How to read “heritability” in the recipe approach to natural selection

There are two ways evolution by natural selection (ENS) is conceptualized in the literature. One provides a ‘recipe’ for ENS incorporating three ingredients: variation, differences in fitness and heritability. The other provides formal equations of evolutionary change and partitions out selection from other causes of evolutionary changes such as transmission biases or drift. When comparing the two approaches there seems to be a tension around the concept of heritability. A recent claim has be made that the recipe approach is flawed and should be abandoned. In this paper I show that the tension is only a superficial one. If one uses a concept of heritability strictly in line with the formal equations of evolutionary change, the recipe approach keeps its validity and generality. To demonstrate that the intuitive concept of heritability is not a general one, I use one formulation of the Price equation formulated by Okasha, show that the concept of heritability in his formulation incorporates both the intuitive notion of heritability as a measure of similarity between parent and offspring characters and a measure of persistence. I advocate that for persistence to be incorporated in the concept of heritability used in recipes for ENS in the same way heredity is, show that this is readily attainable and thereby dissolve any point of tension concerning heritability between the recipe and the analytical approach to ENS

Being Precise about Precision and One-to-one Specificity

Following from my criticisms of Calcott’s analysis on the permissive/instructive distinction, I rebut his claims that 1) he clarifies my measure one-to-one specificity; 2) for all intents and purposes of his analysis his notion of precision is different from my measure of one- to-one specificity; 3) Waddington box is a better and different model than the extension of Woodward’s radio I propose

Being Precise about Precision and One-to-one Specificity

Following from my criticisms of Calcott’s analysis on the permissive/instructive distinction, I rebut his claims that 1) he clarifies my measure one-to-one specificity; 2) for all intents and purposes of his analysis his notion of precision is different from my measure of one- to-one specificity; 3) Waddington box is a better and different model than the extension of Woodward’s radio I propose

Origins and evolution of religion from a Darwinian point of view: synthesis of different theories

The religious phenomenon is a complex one in many respects. In recent years an increasing number of theories on the origin and evolution of religion have been put forward. Each one of these theories rests on a Darwinian framework but there is a lot of disagreement about which bits of the framework account best for the evolution of religion. Is religion primarily a by-product of some adaptation? Is it itself an adaptation, and if it is, does it benefi ciate individuals or groups? In this chapter, I review a number of theories that link religion to cooperation and show that these theories, contrary to what is often suggested in the literature, are not mutually exclusive. As I present each theory, I delineate an integrative framework that allows distinguishing the explanandum of each theory. Once this is done, it becomes clear that some theories provide good explanations for the origin of religion but not so good explanations for its maintenance and vice versa. Similarly some explanations are good explanations for the evolution of religious individual level traits but not so good explanations for traits hard to defi ne at the individual level. I suggest that to fully understand the religious phenomenon, integrating in a systematic way the different theories and the data is a more successful approach

Reconceptualising evolution by natural selection

This thesis examines the theoretical and philosophical underpinnings of the concept of natural selection which is pervasively invoked in biology and other ‘evolutionary’ domains. Although what constitutes the process of natural selection appears to be very intuitive (natural selection results from entities exhibiting differences in fitness in a population), this conceals a number of theoretical ambiguities and difficulties. Some of these have been pointed out numerous times; others have hardly been noticed. One aim of this work is to unpack these difficulties and ambiguities; another is to provide new solutions and clarifications to them using a range of philosophical and conceptual tools. The result is a concept of natural selection stripped down from its biological specificities. I start by revisiting the entangled debates over whether natural selection is a cause of evolutionary change as opposed to a mere statistical effect of other causes, at what level this putative cause operates and whether it can be distinguished from drift. Borrowing tools from the causal modelling literature, I argue that natural selection is best conceived as a causal process resulting from individual level differences in a population. I then move to the question of whether the process of natural selection requires perfect transmission of types. I show that this question is ambiguous and can find different answers. From there, I distinguish the process of natural selection from some of its possible products, namely, evolution by natural selection and complex adaptation. I argue that reproduction and inheritance are conceptually distinct from natural selection, and using individual-based models, I demonstrate that they can be conceived as evolutionary products of it. This ultimately leads me to generalise the concepts of heritability and fitness used in the formal equations of evolutionary change. Finally, I argue that concepts of fitness and natural selection crucially depend on the grains of description at and temporal scales over which evolutionary explanations are given. These considerations reveal that the metaphysical status of the process of natural selection is problematic and why neglecting them can lead to flawed arguments in the levels of selection debate

Dissolving the Missing Heritability Problem

Heritability estimates obtained from genome-wide association studies (GWAS) are much lower than those of traditional quantitative methods. This phenomenon has been called the “missing heritability problem.” By analyzing and comparing GWAS and traditional quantitative methods, we fi rst show that the estimates obtained from the latter involve some terms other than additive genetic variance, while the estimates from the former do not. Second, GWAS, when used to estimate heritability, do not take into account additive epigenetic factors transmitted across generations, while traditional quantitative methods do. Given these two points we show that the missing heritability problem can largely be dissolved

