The Dancing Bees: Karl von Frisch and the Discovery of the Honeybee Language

The Dancing Bees

The Dancing Bees

Karl von Frischand the Discovery of the Honeybee Language

Tania Munz

The University of Chicago Press

CHICAGO AND LONDON

TANIA MUNZ is the vice president for research and scholarship at the Linda Hall Library in Kansas City. Previously, she was a lecturer at Northwestern University and a research scholar at the Max Planck Institute for the History of Science in Berlin.

The University of Chicago Press, Chicago 60637

The University of Chicago Press, Ltd., London

© 2016 by The University of Chicago

All rights reserved. Published 2016.

Printed in the United States of America

25 24 23 22 21 20 19 18 17 16    1 2 3 4 5

ISBN-13: 978-0-226-02086-0 (cloth)

ISBN-13: 978-0-226-02105-8 (e-book)

DOI: 10.7208/chicago/9780226021058.001.0001

Frontispiece: see fig. 8.2, p. 219, for more information.

Library of Congress Cataloging-in-Publication Data

Names: Munz, Tania, 1973– author.

Title: The dancing bees : Karl von Frisch and the discovery of the honeybee language / Tania Munz.

Description: Chicago ; London : The University of Chicago Press, 2016. | Includes bibliographical references and index.

Identifiers: LCCN 2015036905 | ISBN 9780226020860 (cloth : alk. paper) | ISBN 9780226021058 (e-book)

Subjects: LCSH: Frisch, Karl von, 1886–1982. | Bees—Behavior.

Classification: LCC QL31.F7 M959 2016 | DDC 595.79 / 9—dc23 LC record available at http://lccn.loc.gov/2015036905

This paper meets the requirements of ANSI / NISO Z39.48-1992 (Permanence of Paper).

To my parents.

And for Tim.

The life of the bees is a magic well. The more one draws from it, the more richly it flows.

KARL VON FRISCH, Aus dem Leben der Bienen (1953)

Contents

INTRODUCTION

Sensational Findings

BEE VIGNETTE I

Victorian Bees

CHAPTER ONE

Coming of Age in Vienna

CHAPTER TWO

The Bees That Could

BEE VIGNETTE II

Sensing the Senses

CHAPTER THREE

Calm before the Storm

CHAPTER FOUR

In the Service of the Reich

BEE VIGNETTE III

Deep inside the Hive

CHAPTER FIVE

State of Grace

CHAPTER SIX

Picking Up the Pieces in Postwar Germany

CHAPTER SEVEN

Coming to America

BEE VIGNETTE IV

Seeing Bees

CHAPTER EIGHT

Attack on the Dance Language

CONCLUSION

180/60

Acknowledgments

Notes

Bibliography

Index

INTRODUCTION

Sensational Findings

In January of 1946, while much of Europe lay buried under the rubble of World War II, Karl von Frisch penned an excited letter to his friend and fellow animal behaviorist Otto Koehler. In the letter, the Austrian-born experimental physiologist and bee researcher reported his sensational findings about the language of bees. Over the previous two summers, he had discovered that honeybees communicate to their hive mates the distance and direction of food sources by means of the dances they perform after returning from foraging flights. He found that the insects indicate nearby food sources via a circular dance and faraway foods via a figure-eight-shaped waggle dance. The straight-run portion of the waggle dance, he explained, also contains information about direction, and the frequency of its turns correlates closely with distance: the closer the supply, the more rapidly the bees dance. The otherwise reserved von Frisch concluded his missive, And if you now think I’m crazy, you’d be wrong. But I could certainly understand it.¹

Von Frisch was right—his discovery was a sensation, and news of it quickly spread throughout Europe and abroad. Although his interpretation of bee communication would at times face fierce opposition, the work endured as a classic example of animal behavior and succeeded in placing bees alongside primates, dolphins, and birds in the mid-twentieth-century pantheon of communicating beasts. The postwar period saw a flurry of activity: interdisciplinary study groups, edited volumes, and conferences all aimed to tackle the problem of animal communication.² Honeybees held a prominent place in this research. During the 1960s, their language was the most widely studied form of animal communication, and some deemed it the most complex, second only to human speech.³ The stakes were especially high, as language had long been considered a window into human minds and souls, and one of the key distinctions between humans and animals.

