Trophallaxis and the Origin of Society in the Early Twentieth Century

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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 entomologists, 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 “trophallaxis.” 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 relationship. 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 psychiatrist Auguste-Henri Forel (1848–1931) interpreted mutual feeding holistically in a eugenic 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

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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 significance 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 novelty 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. All this evolution of the raising instinct (l’instinct e´ducateur), driven entirely by individual interests, culminated in social behavior:

In the totality of the tribe of the Eumeninae, one can pick out the traces of a continuous evolution of the raising instinct, which has perfected itself in certain types up to the stage now observed among social wasps, in accordance with certain essential inﬂuences. (Roubaud, 1916, p. 80)

Thus Roubaud suggested that, under the appropriate conditions, wasps manifested the four successive provisioning behaviors, progressing by reason of individual (butflexible) instinct from solitary to social behavior. In the fourth and final case, there was potential for sociality as the young were raised in a cohort. When Roubaud looked at social wasps, he concluded that the mutual feeding observed by Janet was the thing actually fulfilling the possibility of keeping the fully social nest together:

One may see the [worker] females pass back and forth three or four times in front of a lot of larvae to which they have given nutriment, in order to imbibe the secretion. The insistence with which they perform this operation is such that there is a ﬂagrant dispro- short portion between the quantity of nourishment distributed among the larvae by the females standard long JHBS—WILEY LEFT BATCH

Top of RH 136 CHARLOTTE SLEIGH Base of RH Top of text and that of the salivary liquid which they receive in return. There is therefore a real Base of text exploitation of the larvae by the nurses. (Roubaud quoted in Wheeler, 1918, p. 321)

In other words, the larvae offered salivary secretions to their adult sisters, and because they were so tasty, the sisters stayed around, looking after the next generation of hatchlings. Why did Roubaud make this connection between extant behavior and evolution? An obvious answer would be to cite the then-recent Darwinian perspective on behavior. However, insect psychologists were more indebted to Lamarckian philosophy than they were to Dar- win’s. In particular, they saw mental adaptations, or psychic energies, as driving physical evolution. Lamarckian psychic evolution can be clearly discerned in the writing of other insect psychologists of the era, such as Roubaud’s teacher, E. L. Bouvier, or the director of the laboratory of psychology at the Sorbonne, Henri Pie´ron (Bouvier, 1922; Pie´ron, 1929).1 Pie´ron described a metaphysical and evolutionary succession from the apparently simple to complex forms of psychology. All might be seen in terms of energy, according to him. Even the stimulus-response circuit might be construed in this way: Energy was brought by the stimulus and transformed by a muscular reaction. (This held as a meaningful model even though the energy brought in by the stimulus, e.g. as light, was millions of times less strong than that given off, typically by muscular action.) In seemingly more complex psychology, it might be the case that the stimulus was itself internal. The inner system of response varied in complexity and form, and could be called “reflex” or “instinctive” activities accordingly, without there being any essential difference between the two. Like Roubaud, Pie´ron had a strong sense of an internal economy of mental energy, and the need for a co–ordination of the mechanisms that lay ready to be tripped off by a stimulus. Reflexes were gradually transformed into instincts, whether responding to external stimuli or internal (e.g. cyclic) prompts. Pie´ron considered that there were many “hereditary equipments” amongst insects and that they were of particular importance for this kind of animal. Thus Pie´ron, who can plausibly be interpreted in the same light as Roubaud, described a Lamarckian evolution, driven by psychological or behavioral demands, and regulated by the psychic economy. Two years after Roubaud’s 1916 paper, the American William Morton Wheeler followed on by characterizing mutual feeding as the means by which society had evolved amongst the Hymenoptera (Wheeler, 1918). Like Roubaud, Wheeler characterized mutual feeding as a parasitic relationship within the colony and assigned it a crucial role in the very origin of insect sociality. He was happy to quote Roubaud extensively to this effect: [T]his peculiar family symbiosis . . . characterized by reciprocal exchanges of nutriment between larvae and parents . . . is the raison d’eˆtre of the colonies of the social wasps. The associations of the higher Vespids has, in our opinion, as its first cause the trophic exploitation of the larvae by the adults. (Roubaud quoted in Wheeler, 1918, p. 322) Roubaud’s chosen subjects, the solitary wasps, were considered to be the phylogenetic precursors of the ants (Wheeler’s specialism), in terms of taxonomy, physiology, and behavior. Wheeler was therefore favorably disposed toward Roubaud’s theory. It was a small step for him, with his Lamarckian view of behavioral evolution, to extend this theory from extant wasps to the evolutionary past of primitive ants, now extinct. Despite his apparently generous citation of Roubaud, Wheeler thought the process of mutual feeding so important that he

1. Wheeler co-edited the translation and publication of some of Lamarck’s manuscripts, remarking of his theory “Its vitality has been so great that despite repeated and apparently devastating onslaughts it still survives” (Wheeler short & Barbour, 1933, pp. xvii–xviii). See Sleigh (forthcoming). standard long JHBS—WILEY RIGHT BATCH

Top of RH BRAVE NEW WORLDS 137 Base of RH Top of text sought to correct Roubaud’s initial description of it, and to replace the Frenchman’s technical Base of text name for it (œcotrophobiosis), with his own mark of priority. Wheeler rechristened this exchange of nutrients with his own term, “trophallaxis” (Wheeler, 1918, p. 322).2

CONTEXTUAL INTERPRETATIONS OF MUTUAL FEEDING Now we turn to the context for this development of biological theory. By the time Wheeler invented his term “trophallaxis” in 1918, it was already well-established that actions within the nest had a social meaning and that there was a whole suite of communicative and feeding behaviors that went together to define and sustain the nest as a social entity.3 ErnstHaeckel was responsible for a major fashion of analogizing biological and social organisms in the latter part of the nineteenth century (Weindling, 1989a).4 Haeckel wrote an early monograph on Radiolaria (1860), a single-celled organism that united into symmetrical colonies. Gradually, he came to interpret Radiolaria’s nature as illustrative of the general lesson that social unification (in his context, German national unification) was a good thing. Furthermore, the study of multicellular organisms proved, according to Haeckel, the importance of duty, the subordination of the individual cell or person to the social whole and the necessity of the division of labor. The French psychologist Henri Pie´ron later described the simple sponge-like organisms, the Metazoa, as having incomplete individualization (i.e. unification), because their various segments sometimes showed independent nervous existence. The sea urchin, according to Jacob von Uexku¨ll, could be regarded as a “republic of reflexes” (Pie´ron, 1929, p. 28). Both Pie´ron and Uexku¨ll were following Haeckel in their metaphors, exploring the problem of unification in progressively more complex organisms by comparison to national or state organization. However, soon after Haeckel pursued analogies between the “cell state” and the organization of a putative German Empire (linked by the nervous system of the German Imperial Telegraph), a political philosopher in France began to consider the exact nature of the bonds that held humans and animals together in a group. This man was Alfred Espinas, and his Des Socie´te´s Animales (1877; 2nd ed., cited henceforward, 1878) was the touchstone for all works of group psychology during the following 50 years. Espinas wrote in a political context very different from Haeckel’s. He did not look forward to the foundation of an empire so much as back to the time of the French Revolution and to the nature of the republic that was thereby founded: the human bonds that precipitated it and held it together (Clark, 1984, pp. 42–43, 68–70 and 118–121). Espinas regarded communal life as “a normal, constant and universal fact” (Espinas, 1878, p. 9), and explained in Socie´te´s Animales the formation of both human and animal societies. To do this, he used functional associations—an explanation very different from the ideology of natural progress that marked even Haeckel’s earliest treatments of phylogenetic evolution. Although Espinas claimed the phylogeny of divergent social forms was more like the branches of a tree than a hierarchy, he organized his book in terms of more- or-less ascending organizations. The first of these was accidental associations between different species and included examples of parasitism and mutualism. Next, he worked upward

