it has its origin in conservation biology, it has become useful across multiple biological disciplines as a means to describe biological variation. It remains, however, unclear what particular biological units the concept refers to. There are currently multiple accounts of which biological features constitute biodiversity and how these are to be measured. In this paper, I draw from the species concept debate to argue for a set of desiderata for the (...) concept of “biodiversity” that is both principled and coheres with the concept’s use. Given these desiderata, this concept should be understood as referring to difference quantified in terms of the phylogenetic structure of lineages, also known as the ‘tree of life’. (shrink)

A phenomenological community is an identifiable assemblage of organisms in a local habitat patch: a local wetland or mudflat are typical examples. Such communities are typically persistent: membership and abundance stay fairly constant over time. In this paper I discuss whether phenomenological communities are functionally structured, causal systems that play a role in determining the presence and abundance of organisms in a local habitat patch. I argue they are not, if individualist models of community assembly are vindicated; i.e., if the (...) presence of one species is not typically explained by the presence or absence of specific other species. I discuss two alternatives to in- dividualism, and conclude by arguing for a dimensional model of phenomenological communities. The causal salience of a phenomenological community depends on three factors: the extent to which it is internally regulated, the extent to which it has robust boundaries, and the extent to which it has emergent properties. I conclude by using this model to frame a natural research agenda for community ecology. (shrink)

In this paper I examine the connection between accounts of biological teleology and the biocentrist claim that all living beings have a good of their own. I first present the background for biocentrists’ appeal to biological teleology. Then I raise a problem of scope for teleology-based biocentrism and, drawing in part on recent work by Basl and Sandler, I discuss Taylor and Varner’s responses to this problem. I then challenge Basl and Sandler’s own response to the scope problem for its (...) reliance on a selectionist account of organismic teleology. Finally I examine the prospects for a biocentrist response to the problem of scope based on an alternative organisational account of internal teleology. I conclude by assessing the prospects for teleology-based biocentrism. (shrink)

The etiological approach to ‘proper functions’ in biology can be strengthened by relating it to Robert Cummins' general treatment of function ascription. The proper functions of a biological trait are the functions it is assigned in a Cummins-style functional explanation of the fitness of ancestors. These functions figure in selective explanations of the trait. It is also argued that some recent etiological theories include inaccurate accounts of selective explanation in biology. Finally, a generalization of the notion of selective explanation allows (...) an analysis of the proper functions of human artifacts. (shrink)

In this paper, I explore the links between liberal political theory and the evaluative nature of medical classification, arguing for stronger recognition of those links in a liberal model of medical practice. All judgments of medical or psychiatric "dysfunction," I argue, are fundamentally evaluative, reflecting our collective willingness or reluctance to tolerate and/or accommodate the conditions in question. Illness, then, is "socially constructed." But the relativist worries that this loaded phrase evokes are unfounded; patients, doctors, and communities will agree in (...) the vast majority of cases about what counts as illness. Where they cannot come to agreement, however, we are faced with precisely the sort of dispute about values and ways of life that the institutions of the liberal state are designed to accommodate. I accordingly sketch a model of medical practice, based loosely on Jürgen Habermas's political theories, designed to maximize both our awareness and our understanding of these disputes. (shrink)

Despite the voluminous literature on biological functions produced over the last 40 years, few philosophers have studied the concept of function as it is used in neuroscience. Recently, Craver (forthcoming; also see Craver 2001) defended the causal role theory against the selected effects theory as the most appropriate theory of function for neuroscience. The following argues that though neuroscientists do study causal role functions, the scope of that theory is not as universal as claimed. Despite the strong prima facie superiority (...) of the causal role theory, the selected effects theory (when properly developed) can handle many cases from neuroscience with equal facility. It argues this by presenting a new theory of function that generalizes the notion of a ‘selection process’ to include processes such as neural selection, antibody selection, and some forms of learning—that is, to include structures that have been differentially retained as well as those that have been differentially reproduced. This view, called the generalized selected effects theory of function, will be defended from criticism and distinguished from similar views in the literature. (shrink)

I distinguish two forms of pluralism about biological functions, between-discipline pluralism and within-discipline pluralism. Between-discipline pluralism holds that different theories of function are appropriate for different subdisciplines of biology and psychology. I provide reasons for rejecting this view. Instead, I recommend within-discipline pluralism, which emphasizes the plurality of function concepts at play within any given subdiscipline of biology and psychology.

