Assessing the fitness landscape revolution

According to Pigliucci and Kaplan, there is a revolution underway in how we understand fitness landscapes. Recent models suggest that a perennial problem in these landscapes—how to get from one peak across a fitness valley to another peak—is, in fact, non-existent. In this paper I assess the structure and the extent of Pigliucci and Kaplan’s proposed revolution and argue for two points. First, I provide an alternative interpretation of what underwrites this revolution, motivated by some recent work on model-based science. Second, I show that the implications of this revolution need to carefully assessed depending on question being asked, for peak-shifting is not central to all evolutionary questions that fitness landscapes have been used to explore.

Spandrels and a pervasive problem of evidence

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 (Evolution 14:116–129, 1960) estimation of an adaptive landscape.

The end of the adaptive landscape metaphor?

The concepts of adaptive/fitness landscapes and adaptive peaks are a central part of much of contemporary evolutionary biology; the concepts are introduced in introductory texts, developed in more detail in graduate-level treatments, and are used extensively in papers published in the major journals in the field. The appeal of visualizing the process of evolution in terms of the movement of populations on such landscapes is very strong; as one becomes familiar with the metaphor, one often develops the feeling that it is possible to gain deep insights into evolution by thinking about the movement of populations on landscapes consisting of adaptive valleys and peaks. But, since Wright first introduced the metaphor in 1932, the metaphor has been the subject of persistent confusion, from equivocation over just what the features of the landscape are meant to represent to how we ought to expect the landscapes to look. Recent advances—conceptual, empirical, and computational—have pointed towards the inadequacy and indeed incoherence of the landscapes as usually pictured. I argue that attempts to reform the metaphor are misguided; it is time to give up the pictorial metaphor of the landscape entirely and rely instead on the results of formal modeling, however difficult such results are to understand in ‘intuitive’ terms.

The two faces of revenge: moral responsibility and the culture of honor

Retributive emotions and behavior are thought to be adaptive for their role in improving social coordination. However, since retaliation is generally not in the short-term interests of the individual, rational self-interest erodes the motivational link between retributive emotions and the accompanying adaptive behavior. I argue that two different sets of norms have emerged to reinforce this link: (1) norms about honor and (2) norms about moral responsibility and desert. I observe that the primary difference between these types of retribution motivators lies in where the normative focus is placed after an offense. In the first form of retribution, the normative focus is on the offended party. In the second, it is on the offender. Next, I show how each class of norms is well tailored to the particular features of the environment in which these forms of retributive behavior emerge. Finally, I consider some philosophical implications of these observations. I suggest that my account, if correct, would pose tough challenges for contemporary philosophical theories of moral responsibility and punishment.

Foresight in cultural evolution

Critics of Darwinian cultural evolution frequently assert that whereas biological evolution is blind and undirected, cultural change is directed or guided by people who possess foresight, thereby invalidating any Darwinian analysis of culture. Here I show this argument to be erroneous and unsupported in several respects. First, critics commonly conflate human foresight with supernatural clairvoyance, resulting in the premature rejection of Darwinian cultural evolution on false logical grounds. Second, the presence of foresight is perfectly consistent with Darwinian evolution, and is found in biology, in the form of open, teleonomic processes such as genetically-biased behavioural learning. Finally, empirical evidence from the social sciences suggests that cultural change appears far less guided and directed, and human foresight far less accurate, than is commonly assumed.

The rise and fall of the adaptive landscape?

The discussion of the adaptive landscape in the philosophical literature appears to be divided along the following lines. On the one hand, some claim that the adaptive landscape is either “uninterpretable” or incoherent. On the other hand, some argue that the adaptive landscape has been an important heuristic, or tool in the service of explaining, as well as proposing and testing hypotheses about evolutionary change. This paper attempts to reconcile these two views.

Individual optimization efforts and population dynamics: a mathematical model for the evolution of resource allocation strategies, with applications to reproductive and mating systems

We develop a formal framework for the optimal allocation of limited resources that includes and clarifies the interplay between individual optimization and the resulting effects at the population level. As an example, in regard to the evolution of sexual recombination, the paradox of the twofold cost of sex is avoided by distinguishing between the evolution of recombination and the subsequent emergence and stability of different mating types as a result of individual optimization within a population that benefits from recombination.

