By strength of character alone no scientist more deserved recognition. Hölldobler was—and remains—the most honest scientist I have ever known. As we filled the hours around tedious replicate experiments with conversation (“Okay, that’s—ah—sixty-three seconds, the forager just entered the nest, got it? Now, I want to go back and say just one more thing about Hennig and the original idea of cladism…”), he endeavored to make every datum in his notebook, every nuance of expression in his published reports, as straight and transparent as he could. If he had a fault worth mentioning it was one which I shared and which made us the more compatible, an obsessiveness of habit in work, expressed as a sometimes unreasonable need to bring one subject to closure before going on to the next.
In science, obsessiveness under psychological control can be a virtue. To a degree I have not encountered elsewhere Hölldobler extended this urge to the design of experiments and a weighing of evidence. Many successful researchers stop with a single well-conducted procedure, which they repeat often enough for the overall result to be statistically persuasive. Then they are prepared to say in print, “I think it likely that such and such is the case.” Others hold back and ask, “What different experiment can I perform, using new kinds of measurements, that will test the conclusion more rigorously?” If they then perform the second procedure and find the second result consistent with the first, they conclude, “That pretty well proves it. Let’s move on.” Hölldobler is a member of the second group. But sometimes during our collaboration he would pause yet again and ask—to my consternation—“Is there a third way?” He did occasionally press on with yet another method. He was the only third-way researcher I have ever known.
He was a scientist’s scientist. He simply loved science as a way of knowing. I believe he would have practiced it without an audience or financial reward. He played no political games. If new data did not fit, he quickly shifted to a new position. He was one of the few scientists I have known actually willing to abandon a hypothesis. He was meticulous about crediting others, quick to praise research when it was original and solid, harsh in his rejection when it was slovenly. The tone of his conversation was explicitly and uncompromisingly ethical, a posture born neither of arrogance nor of self-regard, but from the conviction of his humanistic philosophy that without self-imposed high standards, life loses its meaning.
But a somber picture of Bert Hölldobler would be misleading. He was fun to be with, the younger brother I never had. In periods of relaxation we confided in all things, both scientific and personal. His manner and even his physical appearance were reassuring. Bearded as he approached middle age, he had a burgher’s pleasant countenance fitted upon the short muscular build of a gymnast, the latter a residue of his favorite sport as a youngster. He was deeply devoted to his family, somehow finding time to participate with his wife, Friederike, in every step of the rearing of their three sons. Science was not everything for Hölldobler. A gifted painter and photographer, a good musician, he enjoyed the arts as I never could, locked as I was into my unyielding workaholic’s momentum. In darker moments I envied him that.
Though considerably the younger man, he made me a better scientist. During our work in the laboratory and field, I found myself anxious to meet his standards and to let him know I was trying. I am by nature a synthesizer of scientific knowledge, much better than Hölldobler at this activity, but I confess that in my effort to make sense of everything, to fit every piece into my schemes somewhere no matter how procrustean the result, I often overlooked detail. Hölldobler did not. By temperament and training, he belonged to the Karl von Frisch tradition, expressed to me succinctly one day by Martin Lindauer, Frisch’s student and Hölldobler’s mentor, while I was visiting Würzburg. Lindauer said, grinning as he typically did when speaking of serious matters, “Look for the little things.”
That adjuration Bert and I followed many times during the Harvard years. In 1985 we made our first field trip together to Costa Rica. We drove north from San José to La Selva, the field station of the Organization for Tropical Studies. As we entered the rain forest, I used my more general knowledge of ants to find and identify colonies that might be of exceptional interest in behavioral work. I was looking for a quick and exciting payoff. One candidate was the primitive genus Prionopelta, which I found nesting in rotting logs. No colonies had ever previously been studied in life. I was eager to record the key facts of the social behavior of this ant, the kind of basic data that go comfortably into syntheses and evolutionary constructions. I plunged into the work with Bert’s assistance. We took notes on colony size, the number of queens, division of labor, and the kinds of insects and other small animals captured by the workers. We found, for example, that they preferred silverfishlike creatures called campodeid diplurans. In the course of our work, Bert’s attention fastened on fragments of old cocoon silk plastered on the walls of passageways of the Prionopelta nests. He asked, as much to himself as me, What does this mean? Nothing, just trash, I answered. When the new adults emerge from the cocoons, their nestmates throw out the silk fragments, and they don’t bother to stack them in separate garbage dumps. No, no, he said, look: the pieces are lined up as a smooth layer on the gallery walls. He went on, with close study of his own and the aid of a scanning electron microscope back at Harvard, to show that the cocoon silk is employed as wallpaper. It keeps the chambers of the moist walls drier than would otherwise be the case, and thereby protects the growing brood. Wallpapering was a technique of climate control previously unknown in ants.
