The Mastodon in the room: how Darwinian is neo-Darwinism?

Failing to acknowledge substantial differences between Darwinism and neo-Darwinism impedes evolutionary biology. Darwin described evolution as the outcome of interactions between the nature of the organism and the nature of the conditions, each relatively autonomous but both historically and spatially intertwined. Furthermore, he postulated that the nature of the organism was more important than the nature of the conditions, leading to natural selection as an inevitable emergent product of biological systems. The neo-Darwinian tradition assumed a creative rather than selective view of natural selection, with the nature of the organism determined by the nature of the conditions, rendering the nature of the organism and temporal contingency unnecessary. Contemporary advances in biology, specifically the phylogenetics revolution and evo-devo, underscore the significance of history and the nature of the organism in biology. Darwinism explains more biology better, and better resolves apparent anomalies between living systems and more general natural laws, than does neo-Darwinism. The "extended" or "expanded" synthesis currently called for by neo-Darwinians is Darwinism.     

Confirmation and explaining how possible

Confirmation in evolutionary biology depends on what biologists take to be the genuine rivals. Investigating what constrains the scope of biological possibility provides part of the story: explaining how possible helps determine what counts as a genuine rival and thus informs confirmation. To clarify the criteria for genuine rivalry I distinguish between global and local constraints on biological possibility, and offer an account of how-possibly explanation. To sharpen the connection between confirmation and explaining how possible I discuss the view that formal inquiry can provide a kind of confirmation-theoretic support for evolutionary models, and offer an example of how-possibly explanation interacting with testing practice.     

How-possibly explanations as genuine explanations and helpful heuristics: A comment on Forber

Recently, Forber introduced a distinction between two kinds of how-possibly explanation, global and local how-possibly explanation, and argued that both play genuinely explanatory roles in evolutionary biology. In this paper I examine the nature of these two kinds of how-possibly explanations, focusing on the question whether they indeed constitute genuine explanations. I will conclude that one of Forber's kinds of how-possibly explanation may be thought of as a kind of genuine explanation but not as a kind of how-possibly explanation, while the other kind plays a heuristic role and should not be conceived of as a kind of explanation at all.     

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.     

Constraining the adaptationism debate

This contribution to the adaptationism debate elaborates the nature of constraints and their importance in evolutionary explanation and argues that the adaptationism debate should be limited to the issue of how to privilege causes in evolutionary explanation. I argue that adaptationist explanations are deeply conceptually dependent on developmental constraints, and explanations that appeal to constraints are dependant on the results of natural selection. I suggest these explanations should be integrated into the framework of historical causal explanation. Each strategy explicitly appeals to some aspect of the evolutionary process, while implicitly appealing to others. Thus, adaptationists and anti-adaptationists can offer complementary causal explanations of the same explanandum. This eliminates much of the adaptationism debate and explains why its adversaries regularly agree with each other more than they would like. The adaptationism issue that remains is a species of the general issue of how to privilege causes in explanation. I show how a proposed solution to this general problem might be brought to bear on evolutionary explanations, and investigate some difficulties that might arise due to the nature of the evolutionary process.     

Individual essentialism in biology

A few philosophers of biology have recently explicitly rejected Essential Membership, the doctrine that if an individual organism belongs to a taxon, particularly a species, it does so essentially. But philosophers of biology have not addressed the broader issue, much discussed by metaphysicians on the basis of modal intuitions, of what is essential to the organism. In this paper, I address that issue from a biological basis, arguing for the Kripkean view that an organism has a partly intrinsic, partly historical, essence. The arguments appeal to the demands of biological explanation and are analogous to arguments that I have given elsewhere that a taxon has a partly intrinsic, partly historical, essence. These conclusions about the essences of individuals and taxa yield an argument for Essential Membership. Finally, I cast doubt on LaPorte’s objection to that doctrine arising from the view that a species cannot survive having a daughter.     

Fitness and Propensity’s Annulment?

Recent debate on the nature of probabilities in evolutionary biology has focused largely on the propensity interpretation of fitness (PIF), which defines fitness in terms of a conception of probability known as “propensity”. However, proponents of this conception of fitness have misconceived the role of probability in the constitution of fitness. First, discussions of probability and fitness have almost always focused on organism effect probability, the probability that an organism and its environment cause effects. I argue that much of the probability relevant to fitness must be organism circumstance probability, the probability that an organism encounters particular, detailed circumstances within an environment, circumstances which are not the organism’s effects. Second, I argue in favor of the view that organism effect propensities either don’t exist or are not part of the basis of fitness, because they usually have values close to 0 or 1. More generally, I try to show that it is possible to develop a clearer conception of the role of probability in biological processes than earlier discussions have allowed.     

