FRISCH, KARL RITTER VON

(b. Vienna, Austria, 20 November 1886; d Vienna, 12 June 1982)

experimental zoology, comparative physiology, ethology.

Karl von Frisch is remembered for his pathfinding studies of the sensory capabilities and behavior of animals, particularly fish and honeybees. Throughingenious experiments of classical simplicity, informed by an evolutionary biologist’s sense of function and a physiologist’s concern with biological mechanisms, he greatly enlarged the scientific understanding of animal behavior. His remarkable discovery of the dance “language” of the honeybee is generally regarded by ethologists as the single most important contribution to the study of animal behavior of the twentieth century.

Frisch was the youngest of four sons of Anton Ritter von Frisch and Marie Exner von Frisch. Three generations of physicians preceded him on his father’s side of the family. Careers in academia predominated on his mother’s side of the family. His father was a surgeon and urologist who conducted significant research in anatomy and bacteriology, and became professor of surgery at the Vienna General Polyclinic. His paternal grandfather, also named Anton, was a surgeon in the Austrian Imperial Army who received the Order of the Iron Cross, and with it a hereditary knighthood, for his work in reorganizing the Army Medical Corps.

Karl’s mother, a woman of keen intellect, was the daughter of Franz Exner, professor of philosophy at the University of Prague. All four of her brothers became professors. The most important of these for Karl’s development as a scientist was Sigmund Exner, professor of physiology and director of the Physiological Institute at the University of Vienna. Another of Karl’s uncles, Franz Serafin Exner, was professor of physics and director of the Physics Institute, also at the University of Vienna. This scholarly tradition continued among Karl’s brothers; Hans von Frisch became a professor of constitutional law (initially at Basel, eventually at Vienna); Otto von Frisch became assistant professor of surgery and director of the Rudolfinerhaus clinic in Vienna; and Ernst von Frisch became director of the Salzburg Studienbibliothek.

Frisch attended the Benedictine Schottengymnasium in Vienna and then enrolled in 1905 as a medical student at the University of Vienna, following his father’s wish that he become a physician. Zoology was his primary interest, however, and in 1908 he transferred to the Zoological Institute at Munich to study experimental zoology under Richard Hertwig. In 1909 he returned to Vienna to write his doctoral dissertation under Hans Przibram at the Biological Experimentation Institute. He received his doctorate from the University of Vienna in 1910.Frisch went back to Munich in the fall of 1910 as an assistant to Hertwig and became a Privatdozent at Munich in 1912. During World War I he worked at the Rudolfinerhaus, the hospital directed by his brother Otto. He was appointed assistant professor at the University of Munich in 1919, professor of zoology and director of the Zoological Institute at Rostock University of Breslau in 1923. Upon Hertwig’s retirement in 1925, Frisch succeeded him as professor of Zoology succeeded him as professor of zoology and director of the Zoological Institute at the University of Munich.

In the early 1930’s Frisch secured funds from the International Education Board of the Rockefeller Foundation and from the state of Bavaria to replace the old Zoological Institute at Munich with a new one. Thenew institute, completed in 1932, was the most up-to-date zoological institute in Europe. After the coming to power of the Nazis, however. Frisch was unable to choose his own staff freely, and in 1941 he learned that the Ministry of Education was planning to remove him from his professorship because his maternal grandmother, Charlotte Dusensy Exner, had been of Jewish ancestry. The intended dismissal did not come to pass, however, because the economic and political influence of powerful friends, as reported by Frisch in his autobiography, was brought to bear at the Ministry of Food and at Nazi party headquarters. Frisch’s expertise with bees stood him in good stead at the Ministry of Food, which valued him because an epidemic was devastating the bee populations of Europe and causing serious damage to German agriculture. The ministry gave Frisch an official assignment to study what could be done to combat the problem.

In July 1944 Frisch’s house in Munich was destroyed and the Zoological Institute was severely damaged by Allied bombing. Frisch had already moved much of his equipment to his summer home in Brunnwinkl, Austria, and he continued his research there. In 1946 he accepted the professorship of zoology and directorship of the Zoological Institute at the University of Graz (Austria). In 1950, however, he returned to Munich, where the Zoological Institute had been largely rebuilt. He remained at Munich as professor of zoology and director of the Zoological Institute until his retirement in 1958.