Hölldobler again said, Look, some of the foraging workers appear to be moving more slowly while they drag their hind legs. Again I was unimpressed. Individual ants, I responded, often move slowly or erratically for no particular good reason. Nor is there any cause to believe that these primitive ants lay odor trails in any case. But Hölldobler persevered. He found that not only do the workers lay odor trails—by which they recruit nestmates to new nest sites—but the attractive substance comes from a previously unsuspected gland located in the hind legs. The pheromones are smeared in a line as the ants drag their hind legs over the ground. The existence of the gland provided an important clue to the evolutionary relationships of Prionopelta.
From two weeks of data gathered in the La Selva forest we wrote five scientific articles. During our years of collaboration we made uncounted other discoveries while grubbing around and talking back and forth. First one took the lead, then the other. Our partnership in most respects ended when Bert, after being approached by several institutions in Europe, was offered the professorship at Würzburg. He needed the sophisticated new equipment and skilled assistants promised him at the Theodor Boveri Institute. He had a desire, almost an obsession, to get inside the muscles, glands, and brains of ants to learn how these organs mediate social behavior and organization. He wanted to understand a thousand “little things” to make a great whole. That kind of enterprise is expensive, too much so, it appeared, for the National Science Foundation and private U.S. organizations. Most of their support was either inadequate or unstable or both, contingent on appropriations made in three- to five-year cycles of renewal. Although Bert’s applications to the NSF were consistently given the highest ratings and funded, the amounts provided fell short of sustaining the effort he envisioned.
One day, as Hölldobler grew more serious about leaving, we decided to write a book recounting everything we knew about ants. And while we were at it, we asked ourselves, why not try for a book that has everything everybody ever knew about ants, throughout history? Such a project would take a great deal of effort and time, and it might fall short of the goal we set. But what a worthy conceit! Try for the impossible, as Floyd Patterson, the undersized heavyweight boxing champion of the world once said, in order to accomplish the unusual. The result was The Ants, published by Harvard University Press in 1990. It contained 732 double-columned pages, hundreds of textbook figures and color plates, and a bibliography of 3,000 entries. It weighed 7.5 pounds, fulfillin
g my criterion of a magnum opus—a book which when dropped from a three-story building is big enough to kill a man.
On Tuesday afternoon, the following April 9, the faculty and deans of the College of Arts and Sciences gathered for their monthly meeting in the portrait-encircled main room of University Hall. Just as the meeting was about to be called to order, a secretary entered and handed President Derek Bok a message. Bok announced its content: The Ants had been awarded the 1991 Pulitzer Prize in General Nonfiction. I stood and basked in the applause of the Harvard faculty. Bless my soul, the Harvard faculty. Where could I go from here but down?
I later learned that our book was only the fifth on science ever to receive a Pulitzer Prize, and it was the first with a primarily scientific content, written by specialists for fellow professionals. Soon after I left University Hall that day I called Bert and asked him how it felt to win America’s most famous literary award. Note, I reminded him, not scientific, literary. Wonderful, he replied. They would celebrate in Würzburg. The accent was still there, of course. It made the occasion more special.
*W. H. Bossert and E. O. Wilson, “The Analysis of Olfactory Communication among Animals” Journal of Theoretical Biology 5 (1963):443–69; E. O. Wilson and W. H. Bossert, “Chemical Communication among Animals,” Recent Progress in Hormone Research 19 (1963):673–716.
chapter sixteen
ATTAINING SOCIOBIOLOGY
ON AUGUST 1, 1977, SOCIOBIOLOGY WAS ON THE COVER OF Time. On November 22 I received the National Medal of Science from President Carter for my contributions to the new discipline. Two months later, at the annual meeting of the American Association for the Advancement of Science, held in Washington, demonstrators seized the stage as I was about to give a lecture, dumped a pitcher of ice water on my head, and chanted, “Wilson, you’re all wet!” The ice-water episode may be the only occasion in recent American history on which a scientist was physically attacked, however mildly, simply for the expression of an idea. How could an entomologist with a penchant for solitude provoke a tumult of this proportion? Let me explain.
My interest in sociobiology was not the product of a revolutionary’s dream. It began innocently as a specialized zoology project one January morning in 1956 when I visited Cayo Santiago, a small island off the east coast of Puerto Rico, to look at monkeys. I was accompanied by Stuart Altmann, who had just signed up as my first graduate student. Stuart was an academic anomaly, so much so that upon admittance to the Ph.D. program at Harvard the previous fall, he at first found himself without a sponsor. His problem was not his abilities, which were outstanding, but his proposed thesis research, which was too unusual. He had set his sight on the social behavior of free-living rhesus macaques, and particularly those maintained by the National Institutes of Health on Cayo Santiago, newly dubbed “monkey island.” He came well prepared. He had recently worked on howler monkeys in the Panama rain forest. His command of the relevant literature was complete.
Unfortunately, no one at Harvard knew what he was talking about. The behavior of primates under natural conditions remained virtually unknown in 1955. C. Ray Carpenter, an American psychologist, had laid a foundation in the 1930s with field observations of howlers, rhesus, and gibbons. His published work was well respected by a small circle of biologists and anthropologists but had not spawned a school of research. It was not easy to journey to where wild primates live. Jane Goodall still lived in England, her first visit to the chimpanzees of the Gombe Reserve four years away. Several Japanese researchers, at the time Altmann began his own work, were observing macaques on Mount Takasoki, on the island of Kyushu, but they published reports in their native language, which was virtually unknown to American and European scientists.
