Heritability, Maximization, and Evolutionary Explanations

2015 ◽  
Author(s):  
James A.R. Marshall
Keyword(s):  
Natural Selection ◽  
Social Evolution ◽  
Inclusive Fitness ◽  
Social Behaviors ◽  
Multilevel Selection ◽  
Evolutionary Processes ◽  
Fitness Effects ◽  
Group Fitness ◽  
Inclusive Fitness Theory ◽  
Evolutionary Explanations

This chapter examines which of the equivalent alternative partitions of fitness, including inclusive fitness and group fitness, can be interpreted as being subject to natural selection in a meaningful way. Inclusive fitness theory can deal with subtleties such as nonadditive fitness effects and conditionally expressed phenotypes. However, selection based on inclusive fitness gives equivalent predictions to other models of apparently different evolutionary processes, such as multilevel selection. The chapter considers how we can determine whether inclusive fitness really captures the essence of social evolution and whether inclusive fitness is really maximized by the action of selection, as suggested by William D. Hamilton. It also explains what heritability measures, and whether this makes sense biologically. Finally, it discusses the problem of classifying observed social behaviors in terms of their underlying evolutionary explanations.

Social Behavior and Evolutionary Thought

2015 ◽  
Author(s):  
James A.R. Marshall
Keyword(s):  
Natural Selection ◽  
Mathematical Theory ◽  
Group Selection ◽  
Inclusive Fitness ◽  
Price Equation ◽  
Inclusive Fitness Theory ◽  
Evolutionary Thought ◽  
Multilevel Selection Theory ◽  
The Individual ◽  
Evolutionary Explanations

This book demonstrates the generality of inclusive fitness theory, with particular emphasis on its fundamental evolutionary logic. It presents the basic mathematical theory of natural selection and shows how inclusive fitness theory deals with more complicated social scenarios. Topics include the Price equation, Hamilton's rule, nonadditive interactions, conditional behaviors, heritability, and maximization of inclusive fitness. This chapter provides a brief historical introduction to the problem of apparent design in biology, evolutionary explanations of this, and in particular, evolutionary explanations of individual behaviors that appear designed to benefit not the individual themselves, but other members of their species. It examines how social behaviors can be shaped by natural selection and discusses the problem of providing an evolutionary explanation of self-sacrifice by individuals, altruism in group selection, and multilevel selection theory.

Kin Selection

Ecology ◽  
2012 ◽  
Author(s):  
Andrew F. G. Bourke
Keyword(s):  
Natural Selection ◽  
Social Behavior ◽  
Kin Selection ◽  
Inclusive Fitness ◽  
Social Behaviors ◽  
Selection Theory ◽  
Direct Fitness ◽  
Inclusive Fitness Theory ◽  
Gene Copies ◽  
Genetical Theory

According to Hamilton’s kin selection theory (also known as “inclusive fitness” theory), kin selection is the process by which social evolution occurs in nature. The theory extends the genetical theory of natural selection to social behaviors and finds that their evolution is affected by the likelihood that individuals share genes (relatedness). In biology, a social behavior occurs when one individual (the actor) behaves so as to affect the direct fitness (number of offspring) of itself and another individual (the recipient). For example, altruism occurs when the actor’s behavior decreases the actor’s direct fitness and increases the recipient’s direct fitness. Conversely, selfishness occurs when the actor’s behavior increases the actor’s direct fitness and decreases the recipient’s. Social behaviors are widespread in nature. A classic example is the altruism shown by the sterile workers of social insects such as ants, which sacrifice their own reproduction in order to rear the queen’s offspring. At first sight, altruism poses a problem for the genetical theory of natural selection, which seems to preclude the spread of a gene for reduced reproduction. Kin selection was devised by William Hamilton in the early 1960s to address this “problem of altruism.” The basic principle behind kin selection had been hinted at by Darwin, Fisher, and Haldane, but it was Hamilton who provided the first general model. Hamilton called his idea “inclusive fitness” theory, and it was later dubbed “kin selection” by Maynard Smith in 1964. For most purposes, the two can be considered identical, although inclusive fitness theory technically includes kin selection theory because the relatedness it invokes need not involve kin (genealogical relatives). Kin selection theory solved the problem of altruism by showing that a gene for altruism can spread if altruism is directed at individuals likely to bear the same gene. By definition, kin are likely to share genes. So, a gene for altruism can spread if altruism is directed at kin and the loss of gene copies through the actor’s decreased reproduction is more than offset by the gain in gene copies through the increased reproduction of the recipient. The algebraic version of this condition is termed “Hamilton’s rule.” Although kin selection theory was devised to explain altruism, it also applies to the other forms of social behavior such as selfishness. The theory is therefore now widely used to investigate and explain many kinds of social behavior in living organisms as diverse as bacteria and human beings.

