scholarly journals Darwin's ‘one special difficulty’: celebrating Darwin 200

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.

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

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.


Author(s):  
Heikki Helanterä

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.


Author(s):  
James A.R. Marshall

This chapter examines social behaviors that are expressed conditional on the phenotype of others. David Queller argued that inclusive fitness analyses need to be done on a per-behavior basis, citing as an example the decision over whether to reproduce directly, and whether to aid a reproductive. Queller showed that inclusive fitness predictions are only sensible when one analyzes what an individual should do, given it finds itself in a particular behavioral role. The chapter first provides an overview of implicit and explicit conditionality and presents two classic examples: William D. Hamilton's greenbeard traits and Robert Trivers's theory of reciprocal cooperation. It also introduces an extension of Hamilton's rule to deal with explicitly conditional behaviors; this extension features a measure of phenotypic assortment that appears not to be the classic genetic relatedness of Hamilton's rule.


Author(s):  
James A.R. Marshall

This chapter examines how the logic of inclusive fitness theory can be mathematically formalized using the Price equation, and how that formalization can be used to derive Hamilton's rule in its simplest form, as applied to unconditional behaviors having additive effects on fitness. Various biological phenomena, such as sex allocation and working policing within eusocial insect colonies, have been analyzed by considering what strategies maximize individuals' inclusive fitness, and how observed social behaviors should correlate with quantities such as relatedness. The chapter derives Hamilton's rule by introducing some notation for the effects of behaviors on fitnesses of individuals that interact socially, to make explicit precisely how genes (and later phenotypes) affect fitness, and to give a general form of Hamilton's rule that will apply to any (unconditional, additive) behavior regardless of its details. It shows that inclusive fitness is a genuinely novel extension of the classical fitness studied by Charles Darwin, R. A. Fisher, and others.


2000 ◽  
Vol 355 (1403) ◽  
pp. 1647-1655 ◽  
Author(s):  
David C. Queller

Many of the major transitions in evolution involved the coalescence of independent lower–level units into a higher organismal level. This paper examines the role of kinship, focusing on the transitions to multicellularity in animals and to coloniality in insects. In both, kin selection based on high relatedness permitted cooperation and a reproductive division of labour. The higher relatedness of haplodiploid females to their sisters than to their offspring might not have been crucial in the origin of insect societies, and the transition to multicellularity shows that such special relationships are not required. When multicellular forms develop from a single cell, selfish conflict is minimal because each selfish mutant obtains only one generation of within–individual advantage in a chimaera. Conditionally expressed traits are particularly immune to within–individual selfishness because such mutations are rarely expressed in chimaeras. Such conditionally expressed altruism genes lead easily to the evolution of the soma, and the germ line might simply be what is left over. In most social insects, differences in relatedness ensure that there will be potential conflicts. Power asymmetries sometimes lead to such decisive settlements of conflicts that social insect colonies can be considered to be fully organismal.


2020 ◽  
Author(s):  
António M. M. Rodrigues

AbstractInclusive fitness theory has transformed the study of adaptive evolution since 1964, contributing to significant empirical findings. However, its status as a theory has been challenged by the proposals of several alternative frameworks. Those challenges have been countered by analyses that use the Price equation and the regression method. The Price equation is a universal description of evolutionary change, and the partitioning of the Price equation using the regression method immediately yields Hamilton’s rule, which embodies the main tenets of inclusive fitness. Hamilton’s rule captures the intensity and direction of selection acting on social behaviour and its underlying causal structure. Recent work, however, has suggested that there is an anomaly in this approach: in some cases, the regression method fails to estimate the correct values of the variables in Hamilton’s rule and the causal structure of the behaviour. Here, I address this apparent anomaly. I argue that the failure of the simple regression method occurs because social players vary in baseline fecundity. I reformulate the Price equation and regression method to recover Hamilton’s rule and I show that the method correctly estimates its key variables. I show that games where baseline fecundity varies among individuals represent a more general set of games that unfold in class-structured populations. This framework supports the robustness and validity of inclusive fitness.


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