2. Bouvier described a phenomenon almost identical to trophallaxis which he called “trophobiosis.” (Bouvier, 1922, pp. 333–335.) 3. The earlier work of John Lubbock, which focused on the psychological attributes of individual ants, is therefore no part of this particular story. 4. Rudolf Virchow and other physiologists had previously encouraged interested parties to consider the German short states as bodies requiring medico-physiological policing. See Boyd, 1991, and Weindling, 1989b. standard long JHBS—WILEY LEFT BATCH

Top of RH 138 CHARLOTTE SLEIGH Base of RH Top of text on same-species consociations, starting with simple-celled organisms (like Radiolaria) that Base of text formed colonies for nutritional purposes. These were mostly marine creatures such as sponges, which did not exhibit any form of cellular differentiation but aggregated to form simple sacs or tubes in which digestion could occur, to their mutual benefit. Next came the communities that formed for reasons of reproduction. There were three kinds of these: conjugal societies that came together simply for mating; maternal, domestic societies (such as the ants); and paternal, domestic societies (birds and mammals).5 Finally, Espinas described the most truly social form of existence, the “relational” life of the tribe, which was exhibited by human beings. In the second edition of Socie´te´s Animales (1878), Espinas drew extensively on myrmecological studies in order to make his case. The differences between intraspecific and interspecific societies were given in the final section of Socie´te´s Animales, which spelled out the general laws of sociality (Espinas, 1878, pp. 519–521). The two kinds of consociation worked by similarity and by a delegation of functions, respectively. Emile Durkheim, who wrote in the same sociological social-psychological tradition as Espinas, based his two kinds of societies in The Division of Labor in Society (1893; cited as 1964) upon almost identical concepts.6 “Mechanical solidarity” he defined as solidarity based on the similarity of a society’s members, and the superior “organic solidarity” was the result of labor divided amongst the differently able members of a society. Mechanical solidarity was the property of the most primitive communistic human societies.7 Durkheim actually cited Espinas’ Socie´te´s Animales as explaining the causes of mechanical solidarity, meaning, presumably, aggregation for the purposes of nutrition and reproduction. Both authors analogized societies based on “organic solidarity” in similar ways, too. As Espinas compared the ant-hill to the mammalian brain, so Durkheim analogized the society with divided labor to the body with its organs, specialized for their different tasks. The comparisons were functional and quasi-teleological. In other words, both Durkheim and Espinas thought about component parts and the functions they played in maintaining the phenomenon of the stable society. If Durkheim saw society as a body, Espinas saw it as a mind. The true society, resulting from functional associations, was defined by Espinas as “a living consciousness, or an organism of ideas.” He considered the ant colony to be “truly, a single thought in action (albeit diffuse)” (Espinas, 1878, pp. 531–532). He claimed that all societies existed as thinking organisms because their “ideas and impulsions” (the two components of consciousness) were communicable and could be accumulated. Ideas could be imitated, and emotions could spread by sympathy, almost like an echo; the greater the number of individuals, the greater the force of these impulsive repercussions. By pooling these aspects of consciousness, an animal society could possess many more acts and specializations adapted to the exigencies of life than could an isolated individual of the same species. Eventually, the effect of the group was exercised as a function upon the members whose ancestors had originally formed it. In a true group, the individual was the work rather than the author of society; “what is more,” Espinas continued, “the individual’s action is limited to a short time, while the collective action weighs

5. Compare againstEngels, 1940 (ﬁrstpublished 1884): Engels covers previous theoriesof maternal-basedso- cietal evolution and advances his own, in a striking parallel to the queen-dominated evolution of the formicary. 6. This categorization derives from Farr, 1996, which distinguishes historically between psychological and sociological hermeneutics of social psychology. 7. The seemingly obvious question that Durkheim does not address is this: Should ants not, with their organic solidarity, or division of labor, be regarded as superior to “primitive” human societies? While Durkheim does not address this question explicitly, I have argued elsewhere that ants presented Europeans with precisely this paradox short at the beginning of the twentieth century (Sleigh, 2001). standard long JHBS—WILEY RIGHT BATCH

Top of RH BRAVE NEW WORLDS 139 Base of RH Top of text upon the individual with all the weight of acquired instincts and changes in structure obtained Base of text during the past of the race” (Espinas, 1878, p. 542). Although subsequent authors took issue with Espinas for his supposedly mystical “group-mind” theories, it can be seen that his vo- cabulary was raised upon functional theories to explain the formation and persistence of societies.8 Wheeler’s most obvious contribution to societal-organismic analogies came in 1910, with his famous talk given at the Marine Biological Laboratory, Woods Hole. In this, he deﬁned the ant colony as a unitary organism, or “superorganism” (Wheeler, 1911).9 By making such a comparison, Wheeler was aligning himself with the developmental, holist traditions he had encountered at the Naples Zoological Station, and the quasi-Haeckellian monism he found in his mentor, Auguste Forel (see below). But Wheeler was also staking a claim for the social importance of myrmecology, as well as attempting to place experimentation with the ant-colony on a par with the increasingly popular laboratory-based approaches to more conventional organisms. Thus philosophers and biologists around the turn of twentieth century had been thinking about two related concepts, cultural mirror-images of one another. One set of questions concerned the individual, complex organism as an organized aggregation, while another considered the social aggregation (whether animal or human) as an essentially unitary, organismic phenomenon. These may be regarded as mirror-images of one another because of their shared background of political aims, their overlapping literature and their common context of evolutionary (and, to some extent, developmental) biology.

AUGUSTE FOREL AND THE SOCIAL STOMACH The Swiss psychiatrist Auguste Forel (1848–1931) was, among other things, a eugen- icist, a socialist, a feminist, a paciﬁst and a monist. His love of ants, stretching back to early boyhood, predated all of these commitments, but came to illustrate them all for him (Forel, 1937). In the time left by his psychiatric practice and writing, and particularly after his retirement, Forel published proliﬁcally on ants, a vocation that culminated in his magnum opus Le Monde Social des Fourmis (1921–22; henceforward cited in English translation, Social World of the Ants Compared with that of Man, 1928). One particular aspect of ant behavior that fascinated Forel was the mutual solicitation for food that ants performed almost constantly, using their antennae. If a worker had eaten well, she was approached for food by her nest mates, who would stroke her with rapidly vibrating antennae. If the worker was allowed to feed on syrup stained with blue aniline dye, the abdomens of the entire nest would become tinged with blue after a time, demonstrating clearly the ubiquity of the phenomenon (see Figure 1). Termites engaged in this behavior too, exchanging partially digested wood with one another. This was considered to be the means by which they infected one another with the symbiotic gut protozoans necessary for the