Von Frisch’s findings and approach were praised by many of the period’s best-known scientists, including the sociobiologist and ant specialist E. O. Wilson, the cognitive ethologist Donald Griffin, and the ethologist and Nobel laureate Nikolaas Tinbergen. Generations of bee researchers would go on to discuss and cite his work. And in 1973, he was awarded one of the highest honors bestowed on scientists—his findings earned him a share of the Nobel Prize for Physiology or Medicine, together with Tinbergen and fellow Austrian Konrad Lorenz.⁴

Even before his receipt of the Nobel Prize, von Frisch’s reputation had already reached far and wide. Throughout much of his career, he wrote books and articles for lay readers and edited a series of small books under the title Understandable Science that specifically aimed to bring the work of well-known scientists to the German-speaking public. He gave talks to audiences that ranged from school-age children to retirees and produced a number of pioneering films that used ingenious methods to show the sensory world of animals.⁵ With his boyish build and thick wire-rimmed spectacles, he introduced young and old to the mysterious world of bees.

The Dancing Bees is a dual biography—on the one hand, of von Frisch as one of the most innovative and successful scientists of the twentieth century and, on the other hand, of his honeybees as experimental and, especially, communicating animals that play a rich role in human culture. It tells of von Frisch’s discovery of the dance language in the context of the politics and events that changed Germany and the world over the course of the twentieth century. Others before him, going as far back as Aristotle, had observed the animals’ dances, and beekeepers had long speculated about the possibility of some form of bee communication to mobilize foragers to exploit food sources. But von Frisch was the first to definitively link the bees’ dances to their recruitment abilities. How, we wonder, did von Frisch manage to observe such behaviors where others for centuries had looked but failed to see?

Alongside von Frisch’s story, I include a series of vignettes that explore aspects of the bees’ lives that illustrate the transition the animals underwent in the popular and scientific imaginations over the course of the century. Long before von Frisch revealed the insects’ remarkable dance language, humans and bees had shared an entwined and complex history. From Virgil’s poetry of the first century BC to Bernard Mandeville’s eighteenth-century, subtly subversive fable of the insects’ politics, the hive served as inspiration for how a well-run social polity might function and proved a ready canvas for moral projections onto nature.⁶ I write the bees’ history alongside human history to reveal the changing cultural and scientific understandings of the insects as they transitioned from political to physiological and, finally, to communicating animals.

And yet while students across the world continue to learn about the honeybee dance language, less well known today are the circumstances under which von Frisch’s discoveries were made. As the date of the letter reporting the sensational findings—January 1946—suggests, he performed some of his most important work against the backdrop of the deadliest conflict in human history.

When Hitler took power in 1933, von Frisch was professor of zoology at the University of Munich. Soon after, the city and its institutions were brought under Nazi control through the process of Gleichschaltung (coordination).⁷ In an effort to purge government institutions of Jews and other undesirables, the Nazi government required all civil servants, including professors, to furnish proof of their Aryan descent. In the case of von Frisch, what had begun as vague rumors about him crystallized by late 1940 into ominous fact: after months of searching, the Nazi office responsible for genealogical research located evidence that his maternal grandmother had been of Jewish descent. Like many nineteenth-century assimilated Jews who considered themselves loyal citizens of the Kaiserreich, her parents had converted to Catholicism just a few years before she was born.⁸ Presumably the young couple had done so in the hope of securing a better future for their famly in a Christian-dominated society. But the Nazi concept of what it meant to be Jewish was overwhelmingly rooted in notions of blood and descent rather than cultural heritage or religious belief. As a consequence, people like von Frisch, who had never before considered themselves Jewish, were now officially declared such based on the more or less distant tendrils of their ancestry.

The fate of the Quarter Jews was never as dire as that of the Half and Full Jews.⁹ But gradually they were excluded from higher education and certain professions, such as medicine and law. In the case of von Frisch, the declaration of his status as Quarter Jew by the president of the University of Munich in early 1941 also announced that he was to be ousted from his position at the university. Von Frisch was devastated. He was only fifty-four years old and lived for his work.

Prior to the war and indeed throughout most of his life, he had considered himself staunchly apolitical. Nonetheless, his instincts about whom to turn to and how best to position his work in the scientific landscape of the times would serve him well. Through an odd twist of fate, he performed some of his most important work not so much despite the regime as because of it. And the honeybees played a crucial role.

At the very moment that von Frisch received news of his impending ouster, his favorite research subjects—the honeybees—were also in trouble. A small intestinal parasite, known by its Latin name Nosema apis (or Nosema, for short), was wreaking havoc on the insects’ insides. The invaders multiplied so prolifically that they quite literally burst the animals’ guts. Afflicted bees crawled to their deaths leaving behind a trail of feces. During 1940 and 1941, hundreds of thousands of hives across Germany were destroyed by the parasite.¹⁰ Although the causes of the sudden outbreak were unknown, it was clear that the loss of agriculture’s main pollinators would spell disaster for humans, and especially for a weary German nation at war.