Homology and the hierarchy of biological systems

Homology is the similarity between organisms due to common ancestry. Introduced by Richard Owen in 1843 in a paper entitled "Lectures on comparative anatomy and physiology of the invertebrate animals", the concept of homology predates Darwin's "Origin of Species" and has been very influential throughout the history of evolutionary biology. Although homology is the central concept of all comparative biology and provides a logical basis for it, the definition of the term and the criteria of its application remain controversial. Here, I will discuss homology in the context of the hierarchy of biological organization. I will provide insights gained from an exemplary case study in evolutionary developmental biology that indicates the uncoupling of homology at different levels of biological organization. I argue that continuity and hierarchy are separate but equally important issues of homology.     

Does Biology Need an Organism Concept?

Among biologists, there is no general agreement on exactly what entities qualify as ‘organisms’. Instead, there are multiple competing organism concepts and definitions. While some authors think this is a problem that should be corrected, others have suggested that biology does not actually need an organism concept. We argue that the organism concept is central to biology and should not be abandoned. Both organism concepts and operational definitions are useful. We review criteria used for recognizing organisms and conclude that they are not categorical but rather continuously variable. Different organism concepts are useful for addressing different questions, and it is important to be explicit about which is being used. Finally, we examine the origins of the derived state of organismality, and suggest that it may result from positive feedback between natural selection and functional integration in biological entities.     

Kant's concept of natural purpose and the reflecting power of judgement

This paper examines how in the 'Critique of teleological judgment' Kant characterized the concept of natural purpose in relation to and in distinction from the concepts of nature and the concept of purpose he had developed in his other critical writings. Kant maintained that neither the principles of mechanical science nor the pure concepts of the understanding through which we determine experience in general provide adequate conceptualizations of the unique capacities of organisms. He also held that although the concept of natural purpose was derived through reflection upon an analogy to human purposive activity in artistic production and moral action, it articulates a unique notion of intrinsic purposiveness. Kant restricted his critical reflections on organisms to phenomena that can be given to us in experience, criticizing speculations on their first origins or final purpose. But I argue that he held that the concept of natural purpose is a product of the reflecting power of judgment, rather than an empirical concept, and represents only the relation of things to our power of judgment. Yet it is necessary for the identification of organisms as organized and self-organizing, and as subject to unique norms and causal relations between parts and whole.     

Organisms in evolution

Organisms constitute one of the most remarkable features of our living world. However, they have not yet received any accepted characterization within the framework of the evolutionary theory. The reasons for this contrast between the saliency of organisms in the biological landscape and their theoretical status are multiple and they are analyzed in the first part of this paper. Starting from this contrast, I argue for a theoretically grounded concept of organism within the framework of evolutionary theory itself. To this effect I argue that the theory of major transitions in evolution (Maynard Smith and Szathmáry 1995; Michod 1999) provides us with the theoretical basis for an understanding of the individuality of organisms and I propose a first characterization of organisms as evolutionary units structured by a division of reproductive labor among their parts. I also discuss one of the most important implications of this definition, namely that some colonial entities are to be counted as superorganisms. Finally, I show that though theoretically satisfying, this definition does not suffice in order fully to individuate the organisms and superorganisms in practice. To this end, physiology is needed, because it offers us some criteria for their individuation in ecological space. These criteria, however, are not immune to errors through misidentification and their shortcomings are discussed in the last section. In conclusion, I emphasize the positive implications of these criteria concerning the ecological significance of organisms.     

Conjecture and explanation: A reply to Reydon

Reydon (2012) comments on my account of how-possibly explanation (Forber, 2010). I distinguish between three types of explanation (global how-possibly, local how-possibly, and how actually) and argue that these distinctions track various roles explanations play in evolutionary biology. While Reydon accepts the distinctions, he questions whether the two different types of how-possibly explanation count as genuine explanations. He summarizes his analysis with a slogan: “global how-possibly explanations are explanations but not how-possibly; local explanations are how-possibly but not explanations.” Reydon’s commentary raises a number of insightful points, and I will not be able to address them all. Instead, after clarifying certain points in my original paper (4 1), I will respond to Reydon’s slogan by addressing whether global how-possibly explanations should count as explaining how possible (4 2), and what (so-called) local how-possibly explanations are, if not explanations (4 3).     

The risk of flawed inference in evolutionary studies when detectability is less than one

Addressing evolutionary questions in the wild remains a challenge. It is best done by monitoring organisms from birth to death, which is very difficult in part because individuals may or may not be resighted or recaptured. Although the issue of uncertain detection has long been acknowledged in ecology and conservation biology, in evolutionary studies of wild populations it is often assumed that detectability is perfect. We argue that this assumption may lead to flawed inference. We demonstrate that the form of natural selection acting on body mass of sociable weavers is altered and that the rate of senescence of roe deer is underestimated when not accounting for a value of detectability that is less than one. Because mark-recapture models provide an explicit way to integrate and reliably model the detection process, we strongly recommend their use to address questions in evolutionary biology.     