I present and defend the generalized selected effects theory (GSE) of function. According to GSE, the function of a trait consists in the activity that contributed to its bearer’s differential reproduction, or differential retention, within a population. Unlike the traditional selected effects (SE) theory, it does not require that the functional trait helped its bearer reproduce; differential retention is enough. Although the core theory has been presented previously, I go significantly beyond those presentations by providing a new argument for GSE (...) and defending it from a recent objection. I also sketch its implications for teleosemantics and philosophy of medicine. (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 chapter develops the idea that the germ-soma split and the suppression of individual fitness differences within the corporate entity are not always essential steps in the evolution of corporate individuals. It illustrates some consequences for multilevel selection theory. It presents evidence that genetic heterogeneity may not always be a barrier to successful functioning as a higher-level individual. This chapter shows that levels-of-selection theorists are wrong to assume that the central problem in transitions is always that of minimizing within-group competition. (...) Evidence of intralevel conflict does not qualify as evidence against the existence of a higher level of selection. (shrink)

The concept of ecosystem health is problematic. Do ecosystems as such exist? Is health an objective condition of organisms or is it socially constructed? Can 'health' be unequivocally predicated of ecosystems? Is ecosystem health both objective and valuative? Are ecosystem health and biological integrity identical? How do these concepts interface with the concept of biodiversity? Ecosystems exist, although they are turning out to be nested sets of linked process-functions with temporal boundaries, not tangible superorganisms with spatial boundaries. Ecosystem health – (...) or normal occurrence of ecological processes and functions – is an objective condition of ecosystems, although the concept of ecosystem health allows some room for personal and social determination or construction. Ecosystem health is prudentially, aesthetically, and intrinsically valuable, although the value of ecosystem health is subjectively conferred. Biodiversity and biological integrity are different from, but not unrelated to, ecosystem health. Together these three normative concepts represent complementary conservation goals. (shrink)

A recent development in biology has been the growing acceptance that holobionts, entities comprised of symbiotic microbes and their host organisms, are widespread in nature. There is agreement that holobionts are evolved outcomes, but disagreement on how to characterize the operation of natural selection on them. The aim of this paper is to articulate the contours of the disagreement. I explain how two distinct foundational accounts of the process of natural selection give rise to competing views about evolutionary individuality.

This paper addresses the foundations of Teleological Individualism, the view that organisms, even non-sentient organisms, are goal-oriented systems while biological collectives, such as ecosystems or conspecific groups, are mere assemblages of organisms. Typical defenses of Teleological Individualism ground the teleological organization of organisms in the workings of natural selection. This paper shows that grounding teleological organization in natural selection is antithetical to Teleological Individualism because such views assume a view about the units of selection on which it is only individual (...) organisms that are units of selection. However, none of the Conventionalist, Reductionist, or Multi-Level Realist theories serve to justify such an assumption. Thus, Teleological Individualism cannot be grounded in natural selection. (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)

Philosophy of ecology has been slow to become established as an area of philosophical interest, but it is now receiving considerable attention. This area holds great promise for the advancement of both ecology and the philosophy of science. Insights from the philosophy of science can advance ecology in a number of ways. For example, philosophy can assist with the development of improved models of ecological hypothesis testing and theory choice. Philosophy can also help ecologists understand the role and limitations of (...) mathematical models in ecology. On the other side, philosophy of science will be advanced by having ecological case studies as part of the stock of examples. Ecological case studies can shed light on old philosophical topics as well as raise novel issues for the philosophy of science. For example, understanding theoretical terms such as “biodiversity” is important for scientific reasons, but such terms also carry political importance. Formulating appropriate definitions for such terms is thus not a purely scientific matter, and this may prompt a reevaluation of philosophical accounts of defining theoretical terms. We consider some of the topics currently receiving attention in the philosophy of ecology and other topics in need of attention. Our aim is to prompt further exchange between ecology and philosophy of science and to help set the agenda for future work in the philosophy of ecology. The topics covered include: the role of mathematical models, environmental problem formulation, biodiversity, and environmental ethics. (shrink)