Frisch married Married Margarethe Mohr on 20 July 1917. They had three daughters and one son. In the course of his long and distinguished career, Frisch received many honors, including the Nobel Prize for Medicine or Physiology, which he shared with Konrad Lorenz and Nikolaas Tinbergen in 1973. He was elected a foreign member of the Royal Society of London and belonged to numerous other scientific academies in Europe and the United States. He received honorary degrees from six universities. His other prizes included the Order of Merit for Sciences and Arts (1952), the Kalinga Prize for the popularization of science (1959), the Austrian Medal of Honor for Science and Art (1960), the Balzan Prize for Biology (1963), and the Distinguished Service Cross with Star and Ribbon of the Order of Merit of the German Federal Republic (1974). He was President of the German Zoological Society from 1928 to 1929.

Early Scientific Work. Like most of the main contributors to the biological study of animal behavior in the first half of the twentieth century. Frisch was exceptionally interested in animals as a child. His mother, who possessed a strong love of nature, permitted him to maintain a sizable menagerie, which in his secondary school days included, by his later count, nine different species of mammals, sixteen species of birds, twenty-six species of coldblooded terrestrial vertebrates, twenty-seven species of fish, and forty-five species of invertebrates. He also took great pleasure in collecting animals at his family’s summer home at Brunnwinkl, on Lake Wolfgang in the Salzkammergut region of Austria.

In Frisch’s three years of medical studies at Vienna, the teacher who influenced him most was his uncle Sigmund Exner. Exner, a pioneer in the study of comparative physiology, impressed upon his nephew the importance not only of designing careful experiments but also of being cautious in drawing conclusions. Under his uncle’s direction, Frisch investigated how the position of the pigment cells in the compound eyes of butterflies, beetles, and shrimp shifted in response to light and darkness. These investigations were the subject of his first major scientific paper, which appeared in Biologisches Zentralblatt in 1908.

Experimental zoology was the forte of Richard Hertwig, under whom Frisch studied at Munich. Hertwig stressed that zoology had to go beyond the mere description of forms to an understanding of causal relationships. Such an understanding was to be achieved primarily through experimentation. Frisch also benefited greatly from studying with Richard Goldschmidt, who was Hertwig’s chief assistant, and Franz Doflein, who, as assistant professor of the systematics and biology of animals, led Frisch and his fellow students on numerous excursions into the field.

When Frisch returned to Vienna to write his doctoral dissertation under Hans Przibram at the Biological Experimentation Institute, Przibram selected a morphological topic for him. Frisch chose instead to study nerve pathways and color changes in fish, a topic in which he had become interested after observing the ability of minnows to adapt their coloration to light or dark backgrounds. His Habilitationsschrift at Munich was also on color adaptation in fish. Published in 1912, it included the results of research he had carried out in the spring of 1911 at the Naples Zoological Station.

Crucial for the direction of Frisch’s researches as a young scientist was the stimulus he received from the work of Carl von Hess, director of the Munich Eye Clinic. A prolific researcher, Hess studied phototactic reactions in a wide range of animals. He found that fish and invertebrates appeared to perceive the brightness values of the colors of the spectrum in the same way as a totally colorblind human did: colors in the gold-green to green region appeared brightest, and the spectrum was shortened at the red end. On the basis of this correspondence, Hess concluded that fish and invertebrates were totally color-blind.

Having studied the ability of fish to adapt themselves to the color of their surroundings. Frisch could not believe that they were unable to perceive colors. In June 1911, at the annual meeting of the German Zoological Society, held in Basel, he delivered a lecture entitled “On the Color Sense of Fish,” in which he challenged Hess’s thesis. The fact that many fish possess an ability to change color and many of them assume splendid colors at spawning time made it difficult for Frisch to believe that fish are color-blind, but he wished to judge the issue primarily on experimental grounds. His experiments showed that fish can adapt not only their lightness or darkness but also their color to their surroundings. He was able to show further that fish did not respond simply to the brightness of colors but also to the colors themselves. The gold and red pigment cells expanded in response to gold or red backgrounds but not to a gray background of equal brightness. That this response was mediated by the fish’s eyes was shown by the fact that fish that had been blinded did not display color changes. For Frisch these results strongly confirmed the idea that fish possess a color sense.

Hess responded by calling Frisch’s results “completely mistaken,” and an acrimonious dispute ensued between the two that continued for more than a decade. Though the dispute began over the color sense of fish, it soon opened up on a second front; the question of the color sense of bees.