8. Durkheim too was accused of proposing a mystical group mind to explain the function of society. See Douglas’ foreword in Mauss (Mauss, 1990, pp. xi–xii). Wheeler too felt the need to deny charges of mysticism: “the organization [of complex emergent wholes] is entirely the work of the components themselves and not [external] ‘entel- echies’ (Driesch)...or‘e´lan vital’ (Bergson).” (Wheeler, 1927, p 437; original emphasis). In the 1920s, American social psychologists were divided over the reality of the “group mind,” with Floyd Allport denying its existence, and Charles Horton Cooley and George Herbert Mead amongst the signiﬁcant minority who afﬁrmed it. See the special issue of JHBS of Fall 2000 (Vol. 36, No. 4), “Re–engaging the History of Social Psychology.” 9. Mitchell, 1995, gives a normative comparison between Wheeler’s metaphor and its revival by E. O. Wilson short and EliotSober, etc. standard long JHBS—WILEY LEFT BATCH

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FIGURE 1. Ants of the species Camponotus americanus feeding by regurgitation in the nest (Wheeler, 1910, p. 181). Feeding pairs are circled (added). digestion of wood (Wheeler, 1928c, p. 158; Brunelli, 1904). Some ants were extremely specialized with regard to social feeding. The “honey ants” belonged to a number of genera, usually found in xerophytic (dry) conditions; these were workers whose abdomens were completely distended with their sugary solution, which hung permanently from the roof of the nest ready to be tapped by their more mobile nest-mates (see Figure 2. Forel, 1928, vol. I, pp. 457–462; Wheeler, 1910, pp. 369–375; McCook, 1909, pp. 101–106; Harris, 1934). Other members of the nest were also fed by workers, notably larvae, but besides this, “guests” of other species in the nest, known as symphiles, received nourishment from the workers too. Forel pointed out the differences between the mutual regurgitation of honeydew between workers and the feeding of larvae or symphiles by workers. When exchange occurred between adult ants, the soliciting ant actively licked up the droplet from the motionless donor. When ants fed symphiles and larvae, the donor ant alone was active, though in some species the ant larvae could request food by moving their heads (Forel, 1928, vol. I, pp. 280–291 and 452–3). Forel (Forel, 1928, vol. I, pp. 235–236) and his fellow psychiatrist Rudolf Brun (Brun, 1923) interpreted the mutual type of feeding as highly pleasurable, almost erotic, for all the short ants concerned. “The regurgitating ant with her backward-ﬂung antennae has a look of ecstasy, standard long JHBS—WILEY RIGHT BATCH

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FIGURE 2. Honey antof thespecies Myrmecocystus horti-deorum regurgitating food to ordinary worker (Wheeler, 1910, p. 374). and undoubtedly feels as much enjoyment as the one which is swallowing,” judged Forel. The psychic reading that Forel gave to trophallaxis was crucial; the act relied upon the easy, pleasurable experience of the ant, and thereby gave a model to his whole theory of mind and its pathology. On the positive side, the sharing of food among workers seemed like a good example of socialism to Forel. Indeed, the act of mutual feeding was so central to his account of ants that he used an image of the act as the frontispiece to his two major volumes, The Social World of the Ants Compared with that of Man (see Figure 3). The moral comparison implied in the title was drawn in the aftermath of WWI. Forel recommended that if only humans could abandon their nationalist obsessions and cooperate like the socialist ants, peace would prevail. Mutual feeding was the metonymic behavior of the ants’ socialist community; they possessed, in effect, a “social stomach.” Man, argued Forel, ought to develop a social mind just as the ants had a social stomach. This would be achieved through an education in the importance of social work, the benefits of sexual (i.e. eugenic) hygiene, and of abstinence from alcohol. Forel was in close agreement here with another researcher half a world away: a woman named Adele M. Fielde. Fielde was an American woman, a sometime missionary who had been converted to humanism during her time of service in China. She too upheld the ants as a model of good governance, eugenics, and feminism. For her, the social labor of the workers—like her own efforts in China—was an acceptable, even desirable substitute for moth- erhood (Stevens, 1918). Though Fielde came at myrmecology from the background of an educated East Coast feminist, and Forel understood its significance from the perspective of North European monist, the act of mutual feeding suggested to both the humanistic aid that could be shared between people, the artificiality between national (though not racial) bound- short aries, and the benefits of socialism. standard long JHBS—WILEY LEFT BATCH

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FIGURE 3. short Frontispiece to Forel, The Social World of the Ants Compared with that of Man (Forel, 1921–2, vol. I). standard long 142 JHBS—WILEY RIGHT BATCH

Top of RH BRAVE NEW WORLDS 143 Base of RH Top of text On the negative side, the act of mutual feeding also suggested the origins of mental and Base of text social pathology to Forel. Foreign organisms in the nest—generally other kinds of insects— were able to induce ants to give up their sweet fluids or regurgitated food, thus exploiting their social behavior. (Forel, 1902; idem, 1928, vol. I, pp. 261–309). Some of these symphiles, such as insects of the order Thysanura (silverfish), apparently did this by means of “tapping the ants in a friendly way with their antennae” (see Figure 4). Other intruders, like Lomechusa and Atemeles beetles, were able to reap the benefits of living among the ants by offering them irresistible exudates. The ants would tend these parasites in exchange for their secretions, unaware of the fact that they destroyed the ants’ colony from within, starving or even eating its young, and disrupting its caste arrangements by causing abnormal development of those larvae that survived. Forel compared these two parasitic exploitations to his human concerns. In the first case, where ants were persuaded to support worthless members of society, Forel saw a clear parallel to society’s support of unfit individuals. In the second case, where ants were addicted to the secretions of the harmful invaders, Forel saw a reflection of human alcoholism (Forel, 1902; idem, 1928). Forel’s main trade was the treatment of alcoholics, and his model of the alco- holic’s mind was generated in tandem with his understanding of the ant’s instinct. The nest members simply could notresistthatsweetfluid,justas alcoholics could notforgo thequick buzz of drunkenness. If ants did this, their line would undergo deterioration, just as alcoholics would undergo personal and heritable degeneration. Forel, another Neolamarckian, was convinced that behavioral changes were heritable, and it just so happened that damage to the germ line through alcohol (which he called “blastophthory”) was the only mechanism for this aboutwhich he could be certain. The fact that ants did not automatically feed each other, but first solicited one another for food by means of their antennae, created another layer of meaning for the exchange. A second important aspect of social living for Forel was, accordingly, the so-called “antennal language” of the ants (Forel 1928, vol. I, pp. 239–242 and 447–450). Ants were forever waving their antennae at one another and touching them together; early in the early nineteenth century, Pierre Huber had concluded that this must be a form of communication (Huber,

FIGURE 4. short Thysanura begging for regurgitated food from ants (Fore, 1928, vol. I, p. 307). standard long JHBS—WILEY LEFT BATCH

