I argue that the propensity interpretation of fitness, properly understood, not only solves the explanatory circularity problem and the mismatch problem, but can also withstand the Pandora’s box full of problems that have been thrown at it. Fitness is the propensity (i.e., probabilistic ability, based on heritable physical traits) for organisms or types of organisms to survive and reproduce in particular environments and in particular populations for a specified number of generations; if greater than one generation, “reproduction” includes descendants of (...) descendants. Fitness values can be described in terms of distributions of propensities to produce varying number of offspring and can be modeled for any number of generations using computer simulations, thus providing both predictive power and a means for comparing the fitness of different phenotypes. Fitness is a causal concept, most notably at the population level, where fitness differences are causally responsible for differences in reproductive success. Relative fitness is ultimately what matters for natural selection. (shrink)

The traditional practice of establishing morphological types and investigating morphological organization has found new support from evolutionary developmental biology (evo-devo), especially with respect to the notion of body plans. Despite recurring claims that typology is at odds with evolutionary thinking, evo-devo offers mechanistic explanations of the evolutionary origin, transformation, and evolvability of morphological organization. In parallel, philosophers have developed non-essentialist conceptions of natural kinds that permit kinds to exhibit variation and undergo change. This not only facilitates a construal of species (...) and higher taxa as natural kinds, but also broadens our perspective on the diversity of kinds found in biology. There are many different natural kinds relevant to the investigative and explanatory aims of evo-devo, including homologues and developmental modules. (shrink)

This paper examines a new challenge to neo-Darwinism, a movement known as process structuralism. The process structuralist critique of neo-Darwinism holds 1) that there are general laws in biology and that biologists should search for these laws; 2) that there are general forms of morphology and development and that biologists should attempt to uncover these forms; 3) that organisms are unified wholes that cannot be understood without adopting a holistic perspective; and 4) that no special, causal primacy should be given (...) to the genes in development and morphology. This paper places process structuralism in its historical context, examines the philosophical underpinnings of the movement, and discusses some of the evidence used to support its claims. (shrink)

Following Mayr (1961) evolutionary biologists often maintain that the hallmark of biology is its evolutionary perspective. In this view, biologists distinguish themselves from other natural scientists by their emphasis on why-questions. Why-questions are legitimate in biology but not in other natural sciences because of the selective character of the process by means of which living objects acquire their characteristics. For that reason, why-questions should be answered in terms of natural selection. Functional biology is seen as a reductionist science that applies (...) physics and chemistry to answer how-questions but lacks a biological point of view of its own. In this paper I dispute this image of functional biology. A close look at the kinds of issues studied in biology and at the way in which these issues are studied shows that functional biology employs a distinctive biological perspective that is not rooted in selection. This functional perspective is characterized by its concern with the requirements of the life-state and the way in which these are met. (shrink)

This paper proposes an experiment-based methodology for both classical genetics and molecular biology by integrating Lindley Darden’s mechanism-centered approach and C. Kenneth Waters’s phenomenon-centered approach. We argue that the methodology basing on experiments offers a satisfactory account of the development of the two biological disciplines. The methodology considers discovery of new mechanisms, investigation of new phenomena, and construction of new theories together, in which experiments play a central role. Experimentation connects the three type of conduct, which work as both ends (...) and means, occurring in a circular way and constituting an overall process of scientific practice from classical genetics to molecular biology. (shrink)

In both biology and the human sciences, social groups are sometimes treated as adaptive units whose organization cannot be reduced to individual interactions. This group-level view is opposed by a more individualistic one that treats social organization as a byproduct of self-interest. According to biologists, group-level adaptations can evolve only by a process of natural selection at the group level. Most biologists rejected group selection as an important evolutionary force during the 1960s and 1970s but a positive literature began to (...) grow during the 1970s and is rapidly expanding today. We review this recent literature and its implications for human evolutionary biology. We show that the rejection of group selection was based on a misplaced emphasis on genes as “replicators” which is in fact irrelevant to the question of whether groups can be like individuals in their functional organization. The fundamental question is whether social groups and other higher-level entities can be “vehicles” of selection. When this elementary fact is recognized, group selection emerges as an important force in nature and what seem to be competing theories, such as kin selection and reciprocity, reappear as special cases of group selection. The result is a unified theory of natural selection that operates on a nested hierarchy of units.The vehicle-based theory makes it clear that group selection is an important force to consider in human evolution. Humans can facultatively span the full range from self-interested individuals to “organs” of group-level “organisms.” Human behavior not only reflects the balance between levels of selection but it can also alter the balance through the construction of social structures that have the effect of reducing fitness differences within groups, concentrating natural selection (and functional organization) at the group level. These social structures and the cognitive abilities that produce them allow group selection to be important even among large groups of unrelated individuals. (shrink)