Frisch’s dispute with Hess over the color sense of bees was not without precedent. As early as 1793, in a book entitled Das entdeckte Geheimniss der Natur im Bau und in der Befruchtung der Blumen, Christian Konrad Sprengel proposed that the bright colors and special “honey guides” of flowers were contrivances designed to bring about the fertilization of flowers by insects. Charles Darwin supported Sprengel’s idea that the colors of flowers served to attract insect pollinators, but unlike Sprengel he explained this in terms of natural selection rather than design on the part of a wise Creator. Darwin was more interested in the special mechanisms that assured cross-fertilization in plants than he was in how insects are attracted to flowers in the first place, and he did not test whether bees are capable of distinguishing colors.

Darwin’s contemporary Sir John Lubbock did precisely that. He succeeded in training bees to associate food with certain colors. His experiments led him to conclude not only that bees can distinguish colors but also that they have color preferences. Lubbock’s work was challenged by the French biologist Félix Plateau, who maintained, on the basis of his own experiments, that insects are guided to flowers not by colors but by the smell of nectar or pollen. Between 1900 and 1910 the issue was taken up by scientists in Germany (Albrecht Bethe and Hugo von Buttel-Reepen), Switzerland (AugusteHenri Forel), and the United States (Charles Henry Turner).

Frisch’s debate with Hess over the color sense of bees led him to the first of his classical experiments on honeybee behavior. Beginning in the summer of 1912, he arranged on a feeding table a series of gray squares representing a gradation in brightness from white to black. Among the gray squares he placed a blue square of equal size. He then placed a watch glass on each square and filled the watch glass on the blue square with sugar water. The bees were allowed to feed on the sugar water. New, clean squares and watch glasses were then substituted for the old ones, the relative positions of the squares were changed, and the process was repeated. Frisch found that after a training period of one to two days, the bees would congregate on a blue square when it, like the gray squares among which it was placed, had only a clean and empty watch glass on it. This showed, he believed, that bees are able to distinguish a blue square from a gray square of a comparable brightness, and that bees thus have a color sense. That the bees were not attracted to a special smell associated with a blue square was demonstrated by the fact that the bees were still demonstrated by the fact that the bees were still attracted to the blue square when it and the gray squares were all covered with a large glass plate.

In 1914 Frisch published a 188-page monograph, Der Farbensinn und Formensinn der Biene. He reported that while bees could distinguish the color blue from all shades of gray, they confused red with black and blue-green with gray. Similarly, they mistook red-orange for yellow or green, and blue for violet or purple. He concluded that bees have a color sense, but that it is like that of a red-green color-blind man rather than a man with normal vision. He noted, further, that the colors of flowers and the color sense of bees are closely correlated: colors that are seldom seen in flowers are colors that bees cannot distinguish. This supported the idea that the colors of flowers have developed as adaptations to the color sense of their pollinators. In the following decade, investigators using more refined apparatus than Frisch’s soon revised and extended his findings (identifying, among other things, that bees perceive blue-green and ultraviolet as colors), and in the 1950’s and 1960’s Frisch’s student Karl Daumer further extended the study of the bee’s color sense and the biological significance of floral colors.

Frisch also reported on his efforts to train bees to respond to different shapes. He was unsuccessful in his attempts to get them to distinguish between different geometric shapes, and suggested that they could not distinguish between forms that were foreign to them. This interpretation was modified in the 1920’s and 1930’s by the researches of Mathilde Hertz, who found that what seemed to matter most for the form sense in bees was the degree of “brokenness” of the patterns.

Hess’s response to Frisch’s experiments on the color sense of fish and bees was to maintain, without looking at the experiments, that Frisch’s work was flawed or incorrectly reported. Hertwig supported Frisch by attending the experiments and attesting to the accuracy of the reported results. The debate had the positive effect of bringing attention to Frisch’s work, and it stimulated additional studies on color vision in animals by others. In a paper of 1923 reviewing the debate on the color sense in animals over the previous decade and a half. Frisch was able to list not only eight papers of his own and twenty-four of Hess’s but an additional thirtythree publications produced by twenty-eight other authors.

The sensory physiology of bees offered Frisch further topics for study. During a brief leave from his wartime hospital duties, he began using foodtraining techniques to test the honeybee’s sense of smell. In 1919 he published a 238-page monograph on the bee’s sense of smell and its significance for the biology of flowers. Through a wealth of experiments he found not only that bees could be trained to particular odors but also that their olfactory reactions appeared to be very similar to those of humans, in that substances that smelled similar to humans evidently smelled similar to bees, and substances that had no smell for humans had no smell for bees. He conducted experiments that established that the bee’s sense of smell is located on its antennae. He also undertook comparative experiments that showed that although the color of flowers can attract bees from a greater distance, it is the scent that bees use to identify flowers at close range.

