I propose that, in addition to the commonly recognized increase of entropy, two more time arrows influence living beings. The increase of damage reactions, which produce aging and genetic variation, and the decrease of the rate of entropy production involved in natural selection are neglected arrows of time. Although based on the statistical theory of the arrow of time, they are distinguishable from the general arrow of the increase of entropy. Physiology under healthy conditions only obeys the increase of entropy (...) arrow. But aging, death, and evolution are determined by the other time arrows as well. Paradoxes emerge from conflicts among the specific determinisms associated with each arrow. These conflicts, along with uncertainties intrinsic to the low-number statistics of the arrows of damage reactions and decrease of the rate of production of entropy, highlight the limits of determinism at different levels of biology and its dependence on time span and number of organisms. The recognition of the three time arrows opens new perspectives for the problem of compatibility of the flow of time in physics and psychology. (shrink)

Philosophers of biology have worked extensively on how we ought best to interpret the probabilities which arise throughout evolutionary theory. In spite of this substantial work, however, much of the debate has remained persistently intractable. I offer the example of Bayesian models of divergence time estimation as a case study in how we might bring further resources from the biological literature to bear on these debates. These models offer us an example in which a number of different sources of uncertainty (...) are combined to produce an estimate for a complex, unobservable quantity. These models have been carefully analyzed in recent biological work, which has determined the relationship between these sources of uncertainty, both quantitatively and qualitatively. I suggest here that this case shows us the limitations of univocal analyses of probability in evolution, as well as the simple dichotomy between “subjective” and “objective” probabilities, and I conclude by gesturing toward ways in which we might introduce more sophisticated interpretive taxonomies of probability as a path toward advancing debates on probability in the life sciences. (shrink)

Julian Huxley’s eclipse of Darwinism narrative has cast a long shadow over the historiography of evolutionary theory around the turn of the nineteenth century. It has done so by limiting who could be thought of as Darwinian. Peter Bowler used the eclipse to draw attention to previously understudied alternatives to Darwinism, but maintained the same flaw. In his research on the Non-Darwinian Revolution, he extended this problematic element even further back in time. This paper explores how late nineteenth-century neo-Darwinian conceptualizations (...) of Darwinism were later utilized by several advocates and detractors of the Modern Synthesis. John Beatty has shown how this continuity hinges at least partly on the perceived importance of the creativity of natural selection. The paper provides a more thorough look at Darwin’s two conflicting accounts of variation, ascribed to struggles in explaining quantitative versus qualitative characters. In doing so, it suggests that other forms of Darwinism persisted, in both the non-Darwinian revolution and eclipse periods, because of tension between contingency and creativity in Darwin’s own work. This tension is traced out from Darwin’s conceptions of variation into the work of Alfred Russel Wallace, Hugo de Vries, and Thomas Henry Huxley. Based on this, the eclipse narrative is criticized for not considering the meaning of Darwinism in different geographical locations. Britain and the United States showed few signs of an eclipse. Rather, the rise of German debates about Haeckel’s vision of Darwinism have been mistaken for a universal decline in support. (shrink)

A key issue in the philosophy of biology is evolutionary contingency, the degree to which evolutionary outcomes could have been different. Contingency is typically contrasted with evolutionary convergence, where different evolutionary pathways result in the same or similar outcomes. Convergences are given as evidence against the hypothesis that evolutionary outcomes are highly contingent. But the best available treatments of contingency do not, when read closely, produce the desired contrast with convergence. Rather, they produce a picture in which any degree of (...) contingency is compatible with any degree of convergence. This is because convergence is the repeated production of a given outcome from different starting points, and contingency has been defined without reference to the size of the space of possible outcomes. In small spaces of possibilities, the production of repeated outcomes is almost assured. This paper presents a definition of contingency which includes this modal dimension in a way that does not reduce it to the binary notion of contingency found in standard modal logic. The result is a conception of contingency which properly contrasts with convergence, given some domain of possibilities and a measure defined over it. We should therefore not ask whether evolution is contingent or convergent simpliciter, but rather about the degree to which it is contingent or convergent in various domains, as measured in various ways. (shrink)

In this essay, I examine the usage of the term “just-so story.” I attempt to show that just-so storytelling can be seen as an epistemic concept that, in various ways, tackles the epistemological an...

Much of Stephen Jay Gould’s legacy is dominated by his views on the contingency of evolutionary history expressed in his classic Wonderful Life. However, Gould also campaigned relentlessly for a “nomothetic” paleontology. How do these commitments hang together? I argue that Gould’s conception of science and natural law combined with his commitment to contingency to produce an evolutionary science centered around the formulation of higher-level evolutionary laws.