Top of RH 144 CHARLOTTE SLEIGH Base of RH Top of text 1810). At the turn of the twentieth century, many researchers pointed to Huber as the pioneer Base of text of antennal language. They agreed that antennal communication was a mixture of physical (tapping) and chemical (scented) stimuli, but had very different opinions as to what antennal language meant in essence among the ants, and hence what it signified to the human researcher. The Jesuit priest Erich Wasmann (1905) saw a rather friendly communication in antennal language—ants had developed a profoundly social sense that predisposed them to communicate with their nest-mates in order to help one another. Typical communications might include an invitation to a newly discovered, food source, a warning of danger, a calming or soothing exchange, or a request for regurgitated honeydew. Forel, an ardent socialist, saw above all else “an incitement to collective work, passing from one individual to the other” in the actions of the ants. From a psychological point of view, he praised the ants’ language for being expressive yet simple; it did not permit the dissimulation that big words facilitated among humans. The stroking of antennae also formed part of the grooming routine of ants, or social toilet, as Forel called it (Forel, 1928, vol. I, pp. 455–7). Ants, despite having a “horror of dirt,” could not clean themselves completely and so were dependent upon one another in order to stay immaculate. Two ants would place themselves opposite one another and would begin by rubbing their antennae together. One would then jump upon the other’s back, and whilst the “antennal caresses” were continued, would use her mandibles to clean the back and legs of the ant beneath. Forel was much impressed with the hygiene of the ants; his account of their “social hygiene” corresponds directly with his eugenic views about human society (Jansen, 2001; Weindling, 1989b, pp. 61–152). Antennal stroking also had a function for the security of the nest. The antennae and their acute sense of smell provided ants with a means to establish and check the nest odor. If they were to avoid parasitism, invasion, and other forms of exploitation, ants ought only to give up food to fellow nest-members. To do this, they first had to check who was entitled to feeding, something that they apparently did by odor. Forel referred to this as the period of “quarantine.” Intruders were spotted by their alien scent and attacked. Slave-making ants, on the other hand, raised ants genetically different from their own, yet had to make them rec- ognizable and keep them safe by giving them the identifying scent of the nest. The constant attempts to smear and check the correct odor were the main thing that kept the ant society together. Adele Fielde again contributed to this area of study, conducting a series of related experiments in the first decade of the twentieth century. She performed many experiments where she removed ants from the nest, keeping them out for various lengths of time until they had lost their scent, and then returned them to see whether their erstwhile colleagues would “recognize” them, or whether they would be attacked as intruders with an alien scent. She also re-scented ants with various odorous chemicals to see what reaction they would get when re-introduced to the nest (Fielde, 1903a, 1903b, 1904, 1905a, 1905b). Forel’s most ambitious book, Social World of the Ants (1928), was constructed to highlight Fielde’s studies. Thus, Forel’s “social toilet” was communication of an implicit sort: the verification of nest-identity. Forel thought that the plasticity of scent acceptance or rejection was a natural analogy to the artificiality of national boundaries; once again he drew internationalist and pacifist morals from the life of ants. Thus Forel, within his context of eugenic and prohibitionist interests, saw the ants’ feeding as an intrinsically social process. Their very group existence was defined by “the physiological process of swallowing and regurgitating,...interrupted only by social work, short rest and warfare.” It was “being repeated every moment in the life of the ants” (Forel, 1928, standard long JHBS—WILEY RIGHT BATCH

Top of RH BRAVE NEW WORLDS 145 Base of RH Top of text vol. I, p. 236). Communication, grooming, scent-checking and mutual feeding were all related Base of text behaviors that deﬁned, maintained, and protected the nest.

WHEELER AND TROPHALLAXIS William Morton Wheeler got his professorship in entomology thanks to the profession- alizing efforts of the previous generation of economic entomologists in the United States (Evans & Evans, 1970; Sorensen, 1995; Palladino, 1996; Sleigh, 1999). However, like so many of these men, Wheeler did not want to waste his days testing insecticide in an agricul- tural laboratory. Wheeler had impeccable Chicago credentials in embryology, and his mind was set on higher things. After completing the scientific “grand tour” of Europe (including a period with Forel and another at the Naples Zoological Station) and a stint at the University of Texas, he took a job at Harvard’s school of applied biology in 1908. Wheeler was in no sense an applied biologist, yet for the remainder of his career he remained here, introducing to America European ideas about ants and their societies. In 1910, Wheeler published the book that sealed his reputation in myrmecology, called Ants. Feeding within the nest was central to this book’s account of ant-life, but Wheeler interpreted its importance in a sense rather different from Forel’s eugenic prohibitionism. Wheeler’s perspective was nevertheless social; the principal significance of feeding for him at this time was that it was commonly supposed to produce polymorphism, either through varying the quality or the quantity of larval food (Wheeler, 1910, p. 74). Polymorphism— the existence of multiple ant castes—was the major focus of Ants, and feeding, explaining its proximal causation, therefore took a crucial part in the monograph. The exact method of feeding larvae varied. Most ants fed their larvae on regurgitated foods, exceptthePonerinae, Dorylinae and some Myrmicinae. These sub-families were fed on solid scraps. Of these, the first two were regarded as primitive types, due to their incomplete manifestation of distinct castes. Something seemed to connect feeding by regurgitation with higher forms of ant sociality. In 1910, however, Wheeler was inclined to hedge the question about evolutionary causes. Instead, for the next eight years, he developed a Bergsonian ap- proach to biology. He was convinced that an organism was not a discrete entity—that it could encompass anything that maintained itself as an integrated and replicating whole. He maintained that the objects of science were in fact a flexible hierarchy of aggregations, from atoms to societies. A scientist could pick any level to study meaningfully. It was in this context that he composed his paper in which he described the ant colony as a “superorganism.” When Wheeler read Roubaud’s 1916 article, he combined it with his own holistic theories about ant-life, which were then at their peak. In the light of this, Wheeler began to reconsider some research he had done in 1901, very shortly after he became interested in myrmecology. Wheeler had been studying the primitive sub-family of Ponerine ants and noticed unusual behavior in the species Pachycondyla montezumae. The workers fed the larvae by placing insect fragments upon their ventral surface; when they did so, they were sometimes inundated by a “copious, colorless liquid” exuded by the larvae. This liquid was a digestive agent, but when secreted, it was “eagerly lapped up” by the nurse in question. The larval behavior was to be expected—larvae fed in such a primitive manner needed to digest their rough scraps in some way. But there was apparently no need for the adult nurses to partake in the secretions. The subfamily Ponerinae, looking more like the earliest fossil ants, were regarded as short less social: “the ancestral stirp of the higher subfamilies, and...theoldestexistingexpression standard long JHBS—WILEY LEFT BATCH