By winter of 1941, the stunning results of the blitzkrieg were giving way to a grinding war of attrition. Now, it seemed, the war could not be won by brief displays of force but by the nation that could keep its men armed and fed the longest. Germany redoubled its efforts. From military strategy to the economy and from agricultural policy to propaganda, all resources bled into the war machine. And as the German nation bore down for total war, von Frisch would gain a small foothold in the Nazi bureaucracy. The hungry nation at war depended critically on agriculture to feed bodies in battle and mouths on the home front.¹¹ Agriculture depended on the successful cultivation and growth of food crops, and many of these crops relied on honeybees; only blooms pollinated by the insects would bear fruit. Von Frisch was classically trained in experimental physiology and knew precious little about intestinal parasites and even less about large-scale agriculture. But he knew an awful lot about bees.

When I first read von Frisch’s work, I was drawn in by the elegant simplicity and compelling logic of his experiments. He seemed able to distill the potentially overwhelming complexity of the outdoors into something that more closely resembled the clean and orderly world of the laboratory. And yet the more I read in his papers, letters, and books, the more I found myself wanting to know about the messy world that he seemed to keep at bay when working with his bees. As I followed him on his path, a far less tidy picture emerged; his work, it turned out, was very much of its time and place. Shortly after he received news of his impending ouster, he appealed to friends and colleagues for help. He also switched the focus of his research to a more practical orientation in the hope that the Nazi authorities would deem his work worth continuing. What, I wondered, are we to make of the fact that he performed his most important research while receiving funding from the Nazi Ministry of Food and Agriculture? How should such choices be evaluated in light of what we know today about Nazism and its relationship to science? And what were the postwar consequences of his having been declared a Quarter Jew by the Nazis?

Over the years that I immersed myself in von Frisch’s life and work, I learned that his story resists simple answers. Instead, it offers us a rare glimpse into how he navigated the overwhelming and often frightening Nazi bureaucracy. In considering von Frisch’s biography, we are forced to reject simplistic accounts that paint a regime hostile to all but the crudest and most racialized sciences. His work was neither racially motivated nor pseudoscientific (like much of Nazi genetics, anthropology, psychology, medicine, and physics). And yet the research fit in well with Germany’s long-held aspirations of gaining independence from the import of foods and raw materials to prepare for and wage war.¹²

After the war, von Frisch once again reached out to his colleagues across the Atlantic and Channel. His wartime fate played no small part in easing this contact. Scientists who had worked in the Reich during the war often were excluded or given a chilly reception by their colleagues and old friends in the former Allied countries. But because it became known that the Nazis had declared von Frisch one-quarter Jewish and threatened his job, his political credentials were soon considered beyond reproach; a perceived enemy of the Nazis could easily be considered a friend of the Allies.¹³ To Americans who were looking for good Germans in the wake of the war, especially as their attentions in the late 1940s shifted from denazification to a cold war with the Soviet Union that was heating up, this political exoneration made him an attractive candidate for support and funding. On a 1949 lecture tour to the United States, von Frisch reestablished important connections with access to the deep pockets of the Rockefeller Foundation. These transatlantic allegiances would also support him during heated debates about the meaning of the bee dances in the 1960s and 1970s.

At issue in the dispute that started in the 1960s was not merely whether the bees’ dances served to communicate information about the location of foods but rather the very approach to animals. Were nonhuman animals capable of symbolic communication? And if so, what did this mean for a scientific understanding of the animal-human boundary?

Von Frisch’s approach and answers to these questions ultimately withstood the challenge, and the receipt of the Nobel Prize was a testament to a science of animals that had come into its own. Animal behavior studies had successfully shed its nineteenth-century reputation of lacking objectivity and reveling in anecdotes about morally upstanding worker bees, wise queens, and lazy drones. And yet in their place emerged a new kind of model for humanity. When the Rockefeller Foundation once again sponsored von Frisch’s work in the postwar period, it did so with the explicit desire to understand communication across peoples and cultures in the wake of the war that went down in history as one of humanity’s greatest disasters.

Even today, as science has advanced to give us state-of-the-art access to the hive and its inhabitants, bees still occupy an important cultural and political space. As we must once again face the sobering prospects of the dying bees, the hive serves as a mirror that reflects back to us the values, fears, and aspirations of our time. Colony collapse disorder brings us yet again face-to-face with both the limits of our scientific reach and our dependence on these creatures for our food supply. What is causing the bees to die?¹⁴ We do not know exactly. But we assume that their catastrophically increased death rates are somehow related to our modern ways of life—possible contenders in the search for explanations range from the stresses associated with monocultural factory farming and pesticide exposure, to overuse of antibiotics, to the introduction from Asia of the parasitic Varroa destructor mite. Thus, despite our ongoing efforts to purge modern science of the anthropomorphic pull of previous generations, bees continue to challenge our very understandings of the human us and animal them.