In both biology and the human sciences, social groups are sometimes treated as adaptive units whose organization cannot be reduced to individual interactions. This group-level view is opposed by a more individualistic one that treats social organization as a byproduct of self-interest. According to biologists, group-level adaptations can evolve only by a process of natural selection at the group level. Most biologists rejected group selection as an important evolutionary force during the 1960s and 1970s but a positive literature began to (...) grow during the 1970s and is rapidly expanding today. We review this recent literature and its implications for human evolutionary biology. We show that the rejection of group selection was based on a misplaced emphasis on genes as "replicators" which is in fact irrelevant to the question of whether groups can be like individuals in their functional organization. The fundamental question is whether social groups and other higher-level entities can be "vehicles" of selection. When this elementary fact is recognized, group selection emerges as an important force in nature and what seem to be competing theories, such as kin selection and reciprocity, reappear as special cases of group selection. The result is a unified theory of natural selection that operates on a nested hierarchy of units. The vehicle-based theory makes it clear that group selection is an important force to consider in human evolution. Humans can facultatively span the full range from self-interested individuals to "organs" of group-level "organisms." Human behavior not only reflects the balance between levels of selection but it can also alter the balance through the construction of social structures that have the effect of reducing fitness differences within groups, concentrating natural selection (and functional organization) at the group level. These social structures and the cognitive abilities that produce them allow group selection to be important even among large groups of unrelated individuals. (shrink)

Former discussions of biological generalizations have focused on the question of whether there are universal laws of biology. These discussions typically analyzed generalizations out of their investigative and explanatory contexts and concluded that whatever biological generalizations are, they are not universal laws. The aim of this paper is to explain what biological generalizations are by shifting attention towards the contexts in which they are drawn. I argue that within the context of any particular biological explanation or investigation, biologists employ two (...) types of generations. One type identifies causal regularities exhibited by particular kinds of biological entities. The other type identifies how these entities are distributed in the biological world. (shrink)

Ecological theory is built upon assumptions about the fundamental nature of organism-environment interactions. We argue that two mutually exclusive sets of such assumptions are available and that they have given rise to alternative approaches to studying ecology. The fundamentally different premises of these approaches render them irreconcilable with one another. In this paper, we present the first logical formalisation of these two paradigms.The more widely-accepted approach - which we label the demographic paradigm - includes both population ecology and community ecology (...) (synecology). Demographic ecology assumes that the environment is relatively stable and that biotic processes, governed predominantly by resource availability, are the most important of ecological and evolutionary influences. Moreover, ecological processes are assumed to translate into directional selection pressures that drive significant evolutionary change on a local scale through the process of optimisation. (shrink)

In this paper, I wish to challenge theory-biased approaches to scientific knowledge, by arguing for a study of theorizing, as a cognitive activity, rather than of theories, as abstract structures independent from the agents’ understanding of them. Such a study implies taking into account scientists’ reasoning processes, and their representational practices. Here, I analyze the representational practices of geneticists in the 1910s, as a means of shedding light on the content of classical genetics. Most philosophical accounts of classical genetics fail (...) to distinguish between the purely genetic, or Mendelian level, and the cytological one. I distinguish between them by characterizing them in terms of their respective associated representational practices. I then present how the two levels were articulated within Morgan’s theory of crossing-over, and I describe the representational technique of linkage mapping, which embodies the “merging” of the Mendelian and cytological levels. I propose an analysis of the mapping scheme, as a means of enlightening the conceptual articulation of Mendelian and cytological hypotheses within classical genetics. Finally, I present the respective views of three opponents to Morgan in the 1910s, who had a different understanding of the articulation of cytology and Mendelism, and entertained different views concerning the role and proper interpretation of maps. I propose to consider these diverging perspectives as instantiating what I call different “versions” of classical genetics. (shrink)

