Darwin was a teleologist

It is often claimed that one of Darwin's chief accomplishments was to provide biology with a non-teleological explanation of adaptation. A number of Darwin's closest associates, however, and Darwin himself, did not see it that way. In order to assess whether Darwin's version of evolutionary theory does or does not employ teleological explanation, two of his botanical studies are examined. The result of this examination is that Darwin sees selection explanations of adaptations as teleological explanations. The confusion in the nineteenth century about Darwin's attitude to teleology is argued to be a result of Darwin's teleological explanations not conforming to either of the dominant philosophical justifications of teleology at that time. Darwin's explanatory practices conform well, however, to recent defenses of the teleological character of selection explanations.I would like to thank John Beatty, David Hull and one of this journal's readers for constructive comments on an earlier draft of this paper.     

Teleogy and genes

My aim in this paper is to quickly sketch a teleological approach to the problem of isolating the impact of genes on phenotypic characters. I begin by arguing that it is a mistake to think that there will be only one analysis of genetic input suitable for all theoretical interests. My principle focus is Richard Dawkins' argument for genic selectionism. I argue that a teleological analysis of genetic input is what Dawkins requires to establish the right kind of mapping of gene onto phenotype. This comes at a certain cost, however. Accepting the analysis will threaten Dawkins' claims about the teleogogical priority of gene over phenotype.     

Natural selection and the conditions for existence: representational vs. conditional teleology in biological explanation

Human intentional action, including the design and use of artifacts, involves the prior mental representation of the goal (end) and the means to achieve that goal. This representation is part of the efficient cause of the action, and thus can be used to explain both the action and the achievement of the end. This is intentional teleological explanation. More generally, teleological explanation that depends on the real existence of a representation of the goal (and the means to achieve it) can be called representational teleological explanation. Such explanations in biology can involve both external representations (e.g., ideas in the mind of God) and internal representations (souls, vital powers, entelechies, developmental programs, etc.). However, another type of explanation of intentional action (or any other process) is possible. Given that an action achieving a result occurs, the action can be explained as fulfilling the necessary conditions (means) for that result (end), and, reciprocally, the result explained by the occurrence of those necessary conditions. This is conditional teleological explanation. For organisms, natural selection is often understood metaphorically as the designer, intentionally constructing them for certain ends. Unfortunately, this metaphor is often taken rather too literally, because it has been difficult to conceive of another way to relate natural selection to the process of evolution. I argue that combining a conditional teleological explanation of organisms and of evolution provides such an alternative. This conditional teleology can be grounded in existence or survival. Given that an organism exists, we can explain its existence by the occurrence of the necessary conditions for that existence. This principle of the 'conditions for existence' was introduced by Georges Cuvier in 1800, and provides a valid, conditional teleological method for explaining organismal structure and behavior. From an evolutionary perspective, the conditions for existence are the range of boundary conditions within which the evolutionary process must occur. Moreover, evolutionary change itself can be subjected to conditional teleological explanation, because natural selection theory is primarily a theory about the relation between the conditions for the existence of organisms and the conditions for the existence of traits in populations. I show that failure to distinguish representational from conditional teleological explanation has confused previous attempts to clarify the relation of teleology to biology.     

Theses on Darwin.

The received view that teleology has been successfully eliminated from the modern scientific worldview is challenged. It is argued that both the theory of natural selection and molecular biology presuppose the existence of natural teleology, and so cannot explain it. A number of other issues in the foundations of biology are briefly examined, while stress is laid throughout on empirical evidence of the rational agency inherent in life. It is urged that teleology be rehabilitated and that the reigning functionalist philosophy be replaced by a realistic view of biological functions as emergent properties of living matter within a broad, selforganization framework.     

Ethics in Darwin's melancholy vision

Darwinian natural selection draws on Malthus' harsh vision of human society to explain how organisms come to be adapted to their environments. Natural selection produces the appearance of teleology, but requires only efficient causal processes: undirected, heritable variation combined with effects of the variations on survival and reproduction. This paper draws a sharp distinction between the resulting form of backwards-directed teleology and the future-directed teleology we ascribe to intentional human activity. Rather than dismiss teleology as mere illusion, the paper concludes with an account of how future-directed teleology came to be a justifiable part of how we understand ourselves.     