BEE VIGNETTE I

Victorian Bees

In the darkness of the hive, tens of thousands of bees go about their business. Some tend the young while others clean cells. Still others repair and build combs while a few bees guard the entrance from intruders. Another set meets foragers as they return from their nectar-gathering flights. These insects receive their sisters’ regurgitated food and move it into storage cells. Other gatherers walk deep into the hive with hind legs laden with pollen and kick off the bounty of their forage into dedicated cells. A handful of bees face the queen as she moves through the dense tangle of bodies and dips her abdomen into cell after cell. Each time, she deposits a single tiny egg—glossy and translucent, like a perfect grain of rice. After three days, the eggs quiver and give way to minute, wormlike larvae. They lie curled in their cells and eat. And eat. Nine days later, the larvae weigh more than a thousand times what the original egg weighed. Worker bees now seal the larvae’s cells. The enclosed creatures enter the pupal stage, in which they neither eat nor drink. On the twenty-first day, the transformation is complete and freshly formed worker bees emerge. Such are the workings of the hive and the stuff of bee lore over the centuries.

Bees have long captured the attention of not just beekeepers and naturalists but also economists, poets, political scientists, and social theorists. The animals have been admired for their work ethic, statecraft, and gifts of honey and wax. But above all else, commentators have wondered about the little animals’ remarkable abilities to manage their complex social world. If no single animal—not even the queen—can oversee the whole, then how does the hive arrive at decisions that ensure its survival? For naturalists, bees came to stand at the very pinnacle of instincts as animals whose every move was determined by complex inborn programs of behavior.

If naturalists crowned the bees supreme mistresses of instinct, they appreciated the insects’ comb building as an especially impressive example in their repertoire of accomplishments. So smitten was Charles Darwin with the animals’ elegant combs that he declared him a dull man who can examine the exquisite structure of a comb, so beautifully adapted to its end, without enthusiastic admiration. Dull, indeed. Stacked row upon row, in perfect, machinelike fashion, the cells were deemed marvels of geometric efficiency. The shapes nested in such a way as to distribute their weight equally between neighboring cells. Each hexagon shares its walls with adjacent cells, thereby requiring the least amount of wax for its construction. The eighteenth-century mathematician Colin Maclaurin wrote of the cells’ shape: What is most beautiful and regular, is also found to be most useful and excellent.¹

And yet while all could agree that the combs were elegant and excellent, opinions diverged over what might be required of the bees to execute these constructions. To many, the combs’ seemingly high degree of sophistication demanded an explanation beyond the merely natural; indeed, they believed them to evidence God’s divine plan in nature. The early modern philosopher Thomas Reid argued that the bees work most geometrically, without any knowledge of geometry; somewhat like a child, who, by turning the handle of an organ, makes good music, without any knowledge of music. He concluded: The geometry is not in the Bee, but in the great Geometrician who made the Bee, and made all things in number, weight, and measure.²

Darwin is widely hailed as having put an end to such appeals to God to explain structure and function in nature. But how, if not by the hand of God, had animals and plants become so exquisitely adapted to their surroundings? According to Darwin, instincts, like physical structures, varied and were heritable and therefore subject to natural selection. Since some instincts served organisms better than others, those animals that possessed them would presumably live longer and leave more offspring than their less fortunate kin, and the traits that had offered them an edge in survival and propagation would be passed on to their sons and daughters. Darwin knew well that the success or failure of his theory of evolution depended critically on its ability to explain such complex behaviors as comb building in bees. And indeed, he dedicated considerable space and energy in his Origin of Species to describing just how the comb-building instinct might have arisen from more primitive bees that bored holes into wax.³

Yet even among those who cheered Darwin’s efforts to naturalize explanations of the living world, some observers asked whether there might not be something more—perhaps closer to judgment—that guided the animals’ mysterious behaviors? By the late nineteenth century, at least one naturalist was willing to put a name to this something—intelligence. The English comparative psychologist George John Romanes in his 1882 book, Animal Intelligence, surveyed the animal kingdom from mollusk to primate to marshal examples in support of the book’s title. In a chapter dedicated to bees and wasps, he told of many wondrous feats that he considered too spectacular to be left to mere instincts. Indeed, he argued that these examples proved the existence of the very kinds of foresight and insight that characterize intelligence in humans.