One of the defining features of the classical gene was its position. In molecular genetics, positions are defined instead as nucleotide numbers and there is no clear correspondence with its classical counterpart. However, the classical gene position did not simply disappear with the development of the molecular approach, but survived in the lab associated to different genetic practices. The survival of classical gene position would illustrate Waters’ view about the practical persistence of the genetic approach beyond reductionism and anti-reductionist claims. (...) We show instead that at the level of laboratory practices there are also reductive processes, operating through the rise and fall of different techniques. Molecular markers made the concept of classical gene position practically dispensable, leading us to rethink whether it had any causal role or was just a mere heuristic. (shrink)

I claim that explanations of human behaviour by Edward O. Wilsonand Charles Lumsden are constituted by a religiously functioningmetaphysics: emergent materialism. The constitutive effects areidentified using six criteria, beginning with a metaphorical re-description of dissimilarities between levels of organization interms of the lower level, and consist of conceptual andexplanatory reductions (CER). Wilson and Lumsden practice CER,even though CER is not required by emergent materialism. Theypreconceive this practice by a re-description which conflates thelevels of organization and explain failure of CER in (...) terms oftechnical, not ontological or epistemological reasons. Iinterpret these three practices as a reaction of Wilson againsthis early Christian religious beliefs. Statements by Wilsonindicate this reaction ultimately constitutes his explanations ofsocial, moral and religious behaviour.Tested knowledge about matter at the lower level functions as ametaphysical belief when applied to the higher level becausethere it is untested. I offer twelve criteria for the diagnosisof religious functions of this metaphysical materialism, five ofwhich are satisfied. I show that the constitutive effects of thismaterialism in sociobiology are due to its religious functions,are beneficial for science and do not destroy its public nature. (shrink)

Rosenberg has rightly argued that fitness is supervenient. But he has wrongly assumed that this makes “The fittest survive” nontautologous. Supervenience makes strict reduction impossible. It sheds light on disputes concerning the testability of evolutionary theory.

Since the last decades of the twentieth and the beginning of the twenty-first century, the use of metaphysics by philosophers when approaching conceptual problems in biology has increased. Some philosophers call this tendency in philosophy of biology ‘Metaphysics of Biology’. In this paper, I aim at characterizing Metaphysics of Biology by paying attention to the diverse ways philosophers use metaphysics when addressing conceptual problems in biology. I will claim that there are two different modes of doing Metaphysics of Biology, namely (...) Metaphysics for Biology and Metaphysics in Biology. (shrink)

In addition to the distinction between species as a category and speciesas a taxon, the word species is ambiguous in a very different butequally important way, namely the temporal distinction between horizontal andvertical species. Although often found in the relevant literature, thisdistinction has thus far remained vague and undefined. In this paper the use ofthe distinction is explored, an attempt is made to clarify and define it, andthen the relation between the two dimensions and the implications of thatrelation are examined. (...) Using Darwin's analogy of language evolution forspeciesevolution, and by appealing to a major change in the conception of languagebetween 19th- and 20th-century linguistics, it is argued that the horizontaldimension has priority (pragmatic, epistemological, logical, and ontological)over the vertical dimension. This has immense ramifications for the modernspecies problem. Fundamentally, it favors horizontal species concepts oververtical ones. In so doing it places species realism on a much more securefoundation and largely undercuts species pluralism. In addition it raises aserious problem for the increasingly popular family of phylogenetic speciesconcepts, which generally suffer from a dimensionality confusion. However,thereis a recent trend within this family that attempts to restore the priority ofthe horizontal dimension. It is concluded that this trend should be affirmedandthat the species-as-individuals view should be abandoned. (shrink)

First, a brief history is provided of Popper's views on the status of evolutionary biology as a science. The views of some prominent biologists are then canvassed on the matter of falsifiability and its relation to evolutionary biology. Following that, I argue that Popper's programme of falsifiability does indeed exclude evolutionary biology from within the circumference of genuine science, that Popper's programme is fundamentally incoherent, and that the correction of this incoherence results in a greatly expanded and much more realistic (...) concept of what is empirical, resulting in the inclusion of evolutionary biology. Finally, this expanded concept of empirical is applied to two particular problems in evolutionary biology — viz., the species problem and the debate over the theory of punctuated equilibria — and it is argued that both of them are still mainly metaphysical. (shrink)