Teleology and its constitutive role for biology as the science of organized systems in nature

'Nothing in biology makes sense, except in the light of teleology'. This could be the first sentence in a textbook about the methodology of biology. The fundamental concepts in biology, e.g. 'organism' and 'ecosystem', are only intelligible given a teleological framework. Since early modern times, teleology has often been considered methodologically unscientific. With the acceptance of evolutionary theory, one popular strategy for accommodating teleological reasoning was to explain it by reference to selection in the past: functions were reconstructed as 'selected effects'. But the theory of evolution obviously presupposes the existence of organisms as organized and regulated, i.e. functional systems. Therefore, evolutionary theory cannot provide the foundation for teleology. The underlying reason for the central methodological role of teleology in biology is not its potential to offer particular forms of (evolutionary) explanations for the presence of parts, but rather an ontological one: organisms and other basic biological entities do not exist as physical bodies do, as amounts of matter with a definite form. Rather, they are dynamic systems in stable equilibrium; despite changes of their matter and form (in metabolism and metamorphosis) they maintain their identity. What remains constant in these kinds of systems is their 'organization', i.e. the causal pattern of interdependence of parts with certain effects of each part being relevant for the working of the system. Teleological analysis consists in the identification of these system-relevant effects and at the same time of the system as a whole. Therefore, the identity of biological systems cannot be specified without teleological reasoning.     

Kant on biological teleology: Towards a two-level interpretation

According to Kant's Critique of the power of judgment, teleological considerations are unavoidable for conceptualizing organisms. Does this mean that teleology is more than merely heuristic? Kant stresses the regulative status of teleological attributions, but sometimes he seems to treat teleology as a constitutive condition for biology. To clarify this issue, the concept of natural purpose and its role for biology are examined. I suggest that the concept serves an identificatory function: it singles out objects as natural purposes, whereby the special science of biology is constituted. This relative constitutivity of teleology is explicated by means of a distinction of levels: on the object level of biological science, teleology is taken as constitutive, though it is merely regulative on the philosophical meta level. This distinction also concerns the place of Aristotelian teleology in Kant: on the object level, the Aristotelian view is accepted, whereas on the meta level, an agnostic stance is taken concerning teleology.     

Children’s Intuitive Teleology: Shifting the Focus of Evolution Education Research

Research has shown that children usually provide teleological explanations for the features of organisms and artifacts, from a very early age (3–4 years old). However, there is no consensus on whether teleological explanations are given in the same manner for non-living natural objects as well. The present study aimed to document the teleological explanations of 5- to 8-year-old children for particular features (color and shape) of organisms, artifacts and non-living natural objects. In addition, it was examined if there was any correlation between these explanations and children’s explanations for the usefulness of those features. Our results indicate a developmental shift in children’s teleological explanations, from a non-selective teleology in pre-school to a selective one in the second grade. In the latter case, children provided teleological explanations mostly for the shape of the feet of organisms and for the shape of artifacts, whereas pre-school children provided teleological explanations for non-living natural objects as well, both for the color and for the shape in all cases. Our results are not conclusive and further research is required, including a larger spectrum of students, since teleology is one of the most important conceptual obstacles in understanding evolution that persists even into adulthood. We conclude by proposing a particular research program for this purpose.     

Chemical Ecology and Biomolecular Evolution

The belief in the Darwinian theory of evolution appeared to be shaken when one tried to interpret statements of molecular biology in it. As a consequence there arose a theory of non-Darwinian neutral evolution. The supporters of this theory believe that under natural conditions no factors exist which can distinguish and select organisms on their internal (molecular) structure. In the opinion of these neutralists natural selection cannot in principle control the molecular constitution of organisms. Contrary to the viewpoint of the critics of neutralism it is impossible to admit that nucleic acids, proteins and other biomolecules can evolve without the participation of natural selection. This controversy in contemporary theoretical biology can be solved by integrating the conceptions of molecular ecology with Darwinian theory. Molecular ecology acknowledges the interactions of organisms by means of chemical substances synthesized by them. Such chemical ecological factors play a leading part in the selective stages of biomolecular evolution. These diverse chemical ecological interrelations take place intensively when living beings interact with parasitic microbes.     

Darwin's changing views on evolution: from centres of origin and teleology to vicariance and incomplete lineage sorting

It is a strange fact that in many ways the first edition of Charles Darwin's Origin of Species is closer to modern neodarwinism than the sixth and last edition. Sometimes this is attributed to a decline in the quality of the argument, but the opposite interpretation is given here. It is suggested that Darwin's early work on evolution is naïve and based on the two creationist principles of centre of origin and teleology (panselectionism). This fusion later became the 'modern synthesis'. However, after the first edition of the Origin , Darwin developed a non-teleological synthesis that integrated natural selection with what he called 'laws of growth'– phylogenetic/morphogenetic trends or tendencies. Discussion of Darwin's later, more sophisticated model of evolution has been suppressed in the teleological modern synthesis, but similar ideas are re-emerging in current work on molecular phylogenetics and biogeography. This indicates that the ancestor of a group can be diverse in its morphology and its ecology, that this diversity can be inherited, and that groups usually originate over a broad region and not at a single point.     

The Replicator in Retrospect

The history and theoretical role of the concept of a ``replicator'is discussed, starting with Dawkins' and Hull's classic treatmentsand working forward. I argue that the replicator concept is still auseful one for evolutionary theory, but it should be revised insome ways. The most important revision is the recognition that notall processes of evolution by natural selection require thatsomething play the role of a replicator.     