Social Evolution, Hamilton’s Rule, and Inclusive Fitness

2018 ◽  
Author(s):  
Samir Okasha
Keyword(s):  
Social Behaviour ◽  
Social Evolution ◽  
Inclusive Fitness ◽  
Popular Approach ◽  
Hamilton’S Rule ◽  
Inclusive Fitness Theory ◽  
The Social ◽  
Social Encounters ◽  
The Individual ◽  
Hamilton's Rule

Inclusive fitness theory, originally due to W. D. Hamilton, is a popular approach to the study of social evolution, but shrouded in controversy. The theory contains two distinct aspects: Hamilton’s rule (rB > C); and the idea that individuals will behave as if trying to maximize their inclusive fitness in social encounters. These two aspects of the theory are logically separable but often run together. A generalized version of Hamilton’s rule can be formulated that is always true, though whether it is causally meaningful is debatable. However, the individual maximization claim only holds true if the payoffs from the social encounter are additive. The notion that inclusive fitness is the ‘goal’ of individuals’ social behaviour is less robust than some of its advocates acknowledge.

scholarly journals War and peace: social interactions in infections

2014 ◽  
Vol 369 (1642) ◽  
pp. 20130365 ◽  
Author(s):  
Helen C. Leggett ◽  
Sam P. Brown ◽  
Sarah E. Reece
Keyword(s):  
Species Interactions ◽  
Disease Ecology ◽  
Social Evolution ◽  
Inclusive Fitness ◽  
Experimental Tests ◽  
Microbial Pathogens ◽  
Emergent Properties ◽  
Social Strategies ◽  
Inclusive Fitness Theory ◽  
Social Behaviours

One of the most striking facts about parasites and microbial pathogens that has emerged in the fields of social evolution and disease ecology in the past few decades is that these simple organisms have complex social lives, indulging in a variety of cooperative, communicative and coordinated behaviours. These organisms have provided elegant experimental tests of the importance of relatedness, kin discrimination, cooperation and competition, in driving the evolution of social strategies. Here, we briefly review the social behaviours of parasites and microbial pathogens, including their contributions to virulence, and outline how inclusive fitness theory has helped to explain their evolution. We then take a mechanistically inspired ‘bottom-up’ approach, discussing how key aspects of the ways in which parasites and pathogens exploit hosts, namely public goods, mobile elements, phenotypic plasticity, spatial structure and multi-species interactions, contribute to the emergent properties of virulence and transmission. We argue that unravelling the complexities of within-host ecology is interesting in its own right, and also needs to be better incorporated into theoretical evolution studies if social behaviours are to be understood and used to control the spread and severity of infectious diseases.

scholarly journals Kin and group selection are both flawed but useful data analysis tools

2019 ◽  
Author(s):  
jeff smith ◽  
R. Fredrik Inglis
Keyword(s):  
Empirical Research ◽  
Social Evolution ◽  
Multilevel Selection ◽  
Selection Theory ◽  
Fitness Effects ◽  
Data Driven Approach ◽  
Biological Interpretation ◽  
Multilevel Selection Theory ◽  
Microbial Systems ◽  
Analytical Tools

AbstractFor understanding the evolution of social behavior in microbes, mathematical theory can aid empirical research but is often only used as a qualitative heuristic. How to properly formulate social evolution theory has also been contentious. Here we evaluate kin and multilevel selection theory as tools for analyzing microbial data. We reanalyze published datasets that share a common experimental design and evaluate these theories in terms of data visualization, statistical performance, biological interpretation, and quantitative comparison across systems. We find that the canonical formulations of both kin and multilevel selection are almost always poor analytical tools because they use statistical regressions that are poorly specified for the strong selection and nonadditive fitness effects common in microbial systems. Analyzing both individual and group fitness outcomes helps clarify the biology of selection. We also identify analytical practices in empirical research that suggest how theory might better handle the challenges of microbial data. A quantitative, data-driven approach thus shows how kin and multilevel selection theory both have substantial room for improvement as tools for understanding social evolution in all branches of life.