Gayon's recent claim that Buffon developed a concept of species as physical individuals is critically examined and rejected. Also critically examined and rejected is Gayon's more central thesis that as a consequence of his analysis of Buffon's species concept, and also of Darwin's species concept, it is clear that modern evolutionary theory does not require species to be physical individuals. While I agree with Gayon's conclusion that modern evolutionary theory does not require species to be physical individuals, I disagree with (...) his reasons and instead provide logical rather than historical reasons for the same conclusion. (shrink)

Biological type specimens are a particular kind of voucher specimen stored in natural history collections. Their special status and practical use are discussed in relation to the description and naming of taxonomic zoological diversity. Our current system, known as Linnaean nomenclature, is governed by the International Code of Zoological Nomenclature. The name of a species is fixed by its name-bearing type specimen, linking the scientific name of a species to the type specimen first designated for that species. The name-bearing type (...) specimen is not necessarily a typical example of the species, while establishment of the boundaries of a species requires empirical taxonomic studies. The International Code of Zoological Nomenclature allows for the naming of new species in the absence of preserved specimens. However, photos and DNA sequences should not function as primary type material, while new species should not be described and named without deposition of at least one type specimen in a collection. Philosophically, species are individuals, spatiotemporally restricted entities. Therefore, Linnaean species names are proper names, which do not define the taxon but serve as a label, providing an ostensive definition of a species. Paratypes have no name-bearing function but, nevertheless, are highly valued specimens in natural history collections. Paratypes should be restricted to those specimens originating from the same sample as the holotype. Diagnosis of a species taxon involves establishment of a connection between a Linnaean name and determination of the boundaries of the species. A first step in this process is the choice of an appropriate species concept. It is not the examination of holotypes and paratypes that necessarily provides the best estimate of the taxonomic boundaries of a species, but this is facilitated by a set of voucher specimens known as the hypodigm. Dissatisfaction with the present nomenclatural code led some researchers to propose emendations. Other taxonomists suggested abandoning Linnaean nomenclature and proposed the alternative PhyloCode, albeit that it relegates the naming of species taxa to the traditional nomenclatural codes. (shrink)

Controversies in science have a tendency to be long-lasting. Moreover, they tend to wither rather than be solved by sorting out the arguments pro and con. Barring the sociological dimension, an important factor in the perpetuation of scientific controversies seems to be the contestants' passion for broad philosophical theses when it comes to defending their respective positions. In this paper one such controversy is analysed. It involves the alleged use of Popperian falsificationism to defend a position in (community) ecology some (...) years ago. The upshot of the analysis is that falsificationism is altogether irrelevant to the controversy's solution; philosophy, though, is utterly relevant if one limits it to elementary, uncontroversial, normative methodological principles. (shrink)

At one level, this paper is a lament and a warning. I lament biologists borrowing well-known terms and then drastically and awkwardly changing their meanings, and I warn about the mischief this does. Biology''s public image is at stake, as is its general usefulness. At another level, I attempt to clarify the misnamed concepts, beyond what has been achieved in recent philosophical writings. This helps to account for the mischief, and to see how it might be avoidable. But the most (...) important thing about the paper is that, at a third level, it is an argument against physicalism and materialism, especially those variants which deny the autonomy of organisms and the existence of intrinsic goods. Interpreting biology from the point of view of those denials leads to unsatisfactory and even bizarre results. (shrink)

This paper examines the nature of theory structure in biology and considers the implications of those theoretical structures for theory reduction. An account of biological theories as interlevel prototypes embodying causal sequences, and related to each other by strong analogies, is presented, and examples from the neurosciences are provided to illustrate these middle-range theories. I then go on to discuss several modifications of Nagel''s classical model of theory reduction, and indicate at what stages in the development of reductions these models (...) might best apply. Finally I consider several implications of these analyses of theory structure and reduction for disciplinary integration in biology. (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)