Adaptationism and the adaptive landscape

Debates over adaptationism can be clarified and partially resolved by careful consideration of the ‘grain’ at which evolutionary processes are described. The framework of ‘adaptive landscapes’ can be used to illustrate and facilitate this investigation. We argue that natural selection may have special status at an intermediate grain of analysis of evolutionary processes. The cases of sickle-cell disease and genomic imprinting are used as case studies.

Fitness “kinematics”: biological function,altruism, and organism–environment development

It’s recently been argued that biological fitness can’t change over the course of an organism’s life as a result of organisms’ behaviors. However, some characterizations of biological function and biological altruism tacitly or explicitly assume that an effect of a trait can change an organism’s fitness. In the first part of the paper, I explain that the core idea of changing fitness can be understood in terms of conditional probabilities defined over sequences of events in an organism’s life. The result is a notion of “conditional fitness” which is static but which captures intuitions about apparent behavioral effects on fitness. The second part of the paper investigates the possibility of providing a systematic foundation for conditional fitness in terms of spaces of sequences of states of an organism and its environment. I argue that the resulting “organism–environment history conception” helps unify diverse biological perspectives, and may provide part of a metaphysics of natural selection.

Sexual selection and mate choice in evolutionary psychology

The importance of mate choice and sexual selection has been emphasized by the majority of evolutionary psychologists. This paper assesses three cases of work on mate choice and sexual selection in evolutionary psychology: David Buss on cross-cultural human mate preferences, Randy Thornhill and Steve Gangestad on the link between mate preferences and fluctuating asymmetry, and Geoffrey Miller on the role of Fisher’s runaway process in human evolution. A mixture of conceptual and empirical problems in each case highlights the general weakness of work in evolutionary psychology on these issues.

Forty Years of ‘The Strategy’: Levins on Model Building and Idealization

This paper is an interpretation and defense of Richard Levins’ “The Strategy of Model Building in Population Biology,” which has been extremely influential among biologists since its publication 40 years ago. In this article, Levins confronted some of the deepest philosophical issues surrounding modeling and theory construction. By way of interpretation, I discuss each of Levins’ major philosophical themes: the problem of complexity, the brute-force approach, the existence and consequence of tradeoffs, and robustness analysis. I argue that Levins’ article is concerned, at its core, with justifying the use of multiple, idealized models in population biology.

How ubiquitous is adaptation? A critique of the epiphenomenist program

It is important to distinguish adaptation per se (adaptedness, or being adapted) from the more specific concept of adaptation for some function. Commonly used criteria for adaptation in either sense have limited applicability. There are, however, a number of widely applicable criteria for adaptation per se, such as several kinds of cost, low variation, the maintenance of integration, and the fitness distribution of mutations. Application of these criteria leads to the conclusion that adaptation is overwhelmingly prevalent for features of organisms. Neither the presence nor the absence of adaptation has a privileged status in inference. Effectively neutral evolution can occur on adaptive buttes while maintaining the same degree of adaptation, but it is likely to be relatively minor.

Language processing with dynamic fields

We construct a mapping from complex recursive linguistic data structures to spherical wave functions using Smolensky’s filler/role bindings and tensor product representations. Syntactic language processing is then described by the transient evolution of these spherical patterns whose amplitudes are governed by nonlinear order parameter equations. Implications of the model in terms of brain wave dynamics are indicated. Electronic supplementary material  The online version of this article (doi: ) contains supplementary material, which is available to authorized users.

Intelligent design and mathematical statistics: a troubled alliance

The explanatory filter is a proposed method to detect design in nature with the aim of refuting Darwinian evolution. The explanatory filter borrows its logical structure from the theory of statistical hypothesis testing but we argue that, when viewed within this context, the filter runs into serious trouble in any interesting biological application. Although the explanatory filter has been extensively criticized from many angles, we present the first rigorous criticism based on the theory of mathematical statistics.

