JHBS—WILEY RIGHT BATCH Top of ID Journal of History of the Behavioral Sciences, Vol. 38(2), 133–156 Spring 2002 Published online in Wiley InterScience (www.interscience.wiley.com). DOI: 10:1002/jhbs.10033 ᭧ 2002 Wiley Periodicals, Inc. BRAVE NEW WORLDS: TROPHALLAXIS AND THE ORIGIN OF SOCIETY IN Base of 1st THE EARLY TWENTIETH CENTURY line of ART CHARLOTTE SLEIGH Trophallaxis, the process of feeding by mutual regurgitation amongst insects, was named by the North American entomologist William Morton Wheeler in 1918. I argue that en- tomologists, both before and after 1918, saw mutual feeding as an integral part of the behavioral whole of the nest, and moreover related its explanatory power to theories about human society. In particular, feeding behavior was seen as the key to the riddle of the origin of sociality. I show how entomologists’ precise interpretations of trophallaxis varied and explore the increasingly functional, sociological, and economic constructions of the phenomenon that they developed—without breaking with earlier tradition—into the early 1930s. The article ends by demonstrating how Aldous Huxley’s bleak vision of humanity in the novel Brave New World, and its ambiguous prescription for meaningful life amidst the trappings of modernity, has much in common with metaphors generated by those studying ants. ᭧ 2002 Wiley Periodicals, Inc. This article explores early twentieth-century models of social-insect societies by focusing on one, somewhat strange concept, the “social stomach,” and its associated behavior “troph- allaxis.” Trophallaxis was the process of mutual feeding by regurgitation, and its significance was described in detail by the Harvard-based myrmecologist, William Morton Wheeler (1865–1937). I shall argue that the term trophallaxis, though generated to explain a specifi- cally myrmecological phenomenon, was from the outset understood to cover general features of society. The first section of the article outlines the immediate chain of influences that led up to Wheeler’s description of trophallaxis in 1918. After this, the layers of context are gradually exposed to show why early twentieth-century accounts of ants were inextricably linked with contemporary explanations of human society. One of these is the tradition of social/organismic analogies, which related wholes and parts of various species in nature, generally within a political idiom. The history of this tradition is briefly sketched. Ants in their organized colonies were a particularly clear example of the holistic part-whole relation- ship. The concept of trophallaxis, like all aspects of myrmecology at this time, depended upon an essentially socialized understanding of ants, and as such was developed in conjunc- tion with the inter-specific social analogies which it offered, including the human. The precise analogies to be developed, of course, varied according to context. The psy- chiatrist Auguste-Henri Forel (1848–1931) interpreted mutual feeding holistically in a eu- genic setting. He expressed its human significance openly, as part of his optimistic faith in socialism. In contrast to this, Wheeler and the brothers Aldous and Julian Huxley developed a somewhatcynical economic model for thesame phenomenon. Aldous Huxley’s novel Brave New World (1932) portrays an obvious parallel between human and Hymenopteran sociology. Its source, Ants (1930), was written by his brother Julian—a biological colleague of Wheeler. This means that the novel can be used to illuminate the human dimensions of Wheeler’s work on trophallaxis. I argue that not only Aldous Huxley, but also his scientific sources were re- CHARLOTTE SLEIGH completed her Ph.D. at the University of Cambridge in 1999. Following a one-year fellowship at UCLA, she was appointed to a lectureship in the Centre for History and Cultural Studies of Science at the University of Kent at Canterbury. She writes on various aspects of the history of natural history and the life sciences in the nineteenth and twentieth century and has a particular interest in literature. Her book on ant imagery is forthcoming with Reaktion. short standard 133 Base of DF JHBS—WILEY LEFT BATCH Top ofRH 134 CHARLOTTE SLEIGH Base ofRH Top oftext assessing the world in which they lived through myrmecology, the study ofants. They con- Base oftext templated the possibility that humans, ants, and termites might all be viewed in an economic framework. They faced up to the thought that the individual might be meaningless—that his actions were in fact defined by the society in which he found himself, and not by his private intentions. FROM SOLITARY TO SOCIAL: “L’INSTINCT EDUCATEUR” The paper trail apparently leading to Wheeler’s description oftrophallaxis and its sig- nificance begins in France. In the latter part ofthe nineteenth century there arose across Europe a renewed passion for the observation and study of insect behavior (Sleigh, 1999; Drouin, forthcoming; Lustig, forthcoming). This fashion seized professional naturalists and amateur enthusiasts alike—indeed the two categories cannot truly be separated at this time. The study ofinsect psychology—as