Top of RH 146 CHARLOTTE SLEIGH Base of RH Top of text of social life among the Formicidae” (Wheeler, 1910, pp. 225–226). Ponerines themselves Base of text were considered to have evolved from solitary wasps, such as Janet and Roubaud had studied (Forel 1903; Wheeler, 1910, p. 243–245). Thus whatWheeler had in thePonerines was a glimpse of the evolutionary path towards sociality, marked out by feeding behaviors in extant primitive Hymenoptera. Other ants, belonging to the more highly evolved sub-family Myrmicinae, fed their larvae regurgitated liquid rather than solid fragments. Yet Wheeler remarked that the newly-discovered Myrmicine species, Paedalgus termitolestes, nevertheless exhibited highly developed salivary glands in the larval form. This was more unexpected than when it was observed among the primitive Ponerines. Because Myrmicines were fed pre-digested liquid and not solid food, there was seemingly no need for glands producing digestive secretions in the larvae. Wheeler concluded that the larvae actually acted as a food store for the workers of the nest. Closer examination revealed a variety of strange organs in certain larval stages among a variety of Myrmicine species, organs which Wheeler christened “exudatoria.” All of these secreted substances were licked up by the nurses; their apparent care for the larvae was in their own interest. Social relations depended upon the stable feeding possibilities that it provided for adults, rather than any kind of duty towards the young or the group. Wheeler had now recreated the evolution of sociality from the progressive maternal feeding behaviors of sub-social wasps (Wheeler, 1928c, pp. 11–14), through to the reciprocal feeding of the Ponerines—the link between solitary wasps and modern ants—and found that this latter phenomenon remained a crucial feature of the most social or “eusocial” life. These last insect mothers raised the first brood themselves, sisters who then stayed on to care for subsequent hatchlings, bribed to stay on by that irresistible larval exudate, and becoming the worker caste. In 1918, Wheeler incorporated Roubaud’s theory, building on his own assumption that Ponerine behavior was the more primitive form of sociality and concluding that mutual feeding relations were the true, necessary cause of social forms of life. Wheeler exploited the morally unorthodox element of his theory about the origin of sociality in a satirical lecture written shortly afterward. “The Termitodoxa” (Wheeler, 1920) was originally given as a lecture to the American Society of Naturalists. It was narrated by the leader of the termites (one King Wee-Wee) who told the history of his species. As a race they had been degenerating until some of their biologists made recommendations about the running of society, implementing eugenic guidelines that corresponded to Wheeler’s repre- sentation of actual termite behavior. King Wee-Wee emphasized that trophallaxis was the very mechanism that enabled the emergence of trans-generational societies. Wheeler did not share Forel’s socialistic idealism; he was a humorously cynical man, a kind of Mencken of science. Instead he rather reveled in the suggestion that societies evolved for totally selfish reasons—that instead of being based upon parental love, termite society at least was based upon fatty dermal secretions. It was only bribery or perhaps blackmail that caused the addicted insects to fulfill their familial duties: Our ancestors, like other solitary insects, originally set their offspring adrift to shift for themselves as soon as they hatched, but it was found that the fatty dermal secretions, or exudates of the young, were a delicious food and that the parents could reciprocate with similar exudates as well as with regurgitated, pre-digested cellulose. Thenceforth parents and offspring no longer lived apart . . . (Wheeler, 1920, p. 115)10 Wheeler decided straight away in 1918 that that trophallaxis was not necessarily or short 10. Schneirla (1946, pp. 392–394) likewise described human breastfeeding as trophallaxis. standard long JHBS—WILEY RIGHT BATCH

Top of RH BRAVE NEW WORLDS 147 Base of RH Top of text essentially a larva-nurse exchange of food. Firstly, it encompassed the behavior of the sym- Base of text philes, insects and other kinds of organisms connected with the colony, even those located outside the nest. Indeed, Wheeler rejected Roubaud’s suggested name for the phenomenon partly on the grounds that it implied intraspecific relations only, not the full interspecific range (Wheeler, 1918, p. 322).11 Secondly, Wheeler’s construction of trophallaxis included non– food-based exchanges, especially odorific and communicative ones. In the remainder of this section I shall examine further these two features of Wheelerian trophallaxis. As soon as Wheeler had described the exudatoria of ant larvae, he confirmed their similarity to those of certain symphiles. It is important to notice that this similarity was not based on anatomy. Some of the glands of these symphiles were very similar to those of ant larvae. Others, however, exuded liquid from the so-called fat-body, dispersed underneath the whole of the chitinous cuticle and communicated to the surface by means of small pores. Wheeler glossed over these gross anatomical distinctions; what interested him was function:

...inthese [myrmecophilous] larvae the voluminous fat-body functions as a huge exudatorium....This at once suggests that in many ant larvae the general fat-body may have the same function. . . . [T]here is just as much reason for supposing that the fat-body may function as an exudate organ in the ant-larva as in the larvae of the Lomechusine myrmecophiles. (Wheeler, 1918, p. 315; emphasis added)

Wheeler’s point here was that there were three sources of liquid agreeable to worker ants (salivary glands, exudatoria, and the general fat-body) that were functionally equivalent forms of trophallaxis. Wheeler’s functional interpretation of mutual feeding allowed him to propose a complex web of trophallactic interactions, all of which came under the field of study of the scientist. From the outset, Wheeler considered trophallaxis to be an elastic social phenomenon, covering interspecific, parasitic, and even animal/plant relationships (see Figure 5). In an illustrative diagram, he placed trophallaxis between queen and larvae, or workers and larvae, at the center, echoing his claim that this was the primary form of trophallaxis in evolutionary terms. The next level of trophallaxis, adult–adult exchanges, was Forel’s key behavior. After this came exchanges with symphiles, then between ants of different species (which would occur when a different species had been brought as slaves into the nest), and finally other insects and plants outside the nest. At the final two expansions, ants would typically gain some food from the insect or plant in question, and in return, afford it a certain amount of protection. Wheeler considered this an “incomplete” kind of trophallaxis but nevertheless thought it was of suf- ficient relevance to his elucidation of trophallaxis to include it. Here we can again see Wheeler’s deep debt to the European traditions of holist biology:

Trophallaxis, originally developed as a mutual trophic relation between the mother insect and her larval brood, has expanded with the colony like an ever expanding vortex till ...theantshave drawn their living environment . . . into a trophic relationship. (Wheeler, 1918, p. 326)

All levels of nature were related by their common functional properties, whose laws were there for the scientist to study. Anticipating one possible objection to his 1918 paper, Wheeler suggested that trophal-

11. “Trophallaxis” was derived from two Greek words signifying “nourishment” and “interchange,” whereas Rou- short baud’s “œcotrophobiosis” owed its origin to the words for “home,” “nourishment,” and “life.” standard long JHBS—WILEY LEFT BATCH

Top of RH 148 CHARLOTTE SLEIGH Base of RH Top of text Base of text

FIGURE 5. Wheeler’s diagram of trophallactic relationships (Wheeler, 1918, p. 327 [adapted]).

laxis might be considered to involve non-substantial exchanges. One might choose to question the centrality of trophallaxis by pointing out that ants care just as assiduously for their pupae as they do for their larvae, though the larvae produce no exudate. But Wheeler claimed one could justiﬁably describe their “attractive odor” as a kind of volatile exudate. Thus trophallaxis even explained the care of pupae. Wheeler even tried to make a connection to the functional equivalent of cuticular exudates among humans, suggesting that pubic hairs were responsible for diffusing sexually attractive secretions (Wheeler, 1918, p. 315). In later years, Wheeler continued to emphasize the broadness of the trophallaxis concept, stressing its applicability far beyond the business of food exchanges amongst ants. In 1925, he gave a series of lectures (published as Wheeler, 1928c) at the University of Paris. The centrality of trophallaxis to myrmecology may be seen in the structure of the series. After a short general introduction, Wheeler deﬁned sociality as the possession of a worker caste, which standard long JHBS—WILEY RIGHT BATCH

