The most consistent definition of fitness makes it a static property of organisms. However, this is not how fitness is used in many evolutionary models. In those models, fitness is permitted to vary with an organism’s circumstances. According to this second conception, fitness is dynamic. There is consequently tension between these two conceptions of fitness. One recently proposed solution suggests resorting to conditional properties. We argue, however, that this solution is unsatisfactory. Using a very simple model, we show that it (...) can lead to incompatible fitness values and indecision about whether selection actually occurs. (shrink)

The propensity nature of evolutionary fitness has long been appreciated and is nowadays amply discussed. The discussion has, however, on occasion followed long standing conflations in the philosophy of probability literature between propensities, probabilities, and frequencies. In this paper, I apply a more recent conception of propensities in modelling practice to some of the key issues, regarding the mathematical representation of fitness and how it may be regarded as explanatory. The ensuing complex nexus of fitness emphasises the distinction between biological (...) propensities and the probability distributions over offspring numbers that they give rise to; and how critical it is to distinguish the possession conditions of the underlying dispositional properties from those of their probabilistic manifestations. (shrink)

In several places we have argued that ‘fitness’ is a primitive term with respect to the theory of evolution properly understood. These arguments have relied heavily on the axiomatization of the theory provided by one of us. In contrast, both John Beatty and Robert Brandon have separately argued for a “propensity“ interpretation of “fitness” ; and in Brandon and Beatty they attack our view that “fitness“ is a primitive term in evolutionary theory, concluding that a definition by way of propensities (...) is possible and preferable. Here we reply to their criticisms, and argue that, at most, the view that fitness is a statistical propensity is a terminological variant on our thesis that it is a primitive in evolutionary theory. (shrink)

The propensity account of fitness intends to solve the classical tautologicity issue by identifying fitness with a disposition, the ability to survive and reproduce. As proponents recognized early on, this account requires operational independence from actual reproductive success to avoid circularity and vacuousness charges. They suggested that operational independence is achieved by measuring fitness values through optimality models. Our goal in this article is to develop this suggestion. We show that one plausible procedure by which these independent operationalizations could be (...) thought to take place, and which is in accordance with what is said in the optimality literature, is unsound. We provide two independent lines of reasoning to show this. We then provide a sketch of a more adequate account of the role of optimality models in evolutionary contexts and draw some consequences. (shrink)

In a recent issue of Biology and Philosophy, Kenneth Waters argues that the principle of survival of the fittest should be eliminated from the theory of natural selection, because it is an untestable law of probability, and as such, has no place in evolutionary theory. His argument is impressive, but it does not do justice to the practice of biology. The principle of survival of the fittest should not be eliminated from the theory of natural selection because it is important (...) to biological practice: it plays an essential role in explanation and discovery, and in unffying the theory of natural selection. (shrink)

Susan Mills and John Beatty's propensity interpretation of fitness encountered very different philosophical criticisms by Alexander Rosenberg and Kenneth Waters. These criticisms and the rejoinders to them are both predictable and important. They are predictable as raisingkinds of issues typically associated with disposition concepts (this is established through a systematic review of the problems generated by Carnap's dispositional interpretation of all scientific terms). They are important as referring the resolution of these issues to the development of evolutionary biology. This historical (...) approach to the propensity interpretation of fitness draws attention to the precarious relation between philosophical clarification of scientific concepts and any given state of the empirical arts. (shrink)

This chapter contains sections titled: Highlights of Past Literature Current Work Future Work.

In this paper I discuss recent debates concerning etiological theories of functions. I defend an etiological theory against two criticisms, namely the ability to account for malfunction, and the problem of structural doubles. I then consider the arguments provided by Bigelow and Pargetter (1987) for a more forward looking account of functions as propensities or dispositions. I argue that their approach fails to address the explanatory problematic for which etiological theories were developed.