This article is a systematic critical survey of work done in the philosophy of biology within the logical empiricist tradition, beginning in the 1930s and until the end of the 1950s. It challenges a popular view that the logical empiricists either ignored biology altogether or produced analyses of little value. The earliest work on the philosophy of biology within the logical empiricist corpus was that of Philipp Frank, Ludwig von Bertalanffy, and Felix Mainx. Mainx, in particular, provided a detailed analysis (...) of biology in the 1930s and 1940s in his contribution to the logical empiricists’ Encyclopedia of Unified Science. However, the most important contributions to the philosophy of biology were those of Joseph Henry Woodger and Ernest Nagel. Woodger is primarily remembered for deploying the axiomatic method in biology but he also used semiformal methods for the analysis of many biological problems. While Woodger’s axiomatic work was often derided by some later philosophers of biology (e.g., David Hull and Michael Ruse), this article defends both the biological and the philosophical significance of some of that work, for instance, those aspects that led to the recognition of the conceptual complexity of mereology and temporal identity in biological systems. Woodger’s semiformal analyses were even more important, for instance, his explication of the concepts of the Bauplan and of innateness. Nagel’s importance lies in his analyses of reduction and emergence in the context of all empirical sciences and his use of these analyses in a careful exploration of biological problems. While Nagel’s model of reduction was generally rejected by philosophers of science in the 1970s and 1980s, particularly for biological contexts, it has recently been sympathetically reconstructed by many commentators; this article defends its continued relevance for the philosophy of biology. (shrink)

This survey of major developments in North American philosophy of science begins with the mid-1960s consolidation of the disciplinary synthesis of internalist history and philosophy of science (HPS) as a response to criticisms of logical empiricism. These developments are grouped for discussion under the following headings: historical metamethodologies, scientific realisms, philosophies of the special sciences, revivals of empiricism, cognitivist naturalisms, social epistemologies, feminist theories of science, studies of experiment and the disunity of science, and studies of science as practice and (...) culture. A unifying theme of the survey is the relation between historical metamethodologists and scientific realists, which dominated philosophical work in the late 1970s. I argue that many of the alternative cognitive naturalisms, social epistemologies, and feminist theories that have been proposed can be understood as analogues to the differences between metamethodological theories of scientific rationality and realist accounts of successful reference to real causal processes. Recent work on experiment, scientific practice, and the culture of science may, however, challenge the underlying conception of the field according to which realism and historical rationalism (or their descendants) are the important alternatives available, and thus may take philosophy of science in new directions. (shrink)

This paper argues that the consensus physicalist antireductionism in the philosophy of biology cannot accommodate the research strategy or indeed the recent findings of molecular developmental biology. After describing Wolperts programmatic claims on its behalf, and recent work by Gehring and others to identify the molecular determinants of development, the paper attempts to identify the relationship between evolutionary and developmental biology by reconciling two apparently conflicting accounts of bio-function – Wrights and Nagels (as elaborated by Cummins). Finally, the paper seeks (...) a way of defending the two central theses of physicalist antireductionism in the light of the research program of molecular developmental biology, by sharply reducing their metaphysical force. (shrink)

In The Structure of Biological Science I argued that the theory of natural selection is a statistical theory for reasons much like those which makes thermodynamics a statistical theory. In particular, the theory claims that fitness differences are large enough and the life span of species long enough for increases in average fitness always to appear in the long run; and this claim, I held, is of the same form as the statistical version of the second law of thermodynamics.For the (...) latter law also makes a claim about the long run, and its statistical character is due to this claim: thermodynamic systems must in the long run approach an equilibrium level of organization that maximizes entropy. Over finite times, given local boundary conditions, an isolated mechanical system, like the molecules in a container of gas, may sometimes interact so as to move the entropy of the system further from, instead of closer to the equilbrium level. But given enough interacting bodies, and enough time, the system will always eventually move in the direction prescribed by the law. Thus, we can attach much higher probabilities to the prediction that non-equilibrium systems will reflect greater entropy in future periods than we can to predictions that they will move in the opposite direction. And as we increase the amount of time and the number of bodies interacting, the strength of the probability we can attach to the prediction becomes greater and greater. (shrink)

The correct explanation of why species, in evolutionary theory, are individuals and not classes is the cladistic species concept. The cladistic species concept defines species as the group of organisms between two speciation events, or between one speciation event and one extinction event, or (for living species) that are descended from a speciation event. It is a theoretical concept, and therefore has the virtue of distinguishing clearly the theoretical nature of species from the practical criteria by which species may be (...) recognized at any one time. Ecological or biological (reproductive) criteria may help in the practical recognition of species. Ecological and biological species concepts are also needed to explain why cladistic species exist as distinct lineages, and to explain what exactly takes place during a speciation event. The ecological and biological species concepts work only as sub-theories of the cladistic species concept and if taken by themselves independently of cladism they are liable to blunder. The biological species concept neither provides a better explanation of species indivudualism than the ecological species concept, nor, taken by itself, can the biological species concept even be reconciled with species individualism. Taking the individuality of species seriously requires subordinating the biological, to the cladistic, species concept. (shrink)