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

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.

2017 ◽  
Vol 7 (5) ◽  
pp. 20170005 ◽  
Author(s):  
Andy Gardner

A central feature of Darwin's theory of natural selection is that it explains the purpose of biological adaptation. Here, I: emphasize the scientific importance of understanding what adaptations are for, in terms of facilitating the derivation of empirically testable predictions; discuss the population genetical basis for Darwin's theory of the purpose of adaptation, with reference to Fisher's ‘fundamental theorem of natural selection'; and show that a deeper understanding of the purpose of adaptation is achieved in the context of social evolution, with reference to inclusive fitness and superorganisms.


Author(s):  
James A.R. Marshall

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.


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.


2016 ◽  
Vol 371 (1687) ◽  
pp. 20150085 ◽  
Author(s):  
Erol Akçay ◽  
Jeremy Van Cleve

Inclusive fitness has been the cornerstone of social evolution theory for more than a half-century and has matured as a mathematical theory in the past 20 years. Yet surprisingly for a theory so central to an entire field, some of its connections to evolutionary theory more broadly remain contentious or underappreciated. In this paper, we aim to emphasize the connection between inclusive fitness and modern evolutionary theory through the following fact: inclusive fitness is simply classical Darwinian fitness, averaged over social, environmental and demographic states that members of a gene lineage experience. Therefore, inclusive fitness is neither a generalization of classical fitness, nor does it belong exclusively to the individual. Rather, the lineage perspective emphasizes that evolutionary success is determined by the effect of selection on all biological and environmental contexts that a lineage may experience. We argue that this understanding of inclusive fitness based on gene lineages provides the most illuminating and accurate picture and avoids pitfalls in interpretation and empirical applications of inclusive fitness theory.


Author(s):  
Erin Treanore ◽  
Nathan Derstine ◽  
Etya Amsalem

Abstract Social behavior, although rare, is a highly successful form of living that has reached its most extreme forms in eusocial insects. A tractable framework to understand social evolution is the study of major transitions in social behavior. This includes the transitions between solitary to social living, from species exhibiting intermediate degrees of sociality to species exhibiting true sociality, and from primitive to advanced eusocial species. The latter transition is characterized by the emergence of traits not previously found in primitive eusocial species, such as fixed morphological differences between castes and task specialization within the sterile caste. Such derived traits appear to exist in a binary fashion, present in advanced eusocial species, and absent or rare in primitive ones, and thus do not exist in a gradient that is easily tracked and compared between species. Thus, they may not be viewed as valuable to explore ultimate questions related to social evolution. Here, we argue that derived traits can provide useful insights on social evolution even if they are absent or rare in species with a lower social organization. This applies only if the mechanism underlying the trait, rather than the function it regulates for, can be traced back to the solitary ancestors. We discuss two examples of derived traits, morphological differences in female castes and primer pheromones regulating female reproduction, demonstrating how their underlying mechanisms can be used to understand major transitions in the evolution of social behavior and emphasize the importance of studying mechanistic, rather than functional continuity of traits.


Author(s):  
John Maynard Smith ◽  
Eors Szathmary

Over the history of life there have been several major changes in the way genetic information is organized and transmitted from one generation to the next. These transitions include the origin of life itself, the first eukaryotic cells, reproduction by sexual means, the appearance of multicellular plants and animals, the emergence of cooperation and of animal societies, and the unique language ability of humans. This ambitious book provides the first unified discussion of the full range of these transitions. The authors highlight the similarities between different transitions--between the union of replicating molecules to form chromosomes and of cells to form multicellular organisms, for example--and show how understanding one transition sheds light on others. They trace a common theme throughout the history of evolution: after a major transition some entities lose the ability to replicate independently, becoming able to reproduce only as part of a larger whole. The authors investigate this pattern and why selection between entities at a lower level does not disrupt selection at more complex levels. Their explanation encompasses a compelling theory of the evolution of cooperation at all levels of complexity. Engagingly written and filled with numerous illustrations, this book can be read with enjoyment by anyone with an undergraduate training in biology. It is ideal for advanced discussion groups on evolution and includes accessible discussions of a wide range of topics, from molecular biology and linguistics to insect societies.


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. 20130365 ◽  
Author(s):  
Helen C. Leggett ◽  
Sam P. Brown ◽  
Sarah E. Reece

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.


2011 ◽  
pp. 43-67 ◽  
Author(s):  
Stephan Hartmann ◽  
Pedro Pinto ◽  
Thomas Runkler ◽  
João Sousa

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