Evolutionary biology, indeed any science that attempts to reconstruct prehistory, faces practical limitations on available data. These limitations create the problem of contrast failure: specific observations may fail to discriminate between rival evolutionary hypotheses. Assessing the risk of contrast failure provides a way to evaluate testing protocols in evolutionary science. Here I will argue that part of the methodological critique in the Spandrels paper involves diagnosing contrast failure problems. I then distinguish the problem of contrast failure from the more familiar (...) philosophical problem of underdetermination, and demonstrate how contrast failure arises in scientific practice with an investigation into Lewontin and White’s estimation of an adaptive landscape. (shrink)

We discuss the scientific task of historical reconstruction and the problem of epistemic access. We argue that strong epistemic support for historical claims consists in the consilience of multiple independent lines of evidence, and analyze the impact hypothesis for the End-Cretaceous mass extinction to illustrate the accrual of epistemic support. Although there are elements of the impact hypothesis that enjoy strong epistemic support, the general conditions for this are strict, and help to clarify the difficulties associated with reconstructing the deep (...) past. (shrink)

Path-dependence offers a promising way of understanding the role historicity plays in explanation, namely, how the past states of a process can matter in the explanation of a given outcome. The two main existing accounts of path-dependence have sought to present it either in terms of dynamic landscapes or branching trees. However, the notions of landscape and tree both have serious limitations and have been criticized. The framework of causal networks is both more fundamental and more general that that of (...) landscapes and trees. Within this framework, I propose that historicity in networks should be understood as symmetry breaking. History matters when an asymmetric bias towards an outcome emerges in a causal network. This permits a quantitative measure for how path-dependence can occur in degrees, and offers suggestive insights into how historicity is intertwined both with causal structure and complexity. (shrink)

This essay examines the claim “path dependence entails irreversibility” from the point of view of evolutionary biology. I argue that evolutionary irreversibility possesses many faces, sometimes conflicting with path dependence. I propose an account of path dependence that does not rely on irreversibility and explains why it more naturally coexists with the notion of (contingent) irreversibility developed by the Belgian paleontologist Louis Dollo. However, I argue that we should not conceive of this relationship as necessary.

Walter Bock was committed to developing a framework for evolutionary biology. Bock repeatedly discussed how evolutionary explanations should be considered within the realm of Hempel’s deductive-nomological model of scientific explanations. Explanation in evolution would then consist of functional and evolutionary explanations, and within the latter, an explanation can be of nomological-deductive and historical narrative explanations. Thus, a complete evolutionary explanation should include, first, a deductive functional analysis, and then proceed through nomological and historical evolutionary explanations. However, I will argue that (...) his views on the deductive proprieties of functional analysis and the deductive-nomological parts of evolution fail because of the nature of evolution, which contains a historical element that the logic of deduction and Hempel’s converting law model do not compass. Conversely, Bock’s historical approach gives a critical consideration of the historical narrative element of evolutionary explanation, which is fundamental to the methodology of the historical nature of evolutionary theory. Herein, I will expand and discuss a modern view of evolutionary explanations of traits that includes the currentacknowledgement of the differences between experimental and the historical sciences, including the token and type event dichotomy, that mutually illuminate each other in order to give us a well confirmed and coherent hypothesis for evolutionary explanations. Within this framework, I will argue that the duality of evolutionary explanations is related to two components of character evolution: origin, with its evolutionary pathways along with the history, and maintenance, the function (mainly a current function) for the character being selected. (shrink)

Biologists in the last 50 years have increasingly emphasized the role of historical contingency in explaining the distribution and dynamics of biological systems. However, recent work in philosophy of biology has shown that historical contingency carries various interpretations and that we are still lacking a general understanding of historicity, i.e., a framework from which to interpret why and to what extent history matters in biological processes. Building from examples and analyses of the long-term experimental evolution (LTEE) project, this paper argues (...) that historicity possess three essential conditions: (1) multiple possible pasts, (2) multiple possible outcomes at a given instant, and (3) a relationship of causal dependence between these two sets. These criteria can be further specified in two general forms of historicity: dependence on initial conditions and path dependence. More attention is devoted to developing a rigorous account of the latter, which captures the type of historicity displayed by stochastic processes. This paper also highlights that it is often more productive to adopt an instant-relative approach and think in terms of degree of historicity instead of trying to maintain a rigid and absolute dichotomy between historical and ahistorical (completely convergent) processes. (shrink)

This paper analyzes the relevance and interconnection of two forms of historicity in ecological restoration, namely historical fidelity and path dependence. Historical fidelity is the practice of attempting to restore an ecological system to some sort of idealized past condition. Path dependence occurs when a system can evolve in alternative local equilibria, and that the order and timing of the events that follow from the initial state influence which equilibrium is reached. Using theoretical examples and case studies, the following analysis (...) shows that path dependence can seriously impinge the feasibility of historical fidelity, thus reinforcing the idea that restoration ecology needs to move away from a rigid reliance on historical fidelity and is sometimes justified to endorse an interventionist agenda. Yet, contra recent criticisms, I argue that ecological restoration, directed by prudential and evidence-based reasoning, should maintain historical fidelity as one of its guiding objectives. (shrink)