Top of RH BRAVE NEW WORLDS 149 Base of RH Top of text was the morphological expression of the division of labor within the nest. There followed Base of text two lectures on polymorphism and then one on trophallaxis that described the definitive tasks and exchanges carried out by social insects. “I believe,” Wheeler claimed, “that [trophallaxis] constitutes the most essential characteristic of the social medium” (Wheeler, 1928c, p. 244). The series concluded with a lecture on the variety of exploitative inter-specific relationships made possible by trophallaxis. In the following analysis of those lectures, there are no new definitions of trophallaxis that cannot be discerned in Wheeler’s earlier writing.12 In his lectures, Wheeler continued to rebut the idea that trophallaxis was merely a nutritional exchange. In support of this point, Wheeler argued that taste and smell were intimately related senses (and anthropomorphic ones at that), and that therefore one might as well keep the word “trophallaxis” to refer to both “senses,” which were both chemoreception of one sort or another (Wheeler, 1928c, pp. 230–231). Furthermore, Wheeler countered earlier criticism that the exchange must be necessarily or immediately reciprocal. If the exchange was odor-based, it simply did not make sense to refer to the economy of individual exchanges as Roubaud had done with his food-based interactions. Wheeler had moved beyond Roubaud’s purely “selfish” account of maternal behavior as the creator of society. From eusocial maternal behavior, a novel entity emerged: society.13 Like Espinas before him, he now had a model for the origin and continued function of society that was not just the sum of its constituent members. In this context, Wheeler chose to discuss the researches of Fielde, which were now quite old, on the odor of the brood. Nevertheless, he wanted to emphasize the fact that trophallaxis was a form of communication, performing a role in the protection of the nest, besides that of reciprocal nutrition. But there was even more to it than this: keeping odor grouped together with trophallaxis allowed Wheeler to maintain his holistic metaphysical conception of the nest identity and its evolutionary history. Wheeler created a loop between behavioral cause and effect, and between the nest and the outside world: trophallactic feeding was both the behavior that created the polymorphic caste arrangements of the nest and the behavior that resulted from those arrangements. Through the exchange of food and other chemicals, the nest was maintained in holistic equilibrium. Workers, Wheeler argued, were essentially “hunger forms” of their kind. Although they might have some hereditary predisposition toward their caste, it was their lack of feeding as larvae that played the necessary role in their ontogeny. This deep-rooted hunger defined their adult lives, for they continued to display their enormous appetite in their constant search for food. The size of colony was, therefore, a function of the trophophoric field, the area of food available. In other words, it was intimately related with the outer world: “The social medium obviously comprises not only the regular activities of the workers in the nest and mainly the collection of food and its distribution among themselves, the queens, males and larvae but also the relations to food-yielding insects or plants in the trophophoric field and to the various guests . . . present within the nest itself” (Wheeler, 1928c, p. 229). Yet colony size was also determined by those very workers in the inner world of the nest, through their limited feeding of future workers. The worker caste was the “necessary creator of the social medium” (Wheeler, 1928c, p. 226). Trophallaxis was thus a self-regulating system of stimulus and

12. Thus I disagree with Wilson (1975, pp. 29–30) who claims that Wheeler expanded his deﬁnition of trophallaxis in 1928. Wheeler’s biographers (Evans and Evans, 1970, p. 257) claim that Wheeler’s latter treatment of trophallaxis was provoked by Wasmann’s criticisms; it seems to me, however, that his response was little more than a restatement of his former position. See also Lustig, forthcoming. short 13. On Wheeler and emergence philosophy see Wheeler 1926, 1927, and 1928a. See also Sleigh, 1999. standard long JHBS—WILEY LEFT BATCH

Top of RH 150 CHARLOTTE SLEIGH Base of RH Top of text response that completed the circuit between the inner world of the nest inhabitants and the Base of text outer world—a distinction which Wheeler was long happy to blur, as one can see from his trophallactic diagram of 1918. The trophallactic loop closed under adverse conditions, when the trophophoric field retreated inward to the nest itself and ants consumed their own kind in order that the superorganism might survive. The functioning of the ant-colony super-organism necessitated a communication between its constituent parts, a communication that mapped precisely onto the chemo-receptive phenomenon of trophallaxis.14 This phenomenon was a microcosm of the relational loops among the broader community, including symphiles, and other animals and plants. Trophallaxis was, for Wheeler, any kind of functional exchange among a community, which, holistically construed, went beyond the nest. At this point, one might remind oneself of the continuity between Wheeler and earlier investigators. The first obvious similarity concerns economy, a metaphor that connects Rou- baud’s Lamarckian adaptive psychological effort in individuals and Wheeler’s adaptive actions of the entire trophallactic loop. Just as Wheeler followed Espinas in his non-individualist account of nest relations, so he was not unique in construing the meaningful unit of the nest as containing other organisms than ants: Forel had defined the nest in this way in 1874 (p. iii). Wheeler also followed Durkheim in his physical comparison of trophallaxis to organismic organization: the circulation of nutrition and other stimuli in the superorganism corresponded to the chemical exchanges of blood and tissues in ordinary animals. But because this organization evolved from functional behaviors, it could also be compared to the functional sta- bility of human society. He asked his reader to consider the exchange of goods by coastal and inland people. According to Wheeler, trophallaxis was like this. It did not even have to entail an immediate reciprocation; the important thing was the exchange had the ultimate function of maintaining society in equilibrium. The only difference between the two species was that humans, an evolutionary novelty compared with ants, had not had time to incorporate their functional division of labor into their heritable morphology (Wheeler, 1928c, pp. 22– 23). Wheeler did not appear to see any reason in principle why this should not happen eventually, but more importantly, his functional outlook was supported by his extensive reading of the Italian sociologist Vilfredo Pareto (1848–1923). Wheeler began reading Pareto in 1925 or 1926 and responded to him with enormous enthusiasm and proselytic fervor. Briefly, Pareto’s sociology revolved around the assumption that most people do not live by rational thought, but by non-logical “residues.” Their individual irrationality—their propensity to be swayed by emotional appeals—meant that their behavior could be predicted or even controlled en masse (Pareto, 1980; first published in Italian 1916). Wheeler heartily concurred with this pessimistic assessment of humanity. Wheeler was able to give Pareto’s work a distinctively biological reading, retaining an individual hereditary basis for the mass proclivities of society; evolution explained the formation of society, which in turn explained the “individual” instincts of its component members. It is striking that Wheeler’s interpretation of Pareto resembles nothing so much as a pessimistic version of Espinas’ society-as-group-mind. The residues of the common man thus condemned him to a life that was functionally similar to the ant’s. Wheeler seems to have been convinced by Pareto that the lack of insight on the part of the common man fated society down the degenerate path of fixed instincts. He opened his Parisian lectures with a brief meditation on the “strange analogies” that obtained

14. Trophallaxis thus related to Von Frisch’s more recent work on honey bees whereby returning members of the short nest danced to communicate something about the ﬂoral scents on their bodies. See Evans & Evans, 1970, p. 262. standard long JHBS—WILEY RIGHT BATCH