The chief argument for scientific realism is the no-miracles argument, according to which the approximate truth of our current scientific theories can be inferred from their success through time. To date, anti-realist responses to the argument have been unconvincing, largely because of their anti-realistic presuppositions. In this paper, it is shown that realists cannot pre-emptively dismiss the problem of the underdetermination of theory by evidence, and that the no-miracles argument fails because it does nothing to dispel the threat posed by (...) underdetermination, although it may be effective against other anti-realistic arguments. (shrink)

Darwin’s claim about natural selection is reconstructed as an empirical claim about a causal connection leading from the match of the physiology of an individual and its environment to leaving surviving progeny. Variations in this match, Darwin claims, cause differences in the survival of the progeny. Modern concepts of fitness focus the survival side of this chain. Therefore, the assumption that evolutionary theory wants to explain reproductive success in terms of a modern concept of fitness has given rise to the (...) so-called tautology problem. It is shown that the tautology problem reappears in the treatment of fitness proxies in today’s experimental evolutionary biology when these proxies are considered to indicate fitness only. Taking Darwin’s empirical claim seriously suggests, by contrast, that fitness proxies are first of all measures of the match between organism and environment, which I call the organism’s ‘fittedness’. At the same time, they are indeed related to reproductive success. Thus looking in both directions, at fitness and at fittedness, they are janiform. Acknowledging this situation not only allows for rejection of the tautology objection, but also for integration of Darwin’s argument into current evolutionary biology. It is suggested that this helps reframe and alleviate the dispute between the Modern Synthesis and the Extended Evolutionary Synthesis. (shrink)

The assumption that natural selection alone is sufficient to explain not only which traits get fixed in a population/species, but also how they develop, has been questioned since Darwin’s times, and increasingly in the last decades. Alternative theories, linked to genetic and phenotypic processes, or to the theory of complex systems, have been proposed to explain the rise of the phenotypic variety upon which natural selection acts. In this article, we illustrate the current state of the issue and we propose (...) a logical space based on phenotypic robustness that allows a classification of evolutionary phenomena and can provide a framework for unifying all these accounts. (shrink)

Fitness is a central but notoriously vexing concept in evolutionary biology. The propensity interpretation of fitness is often regarded as the least problematic account for fitness. It ties an individual's fitness to a probabilistic capacity to produce offspring. Fitness has a clear causal role in evolutionary dynamics under this account. Nevertheless, the propensity interpretation faces its share of problems. We discuss three of these. We first show that a single scalar value is an incomplete summary of a propensity. Second, we (...) argue that the widespread method of “abstracting away” environmental idiosyncrasies by averaging over reproductive output in different environments is not a valid approach when environmental changes are irreversible. Third, we point out that expanding the range of applicability for fitness measures by averaging over more environments or longer time scales (so as to ensure environmental reversibility) reduces one's ability to distinguish selectively relevant differences among individuals because of mutation and eco‐evolutionary feedbacks. This series of problems leads us to conclude that a general value of fitness that is both explanatory and predictive cannot be attained. We advocate for the use of propensity‐compatible methods, such as adaptive dynamics, which can accommodate these difficulties. (shrink)

Sustained controversy over a philosophical issue is often times symptomatic of differing commitments at a more fundamental philosophical level. I will argue that two current debates over the foundations of the theory of natural selection are cases in point. Alexander Rosenberg, at times together with Mary Williams, challenges what is becoming received orthodoxy about the foundations of this theory. He argues that the currently popular propensity interpretation of fitness does not legitimize explanations in terms of natural selection, and that furthermore, (...) the biological fields which typically study the ways in which selection pressures originate in the organism/environment complex are not in fact part of the theory of natural selection at all. His foils in the first case are Robert Brandon, John Beatty and Susan Mills;2 on the second issue he is engaged in debate with Elliott Sober.3 We will see that, although the controversies are ostensibly about the interpretation of fitness, the issues, at bottom, involve the nature of scientific explanation. (shrink)

Supervenience is a relationship which has been used recently to explain the physical determination of biological phenomena despite resistance to reduction. Supervenience, however, is plagued by ambiguities which weaken its explanatory value and obscure some interesting aspects of reduction in biology. Although I suspect that similar considerations affect the use of supervenience in ethics and the philosophy of mind, I don’t intend anything I have to say here to apply outside of the physical and biological cases I consider.The main point (...) of this paper is that there is a property of biological systems which makes it both misleading and inappropriate to reduce central biological phenomena to the properties of underlying components. Despite this, reductive explanation has been a major source of innovation in biological theory. The apparent tension can be resolved if underlying properties are explanatorily relevant to the higher level phenomena even though the latter are not strictly reducible to the former. Supervenience, I will argue, is not robust enough to deny reduction while supporting explanatory relevance. (shrink)

