Jumping into the River …

Author(s):  
Jonathan Birch

This chapter provides an introduction to the book. Some brief background on the aims and history of social evolution theory is followed by a brief discussion of Ernst Mayr’s proximate-ultimate distinction. There follows a short overview of the book as a whole. Part I of the book ‘Foundations’, aims to construct a coherent picture of the conceptual structure of social evolution theory, a picture that distinguishes the different explanatory roles of three distinct conceptual innovations due to W. D. Hamilton that are often run together: Hamilton’s rule, kin selection, and inclusive fitness. Part II of the book, ‘Extensions’, turns to the ways in which recent expansions in the explanatory domain of social evolution theory have generated new conceptual challenges.

Author(s):  
Samir Okasha

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.


2014 ◽  
Vol 369 (1642) ◽  
pp. 20130358 ◽  
Author(s):  
Joel W. McGlothlin ◽  
Jason B. Wolf ◽  
Edmund D. Brodie ◽  
Allen J. Moore

Hamilton's theory of inclusive fitness revolutionized our understanding of the evolution of social interactions. Surprisingly, an incorporation of Hamilton's perspective into the quantitative genetic theory of phenotypic evolution has been slow, despite the popularity of quantitative genetics in evolutionary studies. Here, we discuss several versions of Hamilton's rule for social evolution from a quantitative genetic perspective, emphasizing its utility in empirical applications. Although evolutionary quantitative genetics offers methods to measure each of the critical parameters of Hamilton's rule, empirical work has lagged behind theory. In particular, we lack studies of selection on altruistic traits in the wild. Fitness costs and benefits of altruism can be estimated using a simple extension of phenotypic selection analysis that incorporates the traits of social interactants. We also discuss the importance of considering the genetic influence of the social environment, or indirect genetic effects (IGEs), in the context of Hamilton's rule. Research in social evolution has generated an extensive body of empirical work focusing—with good reason—almost solely on relatedness. We argue that quantifying the roles of social and non-social components of selection and IGEs, in addition to relatedness, is now timely and should provide unique additional insights into social evolution.


2014 ◽  
Vol 369 (1642) ◽  
pp. 20130565 ◽  
Author(s):  
Ben J. Hatchwell ◽  
Philippa R. Gullett ◽  
Mark J. Adams

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.


2018 ◽  
Vol 4 (10) ◽  
pp. e1700741 ◽  
Author(s):  
Yusaku Ohkubo ◽  
Tatsuhiro Yamamoto ◽  
Natsuki Ogusu ◽  
Saori Watanabe ◽  
Yuuka Murakami ◽  
...  

Over the past decade, the cause of sociality has been much debated. Inclusive fitness [brin Hamilton’s rule (br−c> 0)] has been criticized but is still useful in the organization of a framework by elucidating mechanisms through whichbr(benefit × relatedness) becomes larger thanc(cost). The beeLasioglossum baleicumis suitable for investigation of this issue because of the sympatric occurrence of both social and solitary nesting in its populations. We show that a large part (approximately 92%) of the inclusive fitness of a eusocial worker can be attributed to the benefits of grouping. A 1.5-fold relatedness asymmetry benefit in singly mated haplo-diploids explains a small part (approximately 8.5%) of the observed inclusive fitness. Sociality enables this species to conduct foraging and nest defense simultaneously, which is not the case in solitary nests. Our results indicate that this benefit of grouping is the main source of the increased inclusive fitness of eusocial workers.


Author(s):  
Jonathan Birch

From microbes to humans, the natural world is full of spectacular examples of social behaviour. In the 1960s, W. D. Hamilton introduced three key innovations—now known as Hamilton’s rule, kin selection, and inclusive fitness—that changed the way we think about how social behaviour evolves, beginning a research program now known as social evolution theory. This is a book about the philosophical foundations and future prospects of that program. Part I, ‘Foundations’, provides a philosophical analysis of Hamilton’s core ideas, with some modifications along the way. We will see that Hamilton’s rule provides a compelling way of organizing our thinking about the ultimate causes of social behaviour; and we will see how, in inclusive fitness, Hamilton found a fitness concept with a special role to play in explaining cumulative adaptation. Part II, ‘Extensions’, shows how these ideas, when extended in certain ways, can help us understand cooperation in micro-organisms, cooperation among the cells of a multicellular organism, and culturally evolved cooperation in the earliest human societies. In all these cases and more, living things cooperate because they are related, where the concept of relatedness picks out relevant statistical patterns of similarity in the transmissible basis (genetic or otherwise) of social traits.


2009 ◽  
Vol 5 (2) ◽  
pp. 214-217 ◽  
Author(s):  
Joan M Herbers

Darwin identified eusocial evolution, especially of complex insect societies, as a particular challenge to his theory of natural selection. A century later, Hamilton provided a framework for selection on inclusive fitness. Hamilton's rule is robust and fertile, having generated multiple subdisciplines over the past 45 years. His suggestion that eusociality can be explained via kin selection, however, remains contentious. I review the continuing debate on the role of kin selection in eusocial evolution and suggest some lines of research that should resolve that debate.


2017 ◽  
Vol 4 (7) ◽  
pp. 170335 ◽  
Author(s):  
Jonathan Birch

This paper attempts to reconcile critics and defenders of inclusive fitness by constructing a synthesis that does justice to the insights of both. I argue that criticisms of the regression-based version of Hamilton's rule, although they undermine its use for predictive purposes, do not undermine its use as an organizing framework for social evolution research. I argue that the assumptions underlying the concept of inclusive fitness, conceived as a causal property of an individual organism, are unlikely to be exactly true in real populations, but they are approximately true given a specific type of weak selection that Hamilton took, on independent grounds, to be responsible for the cumulative assembly of complex adaptation. Finally, I reflect on the uses and limitations of ‘design thinking’ in social evolution research.


2014 ◽  
Vol 369 (1642) ◽  
pp. 20130362 ◽  
Author(s):  
Andrew F. G. Bourke

Hamilton's rule is a central theorem of inclusive fitness (kin selection) theory and predicts that social behaviour evolves under specific combinations of relatedness, benefit and cost. This review provides evidence for Hamilton's rule by presenting novel syntheses of results from two kinds of study in diverse taxa, including cooperatively breeding birds and mammals and eusocial insects. These are, first, studies that empirically parametrize Hamilton's rule in natural populations and, second, comparative phylogenetic analyses of the genetic, life-history and ecological correlates of sociality. Studies parametrizing Hamilton's rule are not rare and demonstrate quantitatively that (i) altruism (net loss of direct fitness) occurs even when sociality is facultative, (ii) in most cases, altruism is under positive selection via indirect fitness benefits that exceed direct fitness costs and (iii) social behaviour commonly generates indirect benefits by enhancing the productivity or survivorship of kin. Comparative phylogenetic analyses show that cooperative breeding and eusociality are promoted by (i) high relatedness and monogamy and, potentially, by (ii) life-history factors facilitating family structure and high benefits of helping and (iii) ecological factors generating low costs of social behaviour. Overall, the focal studies strongly confirm the predictions of Hamilton's rule regarding conditions for social evolution and their causes.


Author(s):  
James A.R. Marshall

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.


2011 ◽  
Vol 278 (1723) ◽  
pp. 3313-3320 ◽  
Author(s):  
Andrew F. G. Bourke

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.


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