No senior members of the Harvard biology faculty considered primate field studies to lie within their province. Some doubted that the subject even belonged in biology. Thus Stuart came to me. In the late fall of 1955 I had been offered an assistant professorship in biology, effective July 1 of the following year. Frank Carpenter, chairman of the department, asked me whether, given my interest in the social behavior of ants, I would sponsor Stuart even before my faculty term began. I accepted happily. I was hardly more than a graduate student myself, just a year older than Altmann, eager to learn the strange new subject he had chosen.
I had decided wisely. The two days Stuart and I lived among the rhesus monkeys of Cayo Santiago were a stunning revelation and an intellectual turning point. When I first stepped ashore I knew almost nothing about macaque societies. I had read Ray Carpenter but was unprepared for the spectacle unfolding. As Altmann guided me on walking tours through the rhesus troops, I was riveted by the sophisticated and often brutal world of dominance orders, alliances, kinship bonds, territorial disputes, threats and displays, and unnerving intrigues. I learned how to read the rank of a male from the way he walked, how to gauge magnitudes of fear, submission, and hostility from facial expression and body posture.
Altmann issued a warning: “Two things. Don’t move too suddenly near an infant, as though you mean to harm it. You might be attacked by a male. And if you do happen to be threatened, don’t look the male in the face. A stare is a threat and might provoke an attack. Just hang your head down and look away.” Sure enough, in a careless moment on the second day, I twisted my body around suddenly while standing next to a very young monkey, and it let out a shriek. At once the number two male ran up to me and gave me a hard stare, with his mouth gaping—the rhesus elevated-threat expression. I froze, genuinely afraid. Before Cayo Santiago I had thought of macaques as harmless little monkeys. This individual, with his tensed, massive body rearing up before me, looked for the moment like a small gorilla. I needed no reminder. I lowered my head and looked away in my most studiously contrite manner, frantically signaling the message “Sorry, didn’t mean anything, sorry.” After a few minutes my challenger left.
In the evenings Altmann talked primates and I talked ants, and we came to muse over the possibility of a synthesis of all the available information on social animals. A general theory, we agreed, might take form under the name of sociobiology. Stuart was already using that word to describe his studies; he had picked it up from the Section on Animal Behavior and Sociobiology, a working subgroup of the Ecological Society of America. A belief floated among zoologists even then that animal societies require a different kind of analysis, that they are properly the subject of a separate, minor discipline. But none could say what the general principles of this sociobiology might be, or how they would relate to the rest of biology. Under the guidance of senior zoologists such as Warder Clyde Allee, Alfred Emerson, and John P. Scott, sociobiology was taking form as a discipline but still consisted largely of descriptions of different kinds of social behavior. As Altmann and I talked over the subject during the pleasant Puerto Rico evenings, we could do no better. Primate troops and social insect colonies seemed to have almost nothing in common. Rhesus monkeys are organized strongly by dominance orders based on individual recognition. That much is also true of primitively social wasps but not of the rest of the social insects, whose colonies are composed of hundreds or thousands of anonymous and short-lived siblings living in harmony. Primates communicate by voice and visual gesture, social insects by chemical secretions. Primates fill temporary roles based on personal relationships; social insects have castes and a relatively rigid, lifelong division of labor.
We knew that no science worthy of the name is built wholly from a checklist of similarities and differences with an overlay of phenomena such as dominance and group action. In 1956 there existed no theory to explain diversity—why various traits have arisen in some groups and not others. Altmann had one good idea. He intended to devise probability transition matrices of behavioral acts, to provide a compendium of the following kinds of information: if a rhesus performs act a, then there is a certain probability that it will perform a again, another probability it will perform b, and so on. I agreed on the concept: a great deal of behavior and social interact
ion can be packed into transition matrices. It should then be possible to use the numbers to compare one kind of society more precisely with another. To quantify social interactions is an important step, but where would it lead? The result would still be a description, offering no explanation of how or why a particular species of monkey or ant arrived at one pattern in the course of its evolution as opposed to another. Neither Altmann nor I had the conceptual tools in 1956 to advance sociobiology further, and we let the subject rest. Stuart pressed on to complete his thesis research.
A congenital synthesizer, I held on to the dream of a unifying theory. By the early 1960s I began to see promise in population biology as a possible foundation discipline for sociobiology. I had entered population biology not to serve sociobiology but to help fashion a counterweight to molecular biology. I believed that populations follow at least some laws different from those operating at the molecular level, laws that cannot be constructed by any logical progression upward from molecular biology. This view of the biological sciences motivated me to collaborate with Lawrence Slobodkin, an alliance that later led to the development of the theory of island biogeography with Robert MacArthur.
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