it was unproblematically known—led to the production ofmany scientific articles and books as naturalists began to focus on questions such as the relationship between instinct and intelligence, the flexibility or automaticity ofinsect behavior, and the problem ofinsect orientation. The best-known observer ofinsect lifeduring this period was Jean-Henri Fabre (1823–1915), whose 10-volume Souvenirs Entomologiques (1879–1907) was the object ofpopular acclaim and extreme scientific wariness. Charles Janet (1849–1932), a Parisian engineer, was another such researcher and had been a keen entomologist since childhood. In 1895 he managed to make more time for his interests and began a series ofpapers on ants, wasps, and bees, focusingmainly on the first two insects. Besides making detailed studies ofant anatomy, Janet designed an artificial ants’ nest that was a success at the Paris exhibition of1900, enabling exhibition visitors to obtain an improved insight into the life of ants. Researchers used his apparatus for many years thereafter. In 1895, Janet observed the following behavior amongst wasps: “One of the first actions [performed by the imago after hatching from its cocoon] . consists in gently tapping her mandibles on the head ofone ofthe first large larvae which she encounters, and in drinking the droplet ofliquid which the latter disgorges” (Janet, 1895, quoted in Berland, 1932, p. 158). This action could also be induced by the observer himself: Ifone lightly touches the heads oflarge larvae with a paintbrush or the tip ofa pencil, one can see them spreading their mandibles, and throwing them backwards, as ifto leave the necessary space between their mouth and alveolar partition to lodge an alimentary globule, and, at the same time, to disgorge a droplet oflimpid liqui d...asimilar globule is disgorged whenever a worker has just nibbled at the head ofa larva. (ibid., pp. 158–9) These descriptions were read by another French researcher, Emile Roubaud (1882–1962). Roubaud is best known for his medical-entomological work in West Africa, but having studied under the insect psychologist E. L. Bouvier, he maintained an interest in the behavior of solitary and social insects throughout his life. His periods in Africa afforded him plenty of opportunities to look at the lives ofthe Hymenoptera and to ponder the meaning ofJanet’s observations, amongst those ofother enthusiasts. One key question in insect psychology concerned the flexibility ofinsect behavior: Could insects appropriately alter their chains ofaction under unforeseencircumstances, or were their short patterns ofaction set in stone? In order to answer such questions, Roubaud (1916) looked at standard long JHBS—WILEY RIGHT BATCH Top of RH BRAVE NEW WORLDS 135 Base of RH Top of text solitary wasps, especially the subfamily Eumeninae (“potter wasps”). These wasps ordinarily Base of text went through a very predictable string of actions in creating a nest, laying an egg within it, provisioning the nest with prey, and then sealing it up. If the egg were removed, would the wasp still provision the nest and seal it up? If the nest were damaged, would it go back to the first step and repair it? Roubaud found that these insects were in fact able to alter their patterns of behavior if they were interrupted and concluded that insects were, in general, frequently able to dissociate strings of habits when conditions demanded. In doing this, Rou- baud proved to his own satisfaction that his compatriot Fabre was wrong in saying that insectan actions were absolutely automatic and would not alter even if their order were dis- rupted by the experimenter. Roubaud connected the ability to manifest flexible behavior with the variety of nesting and feeding techniques, which could actually be observed among various wasp species. He concluded that there were four basic methods of provisioning types undertaken: 1. Rapid and massive provisioning (i.e. before egg hatches) 2. Slower and massive provisioning (i.e. continuing for some time after hatching) 3. Direct, overseen raising of progeny by living paralyzed prey 4. Direct, overseen raising of progeny by malaxated prey (i.e. softened and rendered as a pellet) Insects could move from one stage to another because, as he had proved to his own satisfaction with the potter wasps, their actions were never completely fixed; under new environmental conditions, actions that had previously been associated could be dissociated and performed in a different way. For example, mother wasps would choose large prey in preference to small during times of abundance, since this took less effort overall. One important behavioral nov- elty was that progressive raising efforts (efforts e´ducateurs) tended to be exhibited during good seasons. Mother wasps were able to tailor their efforts to the needs of the growing larvae, rather than going to a lot of effort that might have been wasted. Every instinctual act was changed to economize effort. Roubaud saw these different behaviors, all which could be observed in Africa, as rep- resenting successive evolutionary stages in the development of insect sociality.