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- Ulrich Krohs (2006). The Changeful Fate of a Groundbreaking Insight: The Darwinian Fitness Principle Caught in Different Webs of Belief. Yearbook for European Culture of Science 2:107-124.Darwin’s explanation of biological speciation in terms of variation and natural selection has revolutionised biological thought. However, while his principle of natural selection, the fitness principle, has shaped biology until the present, its interpretation changed more than once during the almost 150 years of its history. The most striking change of the status of the principle is that, in the middle of the 20th century, it transmutated from an often disputed, groundbreaking insight into a tautology. Moreover, not only the interpretation of the fitness principle, but the whole body of biological knowledge was subjected to significant modifications. In this paper, I relate modifications of the fitness principle to those of the respective body of biological knowledge. This body of knowledge is conceived as a Quinean web of belief. After an exposition of Darwin’s conception of the principle, which equated fitness with adaptedness to the environment, several of its changes are analysed with respect to different webs of biological knowledge. It is concluded that the different interpretations and the reshaping of the fitness principle are rational responses to the modified systems of background knowledge, which saved the coherence of the web of biological knowledge in each single case.
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We critically examine Denis Walsh’s latest attack on the causalist view of fitness. Relying on Judea Pearl’s Sure-Thing Principle and geneticist John Gillespie’s model for fitness, Walsh has argued that the causal interpretation of fitness results in a reductio. We show that his conclusion only follows from misuse of the models, that is, (1) the disregard of the real biological bearing of the population-size parameter in Gillespie’s model and (2) the confusion of the distinction between ordinary probability and Pearl’s causal probability. Properly understood, the models used by Walsh do not threaten the causalist view of fitness.
The central point of this essay is to demonstrate the incommensurability of ‘Darwinian fitness’ with the numeric values associated with reproductive rates used in population genetics. While sometimes both are called ‘fitness’, they are distinct concepts coming from distinct explanatory schemes. Further, we try to outline a possible answer to the following question: from the natural properties of organisms and a knowledge of their environment, can we construct an algorithm for a particular kind of organismic life-history pattern that itself will allow us to predict whether a type in the population will increase or decrease relative to other types? Introduction Darwinian fitness Reproductive fitness and genetical models of evolution The models of reproductive fitness 4.1 The Standard Viability Model 4.2 Frequency-dependent selection 4.3 Fertility models 4.4 Overlapping generations Fitness as outcome 5.1 Fitness as actual increase in type 5.2 Fitness as expected increase in type 5.2.1 Expected increase within a generation 5.2.2 Expected increase between generations 5.2.3 Postponed reproductive fitness effects The book-keeping problem Conclusion.
The concept of fitness began its career in biology long before evolutionary theory was mathematized. Fitness was used to describe an organism’s vigor, or the degree to which organisms “fit” into their environments. An organism’s success in avoiding predators and in building a nest obviously contribute to its fitness and to the fitness of its offspring, but the peacock’s gaudy tail seemed to be in an entirely different line of work. Fitness, as a term in ordinary language (as in “physical fitness”) and in its original biological meaning, applied to the survival of an organism and its offspring, not to sheer reproductive output (Paul ////; Cronin 1991). Darwin’s separation of natural from sexual selection may sound odd from a modern perspective, but it made sense from this earlier point of view.
The diversity, complexity and adaptation of the biological realm is evident. Until Darwin, the best explanation for these three features of the biological was the conclusion of the “argument from design.” Darwin's theory of natural selection provides an explanation of all three of these features of the biological realm without adverting to some mysterious designing entity. But this explanation's success turns on the meaning of its central explanatory concept, ‘fitness’. Moreover, since Darwinian theory provides the resources for a purely causal account of teleology, wherever it is manifested, its reliance on the concept of ‘fitness’ makes it imperative that conceptual problems threatening the explanatory legitimacy of this notion be solved.
The diversity, complexity and adaptation of the biological realm is evident. Until Darwin, the best explanation for these three features of the biological was the conclusion of the “argument from design.” Darwin's theory of natural selection provides an explanation of all three of these features of the biological realm without adverting to some mysterious designing entity. But this explanation's success turns on the meaning of its central explanatory concept, ‘fitness’. Moreover, since Darwinian theory provides the resources for a purely causal account of teleology, wherever it is manifested, its reliance on the concept of ‘fitness’ makes it imperative that conceptual problems threatening the explanatory legitimacy of this notion be solved.
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.
Philosophers of biology have been absorbed by the problem of defining evolutionary fitness since Darwin made it central to biological explanation. The apparent problem is obvious. Define fitness as some biologists implicitly do, in terms of actual survival and reproduction, and the principle of natural selection turns into an empty tautology: those organisms which survive and reproduce in larger numbers, survive and reproduce in larger numbers. Accordingly, many writers have sought to provide a definition for ‘fitness’ which avoid this outcome. In particular the definition of fitness as a probabilistic propensity has been widely favored.1 Others, recognizing that no definition both correct and complete can actually be provided, have accepted the consequence that the leading principle of the theory is a definitional truth and attempted to mitigate the impact of this outcome for the empirical character of the theory.2 Still others have argued that ‘fitness’ is properly viewed as a term undefined in the theory of natural selection (on the model of mass—a term undefined in Newtonian mechanics).3 But few have contemplated the solution to this problem proposed by Mohan Matthen and André Ariew (hereafter, MA), in..
Susan Mills and John Beatty proposed a propensity interpretation of fitness (1979) to show that Darwinian explanations are not circular, but they did not address the critics' chief complaint that the principle of the survival of the fittest is either tautological or untestable. I show that the propensity interpretation cannot rescue the principle from the critics' charges. The critics, however, incorrectly assume that there is nothing more to Darwin's theory than the survival of the fittest. While Darwinians all scoff at this assumption, they do not agree about what role, if any, this principle plays in Darwin's theory of natural selection. I argue that the principle has no place in Darwin's theory. His theory does include the idea that some organisms are fitter than others. But greater reproductive success is simply inferred from higher fitness. There is no reason to embody this inference in the form of a special principle of the survival of the fittest.
The principle of natural selection is stated. It connects fitness values (actual reproductive success) with expected fitness values. The term 'adaptedness' is used for expected fitness values. The principle of natural selection explains differential fitness in terms of relative adaptedness. It is argued that this principle is absolutely central to Darwinian evolutionary theory. The empirical content of the principle of natural selection is examined. It is argued that the principle itself has no empirical biological content, but that the presuppositions of its applicability are empirical. They form the empirical biological core of evolutionary theory.
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The theory of natural selection is a rich systematization of biological knowledge without a first principle. When formulations of a proposed principle of natural selection are examined carefully, each is seen to be exhaustively analyzable into a proposition about sources of fitness and a proposition about consequences of fitness. But whenever the fitness of an organic variety is well defined in a given biological situation, its sources are local contingencies together with the background of laws from disciplines other than the theory of natural selection; and the consequences of fitness for the long range fate of organic varieties are essentially applications of probability theory. Hence there is no role and no need for a principle of the theory of natural selection, and any generalities that may hold in that theory are derivative rather than fundamental.
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