Top of RH BRAVE NEW WORLDS 151 Base of RH Top of text between the societies of the social insects and of human beings, indicating explicitly that he Base of text had a Paretan critique in mind (Wheeler, 1928c, p. 2; see also Wheeler, 1928b). Elsewhere he suggested that social stultification was already coming about in post-Revolutionary Russia, on account of the fact that humans were being too perfectly socialized there (Wheeler, 1934). Wheeler himself was always keen to point out how the scientist eluded such generalizations and was a fundamentally anti-social creature; letters in his archive at Harvard reveal his wife and daughter’s frustration at his lack of social skills and Wheeler’s own perverse pride in the fact. (Male colleagues, however, enjoyed his company outside the constraints of the domestic sphere.) His colleague Pitirim Sorokin—with whom he created a Harvard course on comparative sociology—likewise complained about the trophallactic circuit that he had to com- plete upon arrival in Cambridge: “We had to ‘eat our way through Cambridge and Boston’ atmany lunches and dinners given by Harvard professors, “proper Bostonians”and various dignitaries of both cities. I have never much cared for the “social life” of going from party to party; nevertheless, like all newcomers to Harvard, I had to go through this ritual to comply with the established mores” (Sorokin, 1963, p. 241). And so Wheeler gravitated toward an economic, functional model to account for the properties of the ant society. Even if the proximate cause of caste behavior was partially or totally genetic, the origin of the behavior of individual ants within the context of the colony could not be explained in terms of individual inheritance. Moreover, there was nothing on- tologically unique about formicarian society that could not be applied to humans. There are various important differences between mutual feeding as it was construed by Wheeler and Forel. After reading Roubaud, Wheeler’s focus shifted from adult–adult reciprocal feeding to the exchange between adults and larvae. Forel, although recognizing that both phenomena existed, focused primarily on the exchange between adult workers as the normal act. In his Social World of the Ants (1928), Forel did not discuss the fact that, after bringing up her first brood, the queen ant was dependent on food regurgitated by her progeny (Wheeler, 1928c, p. 234). This was apparently a point on which Forel, a believer in the importance of hygienic maternity, did not wish to dwell. Nor, despite publishing after Wheeler’s 1918 paper, did he note the fact that the ant larvae exuded a fatty substance through their chitinous skin, which the nurses would lick up assiduously. In his chapter on “Ant life inside the nest,” Forel’s discussion of trophallaxis was confined to two species, making it sound fairly unusual (Forel, 1928, vol. I, pp. 462–3). Trophallaxis was given as the eighth of 11 worker behaviors; for Forel, the workers were the “true queens of the formicaries, as hard-working communists” (Forel, 1928, vol. I, p. 443). Exchanges involving larvae or the queen were of less interest to him; his sections on mutual feeding between workers, and on the honey-ants, construed as the logical evolutionary extension of mutual feeding, were longer sections and preceded the one on trophallaxis. Further on, Forel listed trophallaxis as one specific kind of feeding amongst many (Forel, 1928, vol. I, p. 516). Wheeler on the other hand, made trophallaxis the centerpiece of his entire analysis of ant feeding. The different choices of paradigmatic, reciprocal feeding arrangements arose partly, perhaps, because Wheeler’s principal interest was in the origin of sociality, while Forel was more interested in its maintenance. For contingent historical reasons, it happened that maternal–offspring relations were more likely to hold the key for Wheeler than worker–worker ones. One reason for difference about which we may be certain was that each man was searching for a different human angle on the phenomenon of mutual feeding. Forel was looking for a high-minded moral while Wheeler was happy to find something that appealed to his sense of humor—in this case, the idea that mothers parasitize their offspring and in so short doing unintentionally created society. Forel, having compared the ants’ addiction to symphilic standard long JHBS—WILEY LEFT BATCH

Top of RH 152 CHARLOTTE SLEIGH Base of RH Top of text exudates with alcoholism, could not bear to explain the raison d’eˆtre of his eugenic utopia Base of text by the same mechanism. Like Forel, Wheeler was now focusing on the workers as the defining feature of the ant colony, buthe had very differentreasons for doing so. Wheeler’s workers emerged from a parasitic relationship within family life. Forel’s workers evolved through a Lamarckian per- fection of co-operative behavior. Wheeler’s workers acted according to irrational, mass-dictated residues; Forel’s were models of civic-minded socialists. But both Forel and Wheeler conceptualized the nest as more than a mono-specific unit: Forel saw Neolamarckian reasons for the evolving relations for different insects within the nest, while Wheeler’s reading of Pareto confirmed his earlier functional holism. Both were participants in a continuous tradition of holistic, social constructions of individual ant behavior.

THE HUXLEYS AND THE ECONOMIC METAPHOR Aldous Huxley remarked that in his novel Brave New World he presented a “picture of society in which the attempt to recreate human beings in the likeness of termites has been pushed almost to the limits of the possible” (A. Huxley, 1968b, p. 266).15 In many ways, this comes as no surprise. With its rigid caste system, its merciless economy, and its dispensability of the individual, the New World is obviously very similar to life in the ant hill or termite mound. But why was Huxley so inspired to write about humans as social insects?16 The immediate answer to this question undoubtedly concerns his brother Julian Huxley’s book on ants, published in 1930, and the information that the brothers shared on social life in the insect world. Perhaps Julian might even have introduced Aldous to Wheeler’s satirical paper, “The Termitodoxa.” Beyond these obvious connections lie a deep debt to Pareto, shared by Aldous Huxley and Wheeler. Both writers took from the sociologist a “top-down” view of behavior (the idea that individual behavior is determined by the group) and a cynical attitude towards human nature. Julian Huxley’s book Ants (cited as 1949) contained no original research. What it did do was refocus current knowledge according to a functional, economic model. The book’s central chapter was called “food economics,” and acted as the lynch-pin to the whole. Julian Huxley compared the arrangement of ant and human societies thus:

The members of a human civilized community are tied together by economic bonds. . . . With us, of course, there is a universal medium of exchange in the shape of money, and by the use of such a medium we raise our system of mutual exchange of services to a far greater level of ﬂexibility than was possible by means of payment in kind or direct barter. The ants in an ant-colony are equally tied into a single economic whole; but the means by which this is accomplished are...unlike and, indeed alien to those employed by man...(J.Huxley, 1949, p. 39)

15. The story of Brave New World, briefly, is as follows. Society is eugenically bred into castes suited for different levels of work. Their obedience and happiness is maintained by subliminal conditioning, the drug “soma,” the constant distraction of entertainment, and by compulsory promiscuity that eliminates the passion of love. The book’s hero, Bernard Marx, comes to doubt whether such things are the route to true human fulfillment, a suspicion that is further provoked by his acquaintance with the “Savage” who, having had the misfortune of being born by natural means (a social taboo) has grown up free from the effects of the New World. The Savage is disgusted by his encounter with “civilization” and the book ends with his ritualized death. short 16. See Lepenies (1988) for an admirable discussion of the “scientific” and “literary” approaches to sociology, standard and Gaziano (1996) for an account of the circulation of metaphors between sociology and biology. long JHBS—WILEY RIGHT BATCH