El proyecto del ‘Darwinismo Formal’ de Alan Grafen propone una formulación matemática de la teoría de Darwin que pretende demostrar que la selección natural moldea los rasgos fenotípicos a través de la maximización de la eficacia. El proyecto de Grafen reposa sobre tres premisas: la selección natural es la única fuerza que moldea los fenotipos; la eficacia es la única medida de le evolución; y el diseño biológico surge como resultado de un proceso de optimización selectiva. En este trabajo argumentamos (...) que estas tres premisas limitan la aplicabilidad del modelo a los hechos evolutivos más simples. Esto se debe a que Grafen implícitamente presupone un concepto de eficacia que es a la vez causa y efecto de las novedades fenotípicas; lo que, por un lado, vacía la teoría de Darwin de poder explicativo y, por el otro, lleva a ignorar la potencial contribución de fuerzas evolutivas no selectivas en la configuración de la complejidad biológica. Para superar estos escollos del proyecto de del Darwinismo Formal - que son además comunes a todos los modelos formales adaptacionistas - proponemos sustituir el concepto causal de eficacia por el de robustez, definiendo así el diseño biológico como forma y función en lugar de como maximización de la eficacia. (shrink)

The assumption that natural selection alone is sufficient to explain not only which traits get fixed in a population/species, but also how they develop, has been questioned since Darwin’s times, and increasingly in the last decades. Alternative theories, linked to genetic and phenotypic processes, or to the theory of complex systems, have been proposed to explain the rise of the phenotypic variety upon which natural selection acts. In this article, we illustrate the current state of the issue and we propose (...) a logical space based on phenotypic robustness that allows a classification of evolutionary phenomena and can provide a framework for unifying all these accounts. (shrink)

Darwin's theory of evolution by natural selection provided the first, and only, causal-mechanistic account of the existence of adaptations in nature. As such, it provided the first, and only, scientific alternative to the “argument from design”. That alone would account for its philosophical significance. But the theory also raises other philosophical questions not encountered in the study of the theories of physics. Unfortunately the concept of natural selection is intimately intertwined with the other basic concepts of evolutionary theory—such as the (...) concepts of fitness and adaptation —that are themselves philosophically controversial. Fortunately we can make considerable headway in getting clear on natural selection without solving all of those outstanding problems. (shrink)

Fitness plays many roles throughout evolutionary theory, from a measure of populations in the wild to a central element in abstract theoretical presentations of natural selection. It has thus been the subject of an extensive philosophical literature, which has primarily centered on the way to understand the relationship between fitness values and reproductive outcomes. If fitness is a probabilistic or statistical quantity, how is it to be defined in general theoretical contexts? How can it be measured? Can a single conceptual (...) model for fitness be offered that applies in all biological cases, or must fitness measures be case-specific? Philosophers have explored these questions over the last several decades, largely in the context of an influential definition of fitness proposed in the late 1970s: the propensity interpretation. This interpretation as first described undeniably suffers from significant difficulties, and debate regarding the tenability of amendments and alternatives to it remains unsettled. (shrink)

Evolution in its contemporary meaning in biology typically refers to the changes in the proportions of biological types in a population over time (see the entry on the concept of evolution to 1872 for earlier meanings). As evolution is too large of a topic to address thoroughly in one entry, the primary goal of this entry is to serve as a broad overview of contemporary issues in evolution with links to other entries where more in-depth discussion can be found. The (...) entry begins with a brief survey of definitions of evolution, followed by a discussion of the different modes of evolution and related philosophical issues, and ends with a summary of other topics in the philosophy of evolution focusing particularly on topics covered in this encyclopedia. (shrink)

Darwin’s claim that Natural Selection, through optimization of fitness, explains complex biological design has not yet been properly formalized. Alan Grafen’s Formal Darwinism Project aims at providing such a formalization and at demonstrating that fitness maximization is coherent with results from Population Genetics, usually interpreted as denying it. We suggest that Grafen’s proposal suffers from some limitations linked to its concept of design as optimized fitness. In order to overcome these limitations, we propose a classification of evolutionary facts based on (...) a bi-dimensional complexity space, which adds robustness to fitness as measure of the quality of a design. In this space, each point represents an organismic architecture, and movement between two points would be an evolutionary fact. Natural Selection explains movement along the fitness axis, while non-selective mechanisms explain movement on the robustness axis. Moreover, we propose a thermodynamic metaphor to draw parallelisms between notions of fitness and entropy, robustness and energy, and movement in this space and reversible and irreversible cycles. We argue that this metaphor illustrates different evolutionary processes usually overlooked by proposals of formalization of Darwinian theory, such Grafen’s Formal Darwinism Project. (shrink)