The Extended Synthesis: The Law of the Conditions of Existence

A unified theory of biology must incorporate a naturalistic explanation for the origin of life, namely that given certain conditions, it was highly probable that life would originate. That theory, however, cannot be solely a theory of the origin of life. The same general mechanisms that allowed life to originate must also explain its subsequent evolution as embodied in the Extended Synthesis, including the origin of the inherent constraints and regularities that allowed natural selection to emerge as a natural process. A naturalistic grounding for the origin of life and its subsequent evolution, what Darwin called the Law of the Conditions of Existence, can be found in the natural law of history, the Second Law of Thermodynamics.     

Cultural transmission and the evolution of human behaviour: a general approach based on the Price equation

Transmitted culture can be viewed as an inheritance system somewhat independent of genes that is subject to processes of descent with modification in its own right. Although many authors have conceptualized cultural change as a Darwinian process, there is no generally agreed formal framework for defining key concepts such as natural selection, fitness, relatedness and altruism for the cultural case. Here, we present and explore such a framework using the Price equation. Assuming an isolated, independently measurable culturally transmitted trait, we show that cultural natural selection maximizes cultural fitness, a distinct quantity from genetic fitness, and also that cultural relatedness and cultural altruism are not reducible to or necessarily related to their genetic counterparts. We show that antagonistic coevolution will occur between genes and culture whenever cultural fitness is not perfectly aligned with genetic fitness, as genetic selection will shape psychological mechanisms to avoid susceptibility to cultural traits that bear a genetic fitness cost. We discuss the difficulties with conceptualizing cultural change using the framework of evolutionary theory, the degree to which cultural evolution is autonomous from genetic evolution, and the extent to which cultural change should be seen as a Darwinian process. We argue that the nonselection components of evolutionary change are much more important for culture than for genes, and that this and other important differences from the genetic case mean that different approaches and emphases are needed for cultural than genetic processes.     

Darwin teleologist? Design in the Orchids

Focusing on the Orchids, this article aims at disentangling the concepts of teleology, design and natural theology. It refers to several contemporary critics of Darwin (Kölliker, Argyll, Royer, Candolle, Delpino) to challenge Huxley's interpretation that Darwin's system was “a deathblow” to teleology. The Orchids seem rather to be a “flank-movement” (Gray): it departs from the Romantic theories of transmutation and the “imaginary examples” of the Origin; it focuses on empirical data and on teleological structures. Although Darwin refers to natural selection, his readers mock him for his fascination for delicate morphological contrivances and co-adaptations – a sign that he was inescapably lured to finality. Some even suggested that his system was a “theodicy”. In the history of Darwinism, the Orchids reveal “another” quite unexpected and heterodox Darwin: freed from the hypothetical fancies of the Origin, and even suggesting a new kind of physico-theology.     

The Role of Teleological Thinking in Learning the Darwinian Model of Evolution

Human beings are predisposed to think of evolution as teleological—i.e., having a purpose or directive principle—and the ways scientists talk about natural selection can feed this predisposition. This work examines the suggestion that students’ teleological thinking operates as an obstacle when the natural selection evolution model is taught. What we mean by obstacle is an established way of thinking that resists change due to its explanatory power. In light of this approach, the challenges of teaching evolution in biology education have been revised, and improved methodological strategies aimed at a better comprehension of the Darwinian evolution model are suggested.     

August Weismann's concept of germ plasma as the basic reason for the inadequacy of neo-Darwinism

Neo-Darwinism is a result of synthesis of Darwinian concept of natural selection with Weismannian concept of germ plasma. The concept of germ plasma is based on a hypothesis that phenotypic traits are completely determined by genes. Hence, neo-Darwinism describes evolution as a process of alternation of gene frequencies under the effect of natural selection. This is an inadequate approach to the study of evolution. In the course of evolution, genes change their functions, whereas phenotypic characters change their corresponding genes. As a result, every step of evolutionary transformation changes the structure of phenotype-to-genotype correspondence. Therefore, phenotypic evolution cannot be described in genetic terms, the same as to human languages cannot be translated one into another whenever the meaning of words is constantly changing. Consequently, Weismannian germ-plasma concept adequately describes the relation of characters to genes only during stasis, but is inapplicable to evolution.     