He too considered the honeybees’ combs the most astonishing products of instinct that are presented in the animal kingdom. But, he continued, it was an instinct not wholly of a blind or mechanical kind but [one that] is constantly under the control of intelligent purpose. In support of this claim, he cited a range of observers who had witnessed the animals making subtle adjustments to their comb building with seemingly humanlike intelligence. He cited one well-known apiculturist who had witnessed some bees tearing down and then correcting sections of comb that their sisters had built, because they found the structures wanting. In another instance, after a piece of comb had fallen to the ground, the bees not only repaired the missing piece but also reinforced other sections of the comb to prevent future calamities. To Romanes, such adjustments evidenced foresight and insight that contradicted the blind execution of instinct that others had postulated.⁴

Examples of a less savory kind also impressed Victorian onlookers as all too human and therefore indicative of faculties beyond mere animal instinct. With regret, they recounted lurid examples of bees engaged in warfare, theft, and murder. Not surprisingly, such descriptions of the animals’ darker side were often saturated with moral opprobrium. When bees with the thieving propensity were observed entering a foreign hive, their intentions could be clearly read. According to Romanes, They show by their whole behavior—creeping into the hive with careful vigilance—that they are perfectly conscious of their bad conduct; whereas the workers belonging to the hive fly in quickly and openly, and in full consciousness of their right. The behavior soon spreads, and the whole bee-nation may develop marauding habits, and when they do this they act in concert to rob by force.⁵

If such accounts called into question the bees’ moral fiber, descriptions of their emotional makeup offered even less comfort. Sir John Lubbock, one of the nineteenth century’s foremost experts on insects, painted a thoroughly grim picture: Far, indeed, from having been able to discover any evidence of affection among them, they appear to be thoroughly callous and utterly indifferent to one another. On one occasion, he reported having been forced to kill a bee for experimental purposes. He crushed the unfortunate animal right next to one of her hive mates. So close were the animals that their wings touched, and yet the survivor took no notice whatever of the death of her sister, but went on feeding with every appearance of composure and enjoyment, just as if nothing had happened.⁶

Despite such chilling accounts of the insects’ indifference toward their hive mates, the animals nevertheless seemed to display remarkable levels of cooperation. Indeed, it was precisely this ability to settle the complex affairs of the hive communally that prompted commentators to wonder about their ability to communicate. For example, observers offered vivid accounts of the collective reaction to the loss of a queen. Since a queenless hive is as good as a dead hive, the news of the loss soon pitches the hive into a frenzy. The German bee researcher Ludwig Büchner reported how in a little time . . . the sad event will be noticed by a small part of the community, and these will stop working and run hastily about over the comb. Soon, the excited bees . . . leave the little circle in which they at first revolved, and when they meet their comrades they cross their antennae and lightly touch the others with them. The bees which have received some impression from this touch now become uneasy in their turn, and convey their uneasiness and distress in the same way to the other parts of the dwelling.⁷ Thus, according to Büchner, the busy bees disseminate the bad tidings with the swiftness of village gossips.

But not everyone agreed that the bees communicate. The utterly sensible Sir John Lubbock, for example, reported to the British journal Nature that his experiments had yielded no evidence of the phenomenon. He had offered food to one of his bees in a location that lay out of sight of the hive. While feeding the animal, he had marked her to see whether she would return and recruit her hive mates to the hidden location. He waited, but to no avail—though the marked bee appeared repeatedly at the food source, her sisters never showed up.⁸

More typical, however, was the view of an American by the name of Josiah Emery who in a letter to Nature countered Lubbock’s skepticism. He recounted the methods of American bee hunters as evidence of the bees’ abilities to communicate. A hunter in search of wild honey, he explained, would first capture a single foraging bee and place her in a box with honey. After allowing the bee to gorge herself, he would release her and wait until she reappeared with some of her hive mates. The hunter would now catch these animals as well and place them in the box so that they too could feed on the honey. Now he could move around the area and release the insects one by one, each time paying close attention to the direction in which they flew. As the animals returned in beeline to their hive, the hunter could determine the location of the bees’ home by triangulation. The letter writer urged that when the bee hunters released their first-caught bee and waited, they relied precisely on the bees’ ability to communicate with and recruit their hive mates to food stores. It is possible that our American bees are more intelligent than European bees, he quipped to a largely British readership, but hardly probable; and I certainly shall not ask an Englishman to admit it.⁹

Were American bees more intelligent than British bees? Unlikely. But it would be another century until careful research revealed by just how great a margin the sober Sir John Lubbock had underestimated the bees’ abilities to communicate.

CHAPTER ONE

Coming of Age in Vienna

Karl’s father, Anton von Frisch, was a nineteen-year-old medical student when he first met Marie Exner.