Evidence, Other Approaches, and Further Topics

2015 ◽  
Author(s):  
James A.R. Marshall
Keyword(s):  
Social Evolution ◽  
Inclusive Fitness ◽  
Kin Recognition ◽  
Replicator Dynamics ◽  
Empirical Support ◽  
Natural World ◽  
Price Equation ◽  
Evolution Theory ◽  
Inclusive Fitness Theory ◽  
The Many

This book has examined the genesis, the logic, and the generality of social evolution theory. In particular, it has presented evolutionary explanations of the many social behaviors we observe in the natural world by showing that William D. Hamilton's inclusive fitness theory provides the necessary generalization of classical Darwin–Wallace–Fisher fitness. This concluding chapter discusses the limitations of the analyses presented in this book and assesses the empirical support for inclusive fitness theory, focusing on microbial altruism, help in cooperative breeders, reproductive restraint in eusocial species, and the evolution of eusociality and cooperative breeding. It also considers more advanced topics in social evolution theory, including sex allocation, genetic kin recognition, spite, and the evolution of organismality. Finally, it reviews theoretical approaches to studying social evolution other than replicator dynamics and the Price equation, such as population genetics, class-structured populations, and maximization approaches.

scholarly journals Male relatedness and familiarity are required to modulate male-induced harm to females in Drosophila

2017 ◽  
Vol 284 (1860) ◽  
pp. 20170441 ◽  
Author(s):  
Sally Le Page ◽  
Irem Sepil ◽  
Ewan Flintham ◽  
Tommaso Pizzari ◽  
Pau Carazo ◽  
...  
Keyword(s):  
Potential Role ◽  
Genetic Relatedness ◽  
Inclusive Fitness ◽  
Kin Recognition ◽  
Female Reproduction ◽  
Fitness Effects ◽  
Inclusive Fitness Theory ◽  
Indirect Fitness ◽  
Female Lifespan

Males compete over mating and fertilization, and often harm females in the process. Inclusive fitness theory predicts that increasing relatedness within groups of males may relax competition and discourage male harm of females as males gain indirect benefits. Recent studies in Drosophila melanogaster are consistent with these predictions, and have found that within-group male relatedness increases female fitness, though others have found no effects. Importantly, these studies did not fully disentangle male genetic relatedness from larval familiarity, so the extent to which modulation of harm to females is explained by male familiarity remains unclear. Here we performed a fully factorial design, isolating the effects of male relatedness and larval familiarity on female harm. While we found no differences in male courtship or aggression, there was a significant interaction between male genetic relatedness and familiarity on female reproduction and survival. Relatedness among males increased female lifespan, reproductive lifespan and overall reproductive success, but only when males were familiar. By showing that both male relatedness and larval familiarity are required to modulate female harm, these findings reconcile previous studies, shedding light on the potential role of indirect fitness effects on sexual conflict and the mechanisms underpinning kin recognition in fly populations.

scholarly journals The validity and value of inclusive fitness theory

2011 ◽  
Vol 278 (1723) ◽  
pp. 3313-3320 ◽  
Author(s):  
Andrew F. G. Bourke
Keyword(s):  
Kin Selection ◽  
Evolutionary Biology ◽  
Social Evolution ◽  
Inclusive Fitness ◽  
Evidence Base ◽  
Alternative Theory ◽  
Selection Theory ◽  
Inclusive Fitness Theory ◽  
Evolution Of Eusociality ◽  
Natural Selection Theory

Social evolution is a central topic in evolutionary biology, with the evolution of eusociality (societies with altruistic, non-reproductive helpers) representing a long-standing evolutionary conundrum. Recent critiques have questioned the validity of the leading theory for explaining social evolution and eusociality, namely inclusive fitness (kin selection) theory. I review recent and past literature to argue that these critiques do not succeed. Inclusive fitness theory has added fundamental insights to natural selection theory. These are the realization that selection on a gene for social behaviour depends on its effects on co-bearers, the explanation of social behaviours as unalike as altruism and selfishness using the same underlying parameters, and the explanation of within-group conflict in terms of non-coinciding inclusive fitness optima. A proposed alternative theory for eusocial evolution assumes mistakenly that workers' interests are subordinate to the queen's, contains no new elements and fails to make novel predictions. The haplodiploidy hypothesis has yet to be rigorously tested and positive relatedness within diploid eusocial societies supports inclusive fitness theory. The theory has made unique, falsifiable predictions that have been confirmed, and its evidence base is extensive and robust. Hence, inclusive fitness theory deserves to keep its position as the leading theory for social evolution.