The insufficience of supervenient explanations of moral actions: Really taking Darwin and the naturalistic fallacy seriously

In a recent paper in this journal (Rottschaefer and Martinsen 1990) we have proposed a view of Darwinian evolutionary metaethics that we believe improves upon Michael Ruse's (e.g., Ruse 1986) proposals by claiming that there are evolutionary based objective moral values and that a Darwinian naturalistic account of the moral good in terms of human fitness can be given that avoids the naturalistic fallacy in both its definitional and derivational forms while providing genuine, even if limited, justifications for substantive ethical claims. Jonathan Barrett (this issue) has objected to our proposal contending that we cannot hold for the reality of supervenient moral properties without either falling foul of the naturalistic fallacy or suffering the consequences of postulating inexplicable moral properties. In reply, we show that Barrett's explicit arguments that we commit either the definitional or derivational form of the naturalistic fallacy fail and that his naturalistic intuitions that supervenience explanations of moral properties by nonmoral properties force us into what we call the explanatory form of the naturalistic fallacy also fail. Positively, his objections help us to clarify the nature of the naturalistic fallacy within an evolutionary based naturalistic ethics and to point out the proper role of both supervenience explanations and moral explanations in such an ethics.     

The genic view of the process of speciation

The unit of adaptation is usually thought to be a gene or set of interacting genes, rather than the whole genome, and this may be true of species differentiation. Defining species on the basis of reproductive isolation (RI), on the other hand, is a concept best applied to the entire genome. The biological species concept (BSC; 84 ) stresses the isolation aspect of speciation on the basis of two fundamental genetic assumptions – the number of loci underlying species differentiation is large and the whole genome behaves as a cohesive, or coadapted genetic unit. Under these tenets, the exchange of any part of the genomes between diverging groups is thought to destroy their integrity. Hence, the maintenance of each species’ genome cohesiveness by isolating mechanisms has become the central concept of species. In contrast, the Darwinian view of speciation is about differential adaptation to different natural or sexual environments. RI is viewed as an important by product of differential adaptation and complete RI across the whole genome need not be considered as the most central criterion of speciation. The emphasis on natural and sexual selection thus makes the Darwinian view compatible with the modern genic concept of evolution. Genetic and molecular analyses of speciation in the last decade have yielded surprisingly strong support for the neo‐Darwinian view of extensive genetic differentiation and epistasis during speciation. However, the extent falls short of what BSC requires in order to achieve whole‐genome ‘cohesiveness’. Empirical observations suggest that the gene is the unit of species differentiation. Significantly, the genetic architecture underlying RI, the patterns of species hybridization and the molecular signature of speciation genes all appear to support the view that RI is one of the manifestations of differential adaptation, as 34 , Chap. 8) suggested. The nature of this adaptation may be as much the result of sexual selection as natural selection. In the light of studies since its early days, BSC may now need a major revision by shifting the emphasis from isolation at the level of whole genome to differential adaptation at the genic level. With this revision, BSC would in fact be close to Darwin’s original concept of speciation.     

Evolution of adaptive phenotypic variation patterns by direct selection for evolvability

A basic assumption of the Darwinian theory of evolution is that heritable variation arises randomly. In this context, randomness means that mutations arise irrespective of the current adaptive needs imposed by the environment. It is broadly accepted, however, that phenotypic variation is not uniformly distributed among phenotypic traits, some traits tend to covary, while others vary independently, and again others barely vary at all. Furthermore, it is well established that patterns of trait variation differ among species. Specifically, traits that serve different functions tend to be less correlated, as for instance forelimbs and hind limbs in bats and humans, compared with the limbs of quadrupedal mammals. Recently, a novel class of genetic elements has been identified in mouse gene-mapping studies that modify correlations among quantitative traits. These loci are called relationship loci, or relationship Quantitative Trait Loci (rQTL), and affect trait correlations by changing the expression of the existing genetic variation through gene interaction. Here, we present a population genetic model of how natural selection acts on rQTL. Contrary to the usual neo-Darwinian theory, in this model, new heritable phenotypic variation is produced along the selected dimension in response to directional selection. The results predict that selection on rQTL leads to higher correlations among traits that are simultaneously under directional selection. On the other hand, traits that are not simultaneously under directional selection are predicted to evolve lower correlations. These results and the previously demonstrated existence of rQTL variation, show a mechanism by which natural selection can directly enhance the evolvability of complex organisms along lines of adaptive change.     

The function of function

Contemporary analyses of biological function almost invariably advocate a naturalistic analysis, grounding biological functions in some feature of the mind-independent world. Many recent accounts suggest that no single analysis will be appropriate for all cases of use and that biological teleology should be split into several distinct categories. This paper argues that such accounts have paid too little attention to the way in which functional language is used, concentrating instead on the types of situation in which it is used. An example of the role of teleology in science is examined and, on the basis of conclusions drawn from this, an alternative unifying analysis is proposed. It is suggested that, contrary to naturalistic accounts, teleology in biology carries no ontological commitment whatsoever to any class of mind-independent entities or properties. Instead, it is best regarded as a methodological device which is used to focus interest, formulate research perspectives and facilitate the structuring of certain questions or types of question that are pertinent in a given context of interest.