Reconceptualising evolution by natural selection

This thesis examines the theoretical and philosophical underpinnings of the concept of natural selection which is pervasively invoked in biology and other ‘evolutionary’ domains. Although what constitutes the process of natural selection appears to be very intuitive (natural selection results from entities exhibiting differences in fitness in a population), this conceals a number of theoretical ambiguities and difficulties. Some of these have been pointed out numerous times; others have hardly been noticed. One aim of this work is to unpack these difficulties and ambiguities; another is to provide new solutions and clarifications to them using a range of philosophical and conceptual tools. The result is a concept of natural selection stripped down from its biological specificities. I start by revisiting the entangled debates over whether natural selection is a cause of evolutionary change as opposed to a mere statistical effect of other causes, at what level this putative cause operates and whether it can be distinguished from drift. Borrowing tools from the causal modelling literature, I argue that natural selection is best conceived as a causal process resulting from individual level differences in a population. I then move to the question of whether the process of natural selection requires perfect transmission of types. I show that this question is ambiguous and can find different answers. From there, I distinguish the process of natural selection from some of its possible products, namely, evolution by natural selection and complex adaptation. I argue that reproduction and inheritance are conceptually distinct from natural selection, and using individual-based models, I demonstrate that they can be conceived as evolutionary products of it. This ultimately leads me to generalise the concepts of heritability and fitness used in the formal equations of evolutionary change. Finally, I argue that concepts of fitness and natural selection crucially depend on the grains of description at and temporal scales over which evolutionary explanations are given. These considerations reveal that the metaphysical status of the process of natural selection is problematic and why neglecting them can lead to flawed arguments in the levels of selection debate

Being Precise about Precision and One-to-one Specificity

Following from my criticisms of Calcott’s analysis on the permissive/instructive distinction, I rebut his claims that 1) he clarifies my measure one-to-one specificity; 2) for all intents and purposes of his analysis his notion of precision is different from my measure of one- to-one specificity; 3) Waddington box is a better and different model than the extension of Woodward’s radio I propose

Fidelity and the grain problem in cultural evolution

High-fidelity cultural transmission, rather than brute intelligence, is the secret of our species’ success, or so many cultural evolutionists claim. It has been selected because it ensures the spread, stability and longevity of beneficial cultural traditions, and it supports cumulative cultural change. To play these roles, however, fidelity must be a causally-efficient property of cultural transmission. This is where the grain problem comes in and challenges the explanatory potency of fidelity. Assessing the degree of fidelity of any episode or mechanism of cultural transmission always depends upon an investigator’s choice of grain of analysis at which cultural traditions are being studied. The fidelity of cultural transmission then appears to be relative to the granularity at which one approaches cultural variation, and since there is a multiplicity of grains of description by which the same tradition can be studied, there results a multiplicity of measures of fidelity for a same event or mechanism of cultural transmission. If this is correct, because fidelity is always relative to the grain of analysis dictated by the local and specific research interests of the investigator, then there seems to be no fact of the matter as to whether cultural transmission is faithful or not, independently from a researcher’s framework of analysis. The aims of this paper are to offer a conceptual clarification of the grain problem in cultural evolution, to assess its causes, to unpack its epistemological implications, and to examine its reach and consequences for a science of cultural evolution

The idea of mismatch in evolutionary medicine

Mismatch is a prominent concept in evolutionary medicine and a number of philosophers have published analyses of this concept. The word ‘mismatch’ has been used in a diversity of ways across a range of sciences, leading these authors to regard it as a vague concept in need of philosophical clarification. Here, in contrast, we concentrate on the use of mismatch in modelling and experimentation in evolutionary medicine. This reveals a rigorous theory of mismatch within which the term ‘mismatch’ is indeed used in several ways, not because it is ill-defined but because different forms of mismatch are.distinguished within the theory. Contemporary evolutionary medicine has unified the idea of ‘evolutionary mismatch’, derived from the older idea of ‘adaptive lag’ in evolution, with ideas about mismatch in development and physiology derived from the Developmental Origins of Health and Disease (DOHaD) paradigm. A number of publications in evolutionary medicine have tried to make this theoretical framework explicit. We build on these to present the theory in as simple and general a form as possible. We introduce terminology, largely drawn from the existing literature, to distinguish the different forms of mismatch. This integrative theory of mismatch captures how organisms track environments across space and time on multiple scales in order to maintain an adaptive match to the environment, and how failures of adaptive tracking lead to disease. Mismatch is a productive organising concept within this theory which helps researchers articulate how physiology, development and evolution interact with one another and with environmental change to explain health outcomes