First Studies of the “Language” of the Bees. In the course of his studies on the senses of color and smell in bees, Frisch observed that a food source set out in a new location often remained undiscovered by scout bees for hours or even days, but that once one bee discovered the food, there would soon be dozens of bees feeding upon it. He also observed that after a troop of bees that had been trained to feed at a particular spot exhausted the food there, the number of bees at the spot quickly dropped off to zero; but as soon as the food was replenished, and a scout bee discovered the renewed supply and returned to the hive laden with nectar, the whole troop would soon be back at the feeding place. It thus appeared to Frisch that bees somehow communicated to their hive mates the existence of worthwhile food sources.

In the spring of 1919 Frisch set out to discover how this communication took place. Using a queenbreeding cage with a single honeycomb that could be observed through glass on both sides, he was able to witness what he later described in his autobiography as “the most far-reaching observation” of his life. He saw a scout bee return from a freshly replenished feeding station and perform on the honeycomb a lively Werbetanz (courting dance), which excited the bees around her and caused them to fly out to the feeding station. He spent the next three years studying this communication system. He set up an observational hive in which the honeycombs were arranged side by side so they could all be observed through a glass window. He also devised a system of numbering bees by painting them with spots of different colors so they could be individually identified.

Frisch first reported on these researches in a lecture to the Society for Morphology and Physiology at Munich in January 1920. He followed this with two more reports in 1921 and 1922, and in 1923 he published a 186-page monograph entitled “Uuml;ber die “Sprache” der Bienen: Eine tierpsychologische Untersuchung,” It turned out that bee “dances” had been described by observers in the eighteenth and nineteenth centuries. Frisch’s special contribution was his analysis of what the dances signified.

In speaking of a “language” of the bees, Frisch was responding to Buttel-Reepen’s suggestion of 1915 that bees make perceptibly different sounds— a flight tone, a stinging tone, a hunger tone, and so forth—that constitute a kind of language for them. Frisch found no evidence for what ButtelReepen called the Lautsprache (audible language) of the bees. He concluded instead that the “language” bees employed to communicate the existence of food sources involved two special dances. The first of these was a “round dance” in which the forager tripped around in a tiny circle on a cell of the honeycomb, alternately to the left and to the right, continuing this for up to half a minute or longer. Thesecond was a Schwänzeltanz (waggle or tail-wagging dance), in which the returning forager performed half-circle turns, alternately to the left and the right, between which she did a short, straightline run while wagging her abdomen rapidly from side to side.

In each case, neighboring bees trooped excitedly along behind the dancer. Frisch interpreted the round dance as a dance performed by nectar collectors and the tail-wagging dance as a dance performed by pollen collectors. Each served, he believed, to recruit additional bees to go out and collect food of the same sort as that brought back to the hive by the dancer. Early in his researches Frisch thought there was a third dance as well, but his later observations failed to confirm this.

It appeared to Frisch that the means by which new bees were recruited to seek a food supply and the means by which they proceeded to find the supply were two different things. His observations showed that the new recruits did not simply accompany the dancer back to the source. Instead, the recruits flew out from the hive in all directions, searching for the particular flower scent that had clung to the dancer. If the source happened to be a flower that was odorless, the recruits were aided in their quest by the scent of their predecessors, for bees in the presence of an excellent food source will discharge their scent organs, thereby impregnating the air with a special aroma that serves to attract their fellow workers.

Frisch acknowledged at a meeting of the German Society for Natural Science and Medicine at Innsbruck in 1942 that at the beginning of his work he had actually contemplated the possibility that the dancer might use a “secret sign” to indicate the direction and distance of the food. This was not the conclusion to which his initial studies on the “language” of the bees had led him. Nonetheless, what he had found was quite remarkable. He had found that honeybees, through their symbolic dances and the comparatively simple means of touch and smell, were able to communicate to bees back in the hive not only the existence of profitable food sources but also the species of flowers with which the food was associated. In this way they could effectively exploit the food sources in their surroundings as different species of plants came into flower. Not until two decades later did he discover that bees can indeed communicate to their hive mates the direction and the distance of their feeding places. Work done primarily in the 1950’s established further that the dances can serve to communicate information not only about food but also about water, resinous substances, and potential nest sites.