The strategy of “The strategy of model building in population biology”

In this essay, I argue for four related claims. First, Richard Levins’ classic “The Strategy of Model Building in Population Biology” was a statement and defense of theoretical population biology growing out of collaborations between Robert MacArthur, Richard Lewontin, E. O. Wilson, and others. Second, I argue that the essay served as a response to the rise of systems ecology especially as pioneered by Kenneth Watt. Third, the arguments offered by Levins against systems ecology and in favor of his own methodological program are best construed as “pragmatic”. Fourth, I consider limitations of Levins’ arguments given contemporary population biology.

Hennigian Phylogenetic Systematics and the ‘Groundplan’ vs. ‘Post-Groundplan’ Approaches: A Reply to Kukalová-Peck

The recent contribution by Jarmila Kukalová-Peck on Hennigian phylogenetics and hexapod limb evolution is critically evaluated.

Productivity,relevance and natural selection

Recent papers by a number of philosophers have been concerned with the question of whether natural selection is a causal process, and if it is, whether the causes of selection are properties of individuals or properties of populations. I shall argue that much confusion in this debate arises because of a failure to distinguish between causal productivity and causal relevance. Causal productivity is a relation that holds between events connected via continuous causal processes, while causal relevance is a relationship that can hold between a variety of different kinds of facts and the events that counterfactually depend upon them. I shall argue that the productive character of natural selection derives from the aggregation of individual processes in which organisms live, reproduce and die. At the same time, a causal explanation of the distribution of traits will necessarily appeal both to causally relevant properties of individuals and to causally relevant properties that exist only at the level of the population.

The adaptive landscape of science

In 1988, David Hull presented an evolutionary account of science. This was a direct analogy to evolutionary accounts of biological adaptation, and part of a generalized view of Darwinian selection accounts that he based upon the Universal Darwinism of Richard Dawkins. Criticisms of this view were made by, among others, Kim Sterelny, which led to it gaining only limited acceptance. Some of these criticisms are, I will argue, no longer valid in the light of developments in the formal modeling of evolution, in particular that of Sergey Gavrilets’ work on adaptive landscapes. If we can usefully recast the Hullian view of science as being driven by selection in terms of Gavrilets’ and Kaufmann’s view of there being “giant components” of high-fitness networks through any realistic adaptive landscape, we may now find it useful to ask what the adaptive pressures on science are, and to extend the metaphor into a full analogy. This is in effect to reconcile the Fisherianism of the Dawkins–Hull approach to selection and replicators, with a Wrightean drift account of social constructionist views of science, preserving, it is to be hoped, the valuable aspects of both.

Creationism in Germany and its Possible Cause

In a widely cited essay published in the journal Science (Aug. 2006), it was stated that in Germany a large majority of adults are convinced of evolution. Based on a new poll, I document that this conclusion is questionable. In Sept. 2005, 60.9% of the German population accepted naturalistic evolution, whereas 37.7% were creationists or adherents of intelligent design. Because among Christians of both confessions the percentage of anti-evolutionists is much larger than among atheists, I suggest that religious indoctrination of children may in part be responsible for the rejection of the concept of evolution, notably in the western part of Germany.

The Concealed Information Test as an Instrument of Applied Differential Psychophysiology: Methodological Considerations

In this interpretative paper, I consider four sets of methodological issues that may be relevant to improving the concealed information test (CIT) as an instrument of applied differential psychophysiology. The first set has to do with psychophysiological measurement in the CIT (e.g., specific sensitivity testing in lab vs. field). Secondly, I consider the relationships between the psychological process of deception and the CIT. Thirdly, I consider the problem of laboratory-to-field generalization of the CIT, a consideration that includes a discussion of whether the lab/field differences are merely quantitative or actually qualitative. Finally, I discuss theories concerning the hypothetical mechanisms underlying the CIT, and argue that while the purely cognitive, Sokolovian, orienting response (OR) account is widely accepted as the sole mechanism, there is evidence to suggest that not just motivational, but even emotional mechanisms are also relevant.