scholarly journals Helping in cooperatively breeding long-tailed tits: a test of Hamilton's rule

2014 ◽  
Vol 369 (1642) ◽  
pp. 20130565 ◽  
Author(s):  
Ben J. Hatchwell ◽  
Philippa R. Gullett ◽  
Mark J. Adams
Keyword(s):  
Social Evolution ◽  
Inclusive Fitness ◽  
Empirical Studies ◽  
Alloparental Care ◽  
Hamilton’S Rule ◽  
Long Term Study ◽  
Inclusive Fitness Theory ◽  
Cooperatively Breeding ◽  
Helping Behaviour ◽  
Hamilton's Rule

Inclusive fitness theory provides the conceptual framework for our current understanding of social evolution, and empirical studies suggest that kin selection is a critical process in the evolution of animal sociality. A key prediction of inclusive fitness theory is that altruistic behaviour evolves when the costs incurred by an altruist ( c ) are outweighed by the benefit to the recipient ( b ), weighted by the relatedness of altruist to recipient ( r ), i.e. Hamilton's rule rb > c . Despite its central importance in social evolution theory, there have been relatively few empirical tests of Hamilton's rule, and hardly any among cooperatively breeding vertebrates, leading some authors to question its utility. Here, we use data from a long-term study of cooperatively breeding long-tailed tits Aegithalos caudatus to examine whether helping behaviour satisfies Hamilton's condition for the evolution of altruism. We show that helpers are altruistic because they incur survival costs through the provision of alloparental care for offspring. However, they also accrue substantial benefits through increased survival of related breeders and offspring, and despite the low average relatedness of helpers to recipients, these benefits of helping outweigh the costs incurred. We conclude that Hamilton's rule for the evolution of altruistic helping behaviour is satisfied in this species.

Eusocial Insects as a Model for Understanding Altruism, Cooperation, and Levels of Selection

2020 ◽  
Author(s):  
Heikki Helanterä
Keyword(s):  
Natural Selection ◽  
Division Of Labor ◽  
Social Insect ◽  
Social Evolution ◽  
Inclusive Fitness ◽  
Family Structures ◽  
Major Transitions ◽  
Eusocial Insects ◽  
Insect Societies ◽  
Evolutionary Success

If the logic of natural selection is applied strictly at the level of individual production of offspring, sterile workers in insect societies are enigmatic. How can natural selection ever produce individuals that refrain from reproduction, and how are traits of such individuals that never produce offspring scrutinized and changed through natural selection? The solution to both questions is found in the family structures of insect societies. That is, the sterile helper individuals are evolutionary altruists that give up their own reproduction and instead are helping their kin reproduce and proliferate shared genes in the offspring of the fertile queen. Selection in such cases is not just a matter of individual’s direct reproduction, and instead of own offspring, the currency of the evolutionary success of sterile individuals is inclusive fitness. The concept of inclusive fitness and the process of kin selection are key to understanding the magnificent cooperation we see in insect societies, and reciprocally, insect societies are key case studies of inclusive fitness logic. In extreme cases, such as the highly advanced and sophisticated societies of ants, honeybees, and termites, the division of labor and interdependence of colony members is so complete, that it is justified to talk about a new level of evolutionary individuality. Such increases in the hierarchical complexity of life are called major transitions in evolution. We see adaptations of the colony, rather than individuals, in, e.g., their communication and group behaviors. The division of labor between morphologically differentiated queens and workers is analogous to germline-soma separation of a multicellular organism, justifying the term superorganism for the extreme cases of social lifestyle. Alongside these extreme cases, there is enormous diversity in the social lifestyles across social insect taxa, which provides a window into the balance of cooperation and conflict, and individual reproduction and helping others, in social evolution. Over the last decades, social insect research has been an area where the theoretical and empirical understanding have been developed hand in hand, together with examples of wonderful natural history, and has tremendously improved our understanding of evolution.

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