Other Experimental Studies. Frisch’s researches on bees were necessarily restricted tot he warm months of the year, when bees are active. For his laboratory work in the colder months, fish were his favorite, though not exclusive, experimental subjects. He also conducted a study on color changes in salamanders, confirming Paul Kammerer’s report that raising salamanders on backgrounds of different colors affected the animals” coloration. In 1923 Frisch took up the long-controverted question of whether fish, which lack the basic auditory structures of the higher vertebrates, are capable of hearing. Using food-training techniques, he was able to teach a catfish to respond to a whistle.

Frisch proceeded to explore in detail the sensory physiology of fish. Combining histological examinations of the retina of fish with color-training experiments of fish in bright and dim light, he was able in 1925 to provide strong support for the duplicity theory of vertebrate vision. He demonstrated that the rods take over for the cones in the fish retina at just the time that the fish in semidarkness becomes unable to distinguish colors. Later, performing surgical operations on the inner ears of fish, he was able to show that one part of the labyrinth serves for hearing while another part serves the sense of balance. He established that minnows cannot localize sound. Along different lines, he discovered that when a minnow’s skin is punctured, a chemical substance is released that serves as a warning to other minnows. This was a fundamental discovery for subsequent work by others on the role of alarm substances in animal social behavior.

These various studies of Frisch’s, like his work on bees, illustrate the kind of comparative physiological investigations at which he excelled. Most zoologists in his day, as he saw it, were preoccupied with the study and description of form. The pioneers of comparative physiology tended to be physiologists working in medical faculties. Perceiving the need for a journal of comparative physiology, Frisch and Alfred Kühn persuaded the publisher Dr. Ferdinand Springer to start the Zeitschrift für vergleichende Physiologie. Frisch served as an editor of this journal, now called Journal of Comparative Physiology, from its inception in 1924 until his death in 1982.

The “Language” and Orientation of the Bees Revisited. Frisch continued his honeybee studies in the 1920’s and 1930’s, devoting his attention in particular to a long course of investigations on the bee’s sense of taste that laid the foundations for work on the subject thereafter. In the 1940’s two factors led him to take up the question of the bee’s “language” again. One was a doctoral dissertation written at Bonn in 1938 by Christoph Henkel, who maintained that Frisch had been mistaken in supposing that the bee’s round dance signified a nectar source while the waggle dance signified a pollen source. Henkel believed that round dances simply indicated an unnatural abundance of food. The second was the reported successes of Russian scientists in increasing clover production by training bees to clover-flavored sugar water. Frisch’s initial testing of Henkel’s view seemed only to confirm the original distinction between the dances of nectar gatherers and pollen gatherers, but in the course of trying to train bees to visit distant feeding places, he made a discovery that opened up a new set of researches on the dance language and orientation of bees.

The discovery was made in August 1944 at Brunnwinkl, where Frisch found it still possible to work despite the dislocations caused by the war. For the first time in the course of his researches, he set up a feeding station at a distance of more than 100 meters from the hive. Previously he had always set the feeding station close enough to the hive so that both could be watched at the same time. He was surprised to find newly recruited bees not searching for food in ever-widening circles from the hive but instead flying to the immediate proximity of the feeding place. Doubting that the direction of a food source could be communicated by a scout to her hive mates, in September 1944 he tested the more likely explanation that the scout bees were emptying their scent organs in flight, thus providing a trail that the other bees followed. This, however, proved not to be the case, for shellacking the bees” scent organs shut did not keep new recruits from arriving at the feeding station.

Frisch then constructed an observation hive to see whether bees returning to the hive from different distances behaved differently from one another. He set up feeding stations at 12 and 280 meters from the hive, and marked the bees that fed at the two stations with blue and red spots, respectively. He found, to his amazement, that all the bees that had been feeding at the nearer source performed “round” dances when they returned to the hive. While all those that had been feeding at the more distant source performed “waggle” dances. He made these observations on 6 October 1944. In the short time that remained before cold weather called a halt to his researches for the rest of the season, he was able to establish that the round dance merged into the waggle dance at a distance of 50 to 100 meters from the hive. He had not resolved the question. however, of whether bees had a means of communicating direction as well as distance.

Eagerly resuming these studies in June 1945, Frisch observed that the direction of the bees” waggle rund on the honeycomb in the hive changed during the course of the day, though the location of the feeding place remained fixed. He discovered that the dances were oriented with respect to the position of the sun. Thus, for example, a waggle run directly upward on the honeycomb in the dark hive indicated that the food source was in the direction of the sun, while a run 60 degrees to the left of the sun, When bees danced outside the hive on a horizontal surface—as opposed to inside the dark hive on a vertical honeycomb—they pointed directly toward the food source. In other words, the sun appeared at the same angle and on the same side to them as it had on their outward flight to the food.

