Contains reports on one research project.National Institutes of Health (Grant 5 PO1 GM14940-07

Gahl, Robert A.

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XIV. COMMUNICATIONS BIOPHYSICSA. Animal CommunicationAcademic and Research StaffProf. H. Steven ColburnProf. Lawrence FrishkopfNathaniel Durlach1. THE SHAKING DANCE OF HONEYBEE WORKERS: EVIDENCEFOR AGE DISCRIMINATIONNational Institutes of Health (Grant 5 PO1 GM14940-07)Robert A. GahlOne of the dances of the honeybee Apis mellifera L. is called the shaking dance.The shaking dance is enacted by a worker bee (called the "shaker") that vibrates itsbody dorsoventrally by means of rapid contractions of the leg muscles. The shakerusually makes contact with another bee that remains motionless during the shake andthen resumes its previous activity. Workers perform this dance on all three castes ofbees,2 but we studied shaker bees only in the worker caste. A bee, after shaking once,will typically crawl across a portion of the comb and soon shake again, either alone, onanother bee, or on several bees together. A sequence of such events is termed a"shaking run. " Occasionally actions of trophallaxis or waggle dances are performedduring a shaking run.MethodsA small observation hive (4. 6 x 16 X 16 cm) was constructed to provide enough roomfor approximately 500 bees. A hybrid queen and several of her workers were placed inthe hive on June 25, 1972. Every day 10-30 newly emerged workers from a stock obser-vation hive were color-marked to enable individual identification,3 and added to the smallcolony. In all, 820 bees were marked, but because of attrition the population peakedbetween 200 and 350. This miniature colony was maintained for observation untilJuly 26, 1972.Observations were made of shaking dances performed by these bees, whose color-marking allowed for quick and accurate determination of age. The colony was observedat random times during the day. The data included the type of shaking situation (i. e.,shaking alone, one-on-one, or one-on-several), the age of the shaker, and if anotherbee was involved, the age of the bee shaken. Also, if trophallaxis occurred during ashaking run, the ages of the participants were recorded, and whether the shaker wasthe receiver or the provider of food was noted.QPR No. 113 169250w 200Zz 150I()L 100OcrLUim5 50z-5 0 5 10 15 20AGE DIFFERENCE (DAYS)Fig. XIV- 1. Number of one-on-one shaking dances as a function of theage difference (age of shaker minus age of bee shaken) ofthe bees involved. n (number of shaking dances) = 2237.3503002502000 5 10 15 20 25AGE (DAYS)Fig. XIV-2 Number ofof shakern = 2237.one-on-one shaking dances as a function of age(solid line) and of bee shaken (dashed line).QPR No. 113 170(XIV. COMMUNICATIONS BIOPHYSICS)High-speed movies of the stock observation hive were taken to provide informationon the duration and frequency of vibrations in shaking dances.ResultsDuring the 28-day period, 4949 shaking dances were observed. Of these, 2039(41. 2%) were performed alone, 2220 (44. 8%) were one-on-one, and 690 (14%) were donewhile straddling more than one bee. The data showed that 96.6% of the one-on-oneshaking dances were performed on bees not older than the shaker (Fig. XIV-1). Morethan 90% of the shaking dances were performed on bees not older than the shaker foreach age of shaker at which an adequate number of one-on-one shaking dances wasobserved. The age range of shakers was 0-24 days, and the age range of bees shakenwas 0-26 days. Histograms of number of shaking dances versus age show peaks at 9 and19 days for shakers and at 2 days for bees shaken (Fig. XIV-2).Only 141 acts of trophallaxis were observed during shaking runs, compared with4949 observed shaking dances. Of these, the shaker was the receiver of food 77.3% ofthe time.The films of 11 shaking dances performed by 9 different bees indicate that the aver-age duration of a dance is 1. 2 s (standard deviation 0. 26 s). The mean frequency ofvibrations within the shaking dance is 16. 3 Hz (standard deviation 5. 8 Hz).DiscussionA relationship has been found between the age of the shaker and of the bee shaken:the shaker is nearly always older. It is concluded that shaking is not performed at ran-dom, but with some discrimination determined by age. This implies some kind of agerecognition in the hive. Such recognition could depend upon physiological variation withage with respect to such traits as strength, health, glandular growth, or labor category(which is partially related to age and partially to food conditions in the hive), or somecombination of these. This behavior may be related to the waggle dance or to trophallac-tic behavior, which are occasionally incorporated into the shaking run.References1. M. D. Allen, "The 'Shaking' of Worker Honeybees by Other Workers," Anim.Behav. 7, 233-240 (1959).2. V. G. Milum, "Honeybee Communication," Amer. Bee J. 95, 97-104 (1955).3. K. V. Frisch, The Dance Language and Orientation of Bees (The Belknap Press ofHarvard University Press, Cambridge, Mass., 1967), pp. 14-17.QPR No. 113 171

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