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The Major Transitions in Evolution$
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John Maynard Smith and Eors Szathmary

Print publication date: 1997

Print ISBN-13: 9780198502944

Published to Oxford Scholarship Online: November 2020

DOI: 10.1093/oso/9780198502944.001.0001

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PRINTED FROM OXFORD SCHOLARSHIP ONLINE (oxford.universitypressscholarship.com). (c) Copyright Oxford University Press, 2021. All Rights Reserved. An individual user may print out a PDF of a single chapter of a monograph in OSO for personal use. date: 02 December 2021

The Origins of Societies

The Origins of Societies

(p.255) 16 The Origins of Societies
The Major Transitions in Evolution

John Maynard Smith

Eors Szathmary

Oxford University Press

The heat generated within a mound of the termite Macrotermes is carried upwards by a central air duct. The air then travels down along narrow channels close to the surface of the mound, where it is cooled, and where, as in a lung, oxygen and carbon dioxide are exchanged. The whole mound is an airconditioning system. Although the mound resembles a human building in having features ensuring the comfort of its inhabitants, it differs in that not one of its builders had a picture of the completed structure before building started. The structure emerged from the rule-governed behaviour of tens of thousands of interacting workers. In this, the mound resembles a human body rather than a human building. The body is built by the rule-governed actions of millions of cells. Nowhere is there anything resembling a blueprint of the body. At most, the genome is a set of instructions for making a body: it is not a description of a body. The resemblance between the development of an insect colony and of an organism has led to the concept of a ‘superorganism’. The analogy has some value. To the extent that individual ants, bees or termites have lost the capacity to reproduce, they can propagate their genes only by ensuring the success of the colony, just as somatic cells can propagate theirs only by ensuring the success of the organism. Hence, the colony can be expected to have features adapted to ensure its success, and it is reasonable to apply concepts of optimization to it, rather than to the individualas was done, for example, by Oster & Wilson (1978) in their book on insect caste systems. But for our purposes the concept of a super organism is of little use. To understand the origins of animal societies, we must ask how individuals capable of reproduction came to cooperate to the extent that most of them lost the ability to reproduce. To understand their maintenance, we must explain why they are not disrupted by cheating. Unlike somatic cells, the individual workers, although related, are not genetically identical. We would therefore expect within-colony conflict to be widespread, as indeed it is: examples discussed below concern egg laying by workers, and conflict over the sex ratio.

Keywords:   Haliotis, Lasioglossum, Myrmecocystus, haplodiploidy, kin recognition, kin selection, social contract game, superorganism

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