Continuing his analysis of how bees communicate distance. Frisch increased the distance of the feeding places from the hive to see if the form of the dance would change again. Perceiving a difference in the rhythm of the dances of bees returning from 100 meters away compared with those of bees returning from 100 to 1, 500 meters away from the hive. He found that the number of round dance movements per unit time decreased with distance, while the number of waggle movements in the waggle runs and the duration of the runs increased.

Having established that nectar collectors performed waggle dances as well as round dances. Frisch tested to see if pollen collectors also performed both dances. This proved to be the case. Contrary \to his earlier conclussion, then, it was not the nature from the hive that determined the form of the dance. with round dances performed at distances of less than 100 meters or so from the hive, and waggle dances at greater distances.

Frisch reported his major new discoveries in two papers published in 1946. He also discussed them in his book of 1947 entitled Duftgelenkte Bienen im Dienst der Landwirtshaft und Imkerei, in which he described the results of his own experiments on how training bees to particular scents, such as that of red clover, could increase crop production.

Frisch’s work, which had attracted considerable attention before the war. became the subject of intense interest. The English ethologist W. H. Thorpe visited Frisch at Brunnwinkl in September 1948 to witness and repeat for himself Frisch’s key experiments. In an article in Nature in 1949. Thorpe confirmed the accuracy of Frisch’s remarkable findings. In the United States. Donald R. Griffin of Cornell University repeated Frisch’s experiments and invited Frisch to lecture at Cornell, an invitation \that developed into a three-month lecture tour of the United States in the spring of 1949.

Frisch’s new discoveries of 1945 and 1946 on the dance language of the bees opened up a host of new research topics for him. Having learned that in long-range flights bees orient themselves with respect to the position of the sun. he asked himself how bees orient themselves when the sun is not visible. A series of experiments led him to conclude— mistakenly, it now seems—that the bees perception of ultraviolet rays allows them to see the sun through a cloud cover that obscures the sun to the human eye. Of more lasting and fundamental importance. he made the remarkable discovery that bees perceive polarized light and are able to orient their dances according to the plane of polarization. Thus. with the sun obscured but a patch of blue sky visible to them, they are able to retain their sense of direction.

In the 1920’s Frisch’s student Ingeborg Beling had conducted pioneering studies on the temporal sense of bees. Frisch in the 1950’s proceeded to establish that thanks to their “internal clock,” bees are able to take into account the changes in the sun’s position during the course of the day and are thus able to use the sun (as well as conspicuous objects in the landscape) for orientation on long flights. This was another performance by bees that Frisch found almost too extraordinary to be true. but carefully designed “displacement” experiments showed that bees, by adjusting for the time of day and the sun’s diurnal course. can indeed use the sun’s position to find a given compass direction.

With his student Martin Lindauer. Frisch under-took a comparative study of the dance language of different varieties and species of bees. This revealed different “dialects” in the dance language of bees, and provided a basis for evaluating how the complex dance language exhibited by the black Austrian honeybee (Apis mellifera carnica). the original subject of Frisch’s researches, could have evolved from a more primitive form of dance, such as that discovered by Lindauer in the Indian dwarf honeybee Apis florea.

In his autobiography Frisch acknowledged that throughout his career he was plagued by periods when he doubted that he would have any more new ideas for research. At the beginning of his career he also worried about whether he would be able to think of interesting dissertation topics for his students The vast majority of the topics he investigated, however, proved immensely fruitful for additional studies of his own and for work conducted by his students and other researchers. His early studies on pigmentation and color adaptation led to his studies of color vision. His experiments on the color sense of bees led to his later studies of bees” senses of smell and taste, their “language,” and their orientation.

In his 1965 book Tanzsprache und Orientierung der Bienen which describes in detail the results of the work of Frisch and others on bees for the twenty years preceding the book’s publication, he was able to list forty-four doctoral students who had written dissertations on bees under his direction. Among them was Martin Lindauer, whose own students by then represented a third generation working in the tradition Frisch had established. Frisch’s skill in devising simple experiments under natural conditions was taken as a model by other scientists—for example the Dutch ethologist Nikolaas Tinbergen.