Causes and consequences of excess resistance in cryptobiotic metazoans

Despite more than 200 yr of recognition that some microscopic metazoans survive environmental conditions far beyond those experienced in nature while in a cryptobiotic state, this phenomenon has received little attention from evolutionary biologists. The excess environmental resistance exhibited by cryptobiotic organisms cannot be viewed as an adaptation within current evolutionary biology. Rather, excess resistance may have evolved as a by-product of natural selection for tolerance to desiccation or other naturally occurring environmental agents. The combined effects of desiccation, metabolic arrest, effective stabilization of dry or frozen cells by protectant molecules, and efficient DNA repair mechanisms may have led to a protection of the organism against conditions far beyond those experienced in nature.     

Kant’s concept of natural purpose and the reflecting power of judgement

This paper examines how in the ‘Critique of teleological judgment’ Kant characterized the concept of natural purpose in relation to and in distinction from the concepts of nature and the concept of purpose he had developed in his other critical writings. Kant maintained that neither the principles of mechanical science nor the pure concepts of the understanding through which we determine experience in general provide adequate conceptualizations of the unique capacities of organisms. He also held that although the concept of natural purpose was derived through reflection upon an analogy to human purposive activity in artistic production and moral action, it articulates a unique notion of intrinsic purposiveness. Kant restricted his critical reflections on organisms to phenomena that can be given to us in experience, criticizing speculations on their first origins or final purpose. But I argue that he held that the concept of natural purpose is a product of the reflecting power of judgment, rather than an empirical concept, and represents only the relation of things to our power of judgment. Yet it is necessary for the identification of organisms as organized and self-organizing, and as subject to unique norms and causal relations between parts and whole.     

``Two' Many Optimalities

In evolutionary biology, a trait is said to be optimal if it maximizes the fitness of the organism, that is, if the trait allows the organism to survive and reproduce better than any other competing trait would. In engineering, a design is said to be optimal if it complies with its functional requirements as well as possible. Cognitive science is both a biological and engineering discipline and hence it uses both notions of optimality. Unfortunately, the lack of a clear methodological stance on this tissue has made it common for researchers to conflate these two kinds of optimality. In this paper, I argue that a strict distinction must kept in order to avoid inaccurate assumptions.     

Do Organisms Exist?

What is the status of organisms in modern evolutionary biology?I argue that this is a question which centers on the questionof reduction, and towards a complete answer, I pursue issuesthrough three different senses of the term: ontological, methodological,and epistemological. The first sense refers to the ultimatestatus of the entities of the organic world, and in this senseI argue that organisms have no special status. The second senserefers to the question of organization, and I argue that inthe light of modern evolutionary biology organisms do have adistinctive "design-like" organization. The third sense refersto the relationship between theories, in particular to whetherthe theories of the biological sciences can be shown to be logicalconsequences of the theories of the physical sciences. I arguethat such reduction may be possible in principle but difficultin practice. However, from the perspective of the working scientist,this hardly matters. In conclusion, I argue that in some respectsorganisms are not distinctive and in other respects they are.Certainly biologists need not worry for the autonomy of theirsubject.     

Darwin's artificial selection as an experiment

Darwin used artificial selection (ASN) extensively and variedly in his theorizing. Darwin used ASN as an analogy to natural selection; he compared artificial to natural varieties, hereditary variation in nature to that in the breeding farm; and he also compared the overall effectiveness of the two processes. Most historians and philosophers of biology have argued that ASN worked as an analogical field in Darwin's theorizing. I will argue rather that this provides a limited and somewhat muddled view of Darwinian science. I say "limited" because I will show that Darwin also used ASN as a complex experimental field. And I say "muddled" because, if we concentrate on the analogical role exclusively, we conceive Darwinian science as rather disconnected from contemporary conceptions of "good science". I will argue that ASN should be conceived as a multifaceted experiment. As a traditional experiment, ASN established the efficacy of Darwin's preferred cause: natural selection. As a non-traditional experiment, ASN disclosed the nature of a crucial element in Darwin's evolutionary mechanics: the nature of hereditary variation. Finally, I will argue that the experiment conception should help us make sense of Darwin's comments regarding the "monstrous" nature of domestic breeds traditionally considered to be problematic.     

The cell theory--history, present state and prospects (on the 150th anniversary of the development of the cell theory)

The main stages of history of this most important biological conception are presented and the state of the modern cell theory and its future prospects are considered. Since 1839, when T. Schwann expounded his conception of the cell, a long pathway in cognition of the cell function and organization has been covered. From the original picture of the complex organism as a "cellular state", made up of relatively independent "elementary organisms", i.e. cells the modern biology has come to the idea of the cell as an integral system either being a part of a complex organism, or living free in the nature (protists). The cell represents certain qualitatively peculiar level in a complex evolutionary established hierarchy of biological systems. Some particular tight relations, existing between cytology, as a fundamental biological science and molecular biology, genetics, ecology and other biological disciplines are considered. The importance of the cell conception is ascertained for practical aims, especially in medicine.