Historical scientists frequently face incomplete data, and lack direct experimental access to their targets. This has led some philosophers and scientists to be pessimistic about the epistemic potential of the historical sciences. And yet, historical science often produces plausible, sophisticated hypotheses. I explain this capacity to generate knowledge in the face of apparent evidential scarcity by examining recent work on Thylacoleo carnifex, the ‘marsupial lion’. Here, we see two important methodological features. First, historical scientists are methodological omnivores, that is, they (...) construct purpose-built epistemic tools tailored to generate evidence about highly specific targets. This allows them to produce multiple streams of independent evidence and thus maximize their epistemic reach. Second, investigative scaffolding: research proceeds in a piece-meal fashion, information only gaining evidential relevance once certain hypotheses are well supported. I illustrate scaffolding in a discussion of the nature of functional ascription in paleobiology. Frequently, different senses of ‘function’ are not discriminated during paleobiological investigation—something which can mar adaptationist investigations of extant organisms. However, I argue that, due to scaffolding, conflating senses of ‘function’ can be the right thing to do. Coarse-grained functional hypotheses are required before it is clear what evidence could discriminate between more fine-grained ones. I draw on omnivory and scaffolding to argue that pessimists make a bad empirical bet. It is a bad idea to bet against the epistemic fortunes of such opportunistic and resourceful scientists, especially when we have reason to think we will systematically underestimate the amount of evidence ultimately available to them. (shrink)

In 1923, Sir Arthur Keith proposed that many common back problems are due to the stresses caused by our evolutionarily novel form of locomotion, bipedalism. In this article, we introduce an updated version of Keith’s hypothesis with a focus on acquired spinal conditions. We begin by outlining the main ways in which the human spine differs from those of our closest living relatives, the great apes. We then review evidence suggesting there is a link between spinal and vertebral shape on (...) the one hand and acquired spinal conditions on the other. Next, we discuss recent studies that not only indicate that two common acquired spinal conditions—intervertebral disc herniation and spondylolysis—are associated with vertebral shape, but also suggest that the pathology-prone vertebral shapes can be understood in terms of the shift from quadrupedalism to bipedalism in the course of human evolution. Subsequently, we place the aforementioned findings under an umbrella hypothesis, which we call the “Evolutionary Shape Hypothesis.” This hypothesis contends that individuals differ in their propensity to develop different acquired spinal conditions because of differences in vertebral shape that relate to the evolutionary history of our species. We end the article with some possible directions for future research. (shrink)

This PhD dissertation examines the conceptual and theoretical foundations of the most general and most widely used framework for understanding social evolution, W. D. Hamilton's theory of kin selection. While the core idea is intuitive enough (when organisms share genes, they sometimes have an evolutionary incentive to help one another), its apparent simplicity masks a host of conceptual subtleties, and the theory has proved a perennial source of controversy in evolutionary biology. To move towards a resolution of these controversies, we (...) need a careful and rigorous analysis of the philosophical foundations of the theory. My aim in this work is to provide such an analysis. I begin with an examination of the concepts behavioural ecologists employ to describe and classify types of social behaviour. I stress the need to distinguish concepts that are often conflated: for example, we need to distinguish simple cooperation from collaboration in collective tasks, behaviours from strategies, and control from manipulation and coercion. I proceed from here to the formal representation of kin selection via George R. Price’s covariance selection mathematics. I address a number of interpretative issues the Price formalism raises, including the vexed question of whether kin selection theory is ‘formally equivalent’ to multi-level selection theory. In the second half of the dissertation, I assess the uses and limits of Hamilton’s rule for the evolution of social behaviour; I provide a precise statement of the conditions under which the rival neighbour-modulated fitness and inclusive fitness approaches in contemporary kin selection theory are equivalent (and describe cases in which they are not); and I criticize recent formal attempts to establish the controversial claim that kin selection leads to organisms behaving as if maximizing their inclusive fitness. (shrink)

A number of areas of biology raise questions about what is of value in the natural environment and how we ought to behave towards it: conservation biology, environmental science, and ecology, to name a few. Based on my experience teaching students from these and similar majors, I argue that the field of environmental ethics has much to teach these students. They come to me with pent-up questions and a feeling that more is needed to fully engage in their subjects, and (...) I believe some exposure to environmental ethics can help focus their interests and goals. I identify three primary areas in which environmental ethics can con- tribute to their education. The first is an examination of who (or what) should be considered to be part of our moral community (i.e., the community to whom we owe direct duties). Is it humans only? Or does it include all sentient life? Or all life? Or ecosystems considered holistically? Often, readings implicitly assume one or more of these answers; the goal is to make the student more sensitive to these implicit claims and to get them to think about the different reasons that support them. The second area, related to the first, is the application of the different answers concerning the extent of the ethical community to real environmental issues and problems. Students need to be aware of how the different answers concerning the moral community can imply conflicting answers for how we should act in certain cases and to think about ways to move toward conflict resolution. The third area in which environmental ethics can contribute is a more conceptual one, focusing on central concepts such as biodiversity, sustainability, species, and ecosystems. Exploring and evaluating various meanings of these terms will make students more reflective and thoughtful citizens and biologists, sensitive to the implications that different conceptual choices make. (shrink)