Frisch not only was exceptionally skilled as an experimenter but also was unusually gifted in his ability to write lucidly about biology. In addition to an introduction to bacteriology that he wrote for nurses during World War I, he produced a number of books of an introductory or semipopular nature, several of whitch have gone through many editions. Among these (with titles of the English translations and dates of the original edition and translations) were Aus dem Leben der Bienen (The Dancing Bees; 1927, 1954): Du und das Leben(Man and the Living World; 1936, 1962); Bees: Their Vision Chemical Senses, and Language (1950, rev. ed. 1971); Biologie (Biology; 2 vols., 1952, 1964); Erinnerungen eines Biologen (A Biologist Remembers; 1957, 1967); and, written with his son Otto, Tiere als Baumeister (Animal Architecture; 1974, 1983).

Frisch was always careful to point out, in speaking of a “language” of the bees, that the language metaphor had its limits, and that the dances of bees, though variable and adapted to different conditions, were mainly instinctive. The 1970’s witnessed considerable discussion of the issue of whether it is appropriate to attribute a “language” to bees, and whether the dances of bees actually serve to communicate information regarding the location of food sources to new recruits. Ethologists in general, however, regard the symbolic, communicative function of the bee dances as well established and, indeed, regard Frisch’s discovery of the dance language of the bees as the single most impressive discovery of classical ethology.

BIBLIOGRAPHY

1.Original Works. A nearly complete bibliography of Frisch’s scientific writings, with relatively few flaws in the citation of page and volume numbers, is in Journal of Comparative Physiology, 147 (1982), 420–422. For Frisch’s authoritative summing up of his own and other researches on bees, see his Tanzsprache und Orientierung der Bienen (Berlin, 1965), translated as The Dance Language and Orientation of Bees (Cambridge, Mass. 1967).

In addition to Frisch’s magnum opus of 1965, the following are among his most important scientific writings; “Studien über die Beziehungen der Pigment verschiebung im Facettenauge,” in Biologisches Zentralblatt, 28 (1908), 662–671, 698–704; “über die Beziehungen der Pigmentzellen in der Fischhaut zum sympathischen Nervensystem,” in Festschrift zum sechzigsten Geburstag Richard Hertwigs, III (Jena, 1910), 17–28; “Beiträge zur Physiologie der Pigmentzellen in der Fischhaut,” in Pflügers Archiv für die gesamte Physiologie des Menschen und der Tiere, 138 (1911), 319–387; “überden Farbensinn der Fische,” in Verhandlungen der Deutschen zoologischen Gesellschaft, 21 (1911), 220–225; “über farbige Anpassung bei Fischen, “in Zoologische Jahrbücher. Abteilung für allgemeine Zoologie und Physiologie, 32 (1912), 171–230; “Sind die Fische farbenblind?” ibid. 33 (1912), 107–126; “Der Farbensinn und Formensinn der Biene,” ibid. 35 (1914), 1–182, also separately reprinted (Jena, 1914); “Zur Streitfrage nach dem Farbensinn der Bienen,” in Biologisches Zentralblatt, 39 (1919), 122–139; “über den Geruchsinn der Biene und seine blütenbiologische Bedeutung,” in Zoologische Jahrbücher, Abteilung für allgemeine Zoologie and Physiologie, 37 (1919), 1–238; “über den Einfluss der Bodenfarbe auf die Fleckenzeichnung des Feuersalamanders,” in Biologisches Zentralblatt. 40 (1920), 390–414; “über die; Sprache” der Bienen. 1-III,” in Münchener medizinische wochenschrift, 67 (1920), 566–569, 68 (1921), 509–511, 69 (1922), 781–782; “über die “Sprache” der Bienen. Eine tierpsychologische Untersuchung,” in Zoologische Jahrbücher. Abteilung für allgemeine Zoologie und Physiologie, 40 (1923), 1–186; and” Das Problem des tierischen Farbensinnes,” in Naturwissenschaften, 11 (1923), 470–476.

These Writings were followed by “Ein Zwergels, der kommt, wenn man ihm pfeift,” in Biologisches Zentralblatt, 43 (1923), 439–446; “sinnesphysiologie und “Sprache” der Bienen,” in Naturwissenschaften, 12 (1924), 981–987; “Farbensinn der Fische und Duplizitätstheorie,” in Zeitschrift für vergleichende Physiologie, 2 (1925), 393–452; “Neue Versuche über die Bedeutung von Duftorgan und Pollenduft für die Verständigung im Bienenvolk,” ibid., 4 (1926), 1–21, with G, A. Rösch; “Versuche über den Geschmacksinn der Bienen. 1-III, “in Naturwissenschaften, 15 (1927). 321–327, 16 (1928), 307–315, 18 (1930). 169–174;’äber die Labyrinth-Funktionen bei Fischen,” in Verhandlungen der Deutschen zoologischen Gesellschaft, 33 (1929), 104–112; “Untersuchungen öber den Sitz des Gehörsinnes bei der Elritze,” in Zeitschrift för vergleichende Physiologie, 17 (1932), 686–801, with H. Stetter; “öber den Geschmacksinn der Biene,” ibid. 21 (1934), 1–156; “öber den Gehörsinn der Fische,” in Biological Reviews, 11 (1936), 210–246; “Psychologie der Bienen, “in Zeitschrift för Tierpsychologie, 1 (1937), 921; and “Zur Psychologie des Fisch-Schwarmes,” in Naturwissenschaften, 26 (1938), 601–606.