Top of RH BRAVE NEW WORLDS 153 Base of RH Top of text Thus Huxley, like Wheeler and Forel, saw exchange as the thing holding the nest Base of text together.17 Huxley’s characterization of trophallaxis as an alien form of exchange was not entirely fair, as we can imagine his brother Aldous thinking. For what was soma but trophic exchange? It was precisely what King Wee-Wee the termite described in 1920: ...anelaborate exchange of exudates, veritable social hormones . . . which, continually circulating through the community, bound all its individuals together in one blissful, indissoluble, syntrophic whole, satisfied to make the comminution and digestion of wood and mud the serious occupation of existence, but the swapping of exudates the delight of every leisure moment. (Wheeler, 1920, p. 115) The capacity of the shared honey-dew to pacify the individual, and to maintain the greater harmony of the colony, meant that it performed exactly the same function as the ubiquitous drug of Aldous Huxley’s novel. Although freely available from the State, characters in the book gave soma to one other, quoting one of the relevant phatic aphorisms which they had been taught from childhood: “a gramme is better than a damn,” “a gramme in time saves nine,” and “one cubic centimetre cures ten gloomy sentiments” (A. Huxley, 1968a, pp. 58– 61 and 87–90). Soma is the all-purpose reward, consolation and pacification without which the inhabitants of the Brave New World could not stably exist. And so we come to the way in which the instincts of ants and humans were necessarily subordinated to the functional organization of their respective Brave New Worlds. Soma was even at the core of the “solidarity services” at the Community Singery. In a parody of the Eucharist, a cup of strawberry ice-cream soma was passed around, with the 12- times repeated formulation “I drink to my annihilation” (A. Huxley, 1968a, pp. 79–85). It was the individual that was annihilated in this ritual; he submerged himself into the community, losing even his sexual identity, and individuality in the “orgy-porgy” that followed. The ideology of Brave New World was thus that of the superorganism: “the social body persists although the component cells may change” (A. Huxley, 1968a, p. 94; idem, 1968b, p. 326). Like the individual ant, the inhabitant of Mond’s world was singly insignificant, and his role was constructed through the use of a social drug. The distribution of soma thus fulfilled the same social function as the honey-dew of the ants. Its dispersal in the Brave New World was trophallaxis by another name. More generally, Julian Huxley’s use of an economic system as the chief model of what holds together the insect society echoed the mandatory over-production and over-consumption that bound and perpetuated the system of the Brave New World. In Aldous Huxley’s world, people were exhorted to use fantastical goods and services, which were constantly being “improved” and added to. They were never supposed to mend things, but always to throw away and consume anew. This constant obsession with material goods gave everyone something literally and metaphorically in common—a version of socio–economics very different from the liberal Smithian lesson that used to be provided by the beehive (Hundert, 1994; cf. Wheeler, 1928c, pp. 306–7). In that case, each bee faithfully labored in order to contribute to the common good. Now, the ants seemed to be unwitting slaves of the system instead of its authors. The processes associated with the colony’s benefit were not their aim; rather, they were the addictions that happened to define it. There was a precedent for looking at so-called “alien exchange” within human society.

17. Julian Huxley had read Wheeler’s seminal work Ants some time between its publication in 1910 and writing short his own book, The Individual in the Animal Kingdom, which came outin 1912. In thisbook, Julian Huxley also played with the idea that anything could be called an organism so long as it performed the functions associated with standard organism-status. long JHBS—WILEY LEFT BATCH

Top of RH 154 CHARLOTTE SLEIGH Base of RH Top of text And here we take a step back into rather Victorian thought: the savage as mirror to civilized Base of text man in his baser moments. Bronislaw Malinowski devoted his 1922 book, Argonauts of the Western Pacific, to a form of exchange prevalent in the Trobriands named “Kula.” He wanted to dispel certain misconceptions about savage life: (1) that the savage was “happy-go-lucky” yet governed himself by rational, utilitarian motives, and (2) that he was not capable of organized labor or its corollary, trade and economics. On the contrary, argued Malinowski, there was a strict though non-utilitarian exchange which was a form of economics. He sum- marized: the whole tribal life [of the Trobriands people] is permeated by a constant give and take; . . . every ceremony, every legal and customary act is done to the accompaniment of material gift and counter gift; . . . wealth, given and taken, is one of the main instru- ments of social organization, of the power of the chief, of the bonds of kinship, and of relationship in law. (Malinowski, 1978, p. 167; original emphasis) What we have here is the ant as primitive version of human: irrationally economic. It is a surprising comparison, for we have just seen the ant-people of Brave New World as the apex of modernity. Yet Wheeler too read the Argonauts and cited Malinowski in his writings about the ants (Wheeler, 1926, note 4). His reading of Pareto had convinced him that there was not so very much to distinguish between the savage and the American—or, indeed, the ant. Meanwhile, Wheeler’s Paretan contemporaries in sociology, Homans and Curtis (1934), discussed the importance of institutions and social exchange among “civilized” humans. The Huxleys’ constructions of trophallaxis were thus indebted to a long tradition linking studies of ant and human societies. Espinas’ political philosophy was heavily based on myrmecological study, and in turn influenced the human sociology of Durkheim. Durkheim’s disciple Marcel Mauss took his master’s method as an injunction to record the entire exchange system of a human culture to see how it was held together, functionally, in the absence of a “market” as such, by acts of giving (Mauss, 1990; originally published 1950). Meanwhile, Malinowski was providing myrmecologists with a plausible model of non-rational economics for ants and termites. The publisher, C. K. Ogden, grouped this network of scientists and philosophers together in his International Library of Psychology, Philosophy and Scientific Method, a series of around 100 books, which featured titles by several insect psychologists, including Wheeler, as well as two titles by Malinowski. Ogden was moreover the translator of Forel’s Social World of the Ants, a personal friend of Malinowski, and partof a social scientific circle including the entomologist J. G. Myers and Julian Huxley. Ogden corresponded with and met Wheeler between 1927 and 1935; Wheeler praised Ogden’s philo- sophical grasp of biology and enjoyed his company. Trophallaxis, then, took its place in this scholarly/social context, and by the 1930s was constructed as a behavior born out of group- dictated, economic norms. A circle of intellectuals enabled metaphors to move back and forth between different areas of social and literary scholarship, human and myrmecological.

CONCLUSION:THE SIGNIFICANCE OF TROPHALLAXIS I have demonstrated that sociologically holist interpretations of ant behavior existed from the latter part of the nineteenth century. The exact reading of ant behavior varied for different authors in their speciﬁc contexts. Before Wheeler even coined the term trophallaxis, Forel saw mutual feeding as the most interesting feature of ant life, analogizing it in the negative sense to the behavior of alcoholics and in the positive sense to socialist practices. Forel, Fielde, and others had been treating it as part of a range of related behaviors—including odor short and communication—that deﬁned the nest. Wheeler too treated trophallaxis as a holist con- standard cept instantiating the emergent, social properties of the nest—properties that for him were long JHBS—WILEY RIGHT BATCH

Top of RH BRAVE NEW WORLDS 155 Base of RH Top of text somewhat elitist in their construction. Wheeler chose to place trophallaxis at the center of his Base of text humorous, cynical critique of human mores, while Julian and Aldous Huxley used the behavior, constructed in fundamentally Wheelerian terms, as the central metaphor for contemporary society’s basis in economics. Thus Wheeler and Julian Huxley were not unique in believing that trophallaxis, broadly construed, was the process that sustained the life of the formicary; what they did do was give trophallaxis the additional function of defining its constituent members through their social role. The social, economic reading of ants in the early twentieth century yielded an ambiguous moral: they were no longer a natural-theological complement to human reason. Instead ants reflected a fear that Westerners had not left primitive ways so very far behind—or contrarily, that modernity might consume them. Ant-human metaphors, which we have seen freely ex- changed between sociologists and entomologists in the early twentieth century, were clearly very complex. They involved civilized and savage humans, and constructed ants as simul- taneously super-civilized and ridiculously simple animals. The ant was both model and warning for Forel’s patients. Ants and the ant-like people in the Brave New World were the antithesis of what Wheeler and the Huxleys wanted to be, while the Savage was the Christ- like figure of true civilization. Yet Malinowski, Wheeler, and their colleagues in Ogden’s Library would not dream of looking up to the Trobrianders, the real life “savages” acting as their model for ants. Whatever the solution of this semiotic riddle, it is clear that trophallaxis long formed part of a socialized understanding of ants, and hence humanity.

ACKNOWLEDGMENTS I would like to acknowledge the helpful criticisms and suggestions I received from a number of people whilst this article was in its various draft forms, particularly Raymond Fancher, John Forrester, Ian Higginson, Abigail Lustig, and two anonymous referees.

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