Frisch’s publications of the 1940’s include “Die Tänze und das Zeitgedächtnis der Bienen im Widerspruch,” in naturwissenschaften, 28 (1940), 65–69; “Die Bedeutung des Geruchsinnes im Leben der Fische,” ibid. 29 (1941), 321–333;’Über einen Schreckstoff der Fischhaut und seine biologische Bedeutung, ’in Zeitschrift fiir vergleichende Physiologie, 29 (1941), 46–145;’Die Werbetänze der Bienen und ihre Auslösung, ’in Naturwissenschaften. 30 (1942), 269–277;’versuche über die Lenkung des Bienenfluges durch Duftstoffe, ’ibid., 31 (1943), 445–460;’Die “sprache” der Bienen und ihre Nutzanwendung in der Landwirtschaft,” in Experientia. 2 (1946), 397–404; “Die Tänze der Bienen,” in Österreichische zoologische Zeitschrift, 1 (1946), 1–48, translated as “The Dances of the Honey Bee,” in Bulletin of Animal Behaviour, spec. no. (December 1947), 5–32; Duftgelenkte Bienen im Dienst der Landwirtschaft und Imkerei (Vienna, 1947); “Gelö und ungelöste Rätsel der Bienensprache,” in Naturwissenschaften, 35> (1948), 12–23, 38–43; and “Die Polarisation des Himmeslslichtes als orientierender Faktor bei den Tänzen der Bienen,” in Experientia, 5 (1949), 142–148.

Among Frisch’s later publications were “Die Sonne als Kompass im Leben der Bienen,” in Experientia6 (1950), 210–221; “Orientierungsvermögen und Sprache der Bienen,” in Naturwissenschaften. 38 (1951), 105–112; “Gibt es in der “Sprache” der Bienen eine Weisung nach oben oder unten?” in Zeitschrift für vergleichende Physiologie, 35 (1953), 219–245, with H. Heran and M. Lindauer;’Himmel und Erde in Konkurrenz bei der Orientierung der Bienen,” in Naturwissenschaften. 41 (1954), 245–253, with M. Lindauer; “Über die Fluggeschwindigkeit der Bienen und über ihre Richtungsweisung bei Seitenwind,” ibid., 42 (1955), 377–385, with M. Lindauer; “The “Language” and Orientation of the Honey Bee,” in Annual Review of Entomology, 1 (1956). 4558, with M. Lindauer;’Lernvermögen und erbgebundene Tradition im Leben der Bienen,” in Marieo Autuori et al., L’instinct dans le comportement des animaux et del’homme (paris, 1956), 345–386;’Über die “Missweisung” bei den richtungsweisenden Tänzen der Bienen, ’in Naturwissenschaften, 48 (1961), 585–594, with MLindauer; “Dialects in the Language of the Bees,” in Scientific American, 207 (1962), 78–87;’Honeybees: Do They Use Direction and Distance Information Provided by Their Dancers?” in Science, 158 (1967), 1072–1076; and “Decoding the Language of the Bee, ’ibid., 85 (1974), 663–668

II. Secondary Literature. On Frisch’s life and career the best source is his autobiography, Erinnerungen eines Biologen (Berlin, 1957), translated by Lisbeth Gombrich as A Biologist Remembers (Oxford, 1967).W.H. Thorpe, “Karl von Frisch, ’in Biographical Memoirs of Fellows of the Royal Society, 29 (1983).197–200, is a brief notice that borrows heavily from the autobiography. For a sampling of current work directly or indirectly in the tradition of research established by Frisch.and for other comments on Frisch as a scientist, see Bert Holldobler and Martin Lindauer, eds., Experimental Behavioral Ecology and Sociobiology; In Memoriam Karl von Frisch 1886–1982 (New york, 1985).

The author gratefully acknowledges the helpful comments of Donald R.Griffin and Martin Lindauer on an earlier draft of this paper.

Richard W. Burkhardt, JR.