Foraging behavior of honeybees and hormones

Dr. Janko Bozic

University of Ljubljana, Biotechnical faculty, Department of Biology

 Scientific interest

Managing and planning foraging of honeybee colonies is a priority of any beekeeper in the world. In early eighties I started investigating basic concepts of foraging behavior of honeybees. Dance communication is mayor biological mechanism that promotes foraging success of a honeybee colony. Two honeybees are always involved in the dance communication, the dancer bee and bee that collect the dance message. The dancer bee emits the direction and distance message whereas the bees around the dancer receive the information. In the eighties the most popular hypothesis of the message transfer proposed that the message is transmitted via sound vibrations that are detected by bees in the near acoustic field around the dancer (Kirchner, Michelsen). A novel definition of behavior of bees surrounding the dancer was needed. The followers bees move along with the waggling dancer always touching its thorax and abdomen with both antennae.The attender bees stand around the dance field, do not follow and do not touch the dancer. The initially inactive bee starts attending the dancer, gets excited and potentially starts following the dancer. Not all the attenders will follow the dancer, however only the follower bees can find a distant food source successfully. Before flying in the direction of the foraging site the follower bees leave the hive and orient around the hive several times in succession. The honeybee foraging behavior is always accompanied by large changes in hemolymh sugar concentrations, concentrations of glucose, fructose and trehalose in hemolymph are activity dependent. The concentration of trehalose is high during departure from the hive and it decreases during the hive return. The concentrations of glucose and fructose are high in the follower bees and in the foraging bees. Biogenic amines and juvenile hormone also control some aspects of the foraging behavior. Concentrations of biogenic amines change during the waggle dance activity. The follower bees have higher activity of dopamine than the inactive bees. Increased octopamine concentrations are linked to feeding arousal of honeybees. Serotonin concentrations increase during the decline of foraging activity. The queen pheromone is transferred passively from the queen to its attending bees, and further from the queen attending bees to other bees in the hive.

Ripe fruits frequently contain ethanol and facilitate natural ethanol exposure of honeybees. With the group of C. Abramson from Oklahoma State University of we started investigating ethanol effects in honeybee behavior. Honeybees are thus becoming a model for drug abuse and toxicity studies in insects.


1. BOŽIČ, Janko, VALENTINČIČ, Tine. Attendants and followers of honey bee waggle dances. J. Apic. Res., 1991, (30): 3/4, pp. 125-131.
2. BOŽIČ, Janko, VALENTINČIČ, Tine. Quantitative analysis of social grooming behavior of the honey bee Apis mellifera carnica. Apidologie, 1995, (26) pp. 141-147.
3. BOŽIČ, Janko, WOODRING, Joseph. Effect of activity on the haemolymph sugar titres in honey bees. J. Apic. Res., 1997, (36):1, pp. 33-39.
4. BOŽIČ, Janko, WOODRING, Joseph. Variations of brain biogenic amines in mature honeybees and induction of recruitment behavior. Comp. biochem. physiol., Part A Physiol., 1998, (120) pp. 737-744.
5. BOŽIČ, Janko, WOODRING, Joseph. Variation in JH synthesis rate in mature honeybees and its possible role in reprogramming of hypopharingeal gland function. Pflügers Arch, 2000, (439):3, suppl., pp. R163-R164.
6. KRALJ, Jasna, BOŽIČ, Janko. Activity of attendants after licking and palpating the queen in honeybee (Apis mellifera carnica). Neth. j. zool., 2001, (51):4, pp. 415-419.
7. BOŽIČ, Janko, ABRAMSON, Charles I. Following and attending - two distinct behavior patterns of honeybees in a position to collect the dance information. Mellifera, 2003, (3):6, pp. 48-55.
8. ABRAMSON, Charles I., KANDOLF, Andreja, SHERIDAN, Audrey, DONOHUE, Darius, BOŽIČ, Janko, MEYERS, Julia E, BENBASSAT, Danny. Development of an ethanol model using social insects : III. preferences for ethanol solutions. Psychol. rep., 2004, (94) pp. 227-239.
9. BOŽIČ, Janko, ABRAMSON, Charles I., BEDENČIČ, Mateja. Reduced ability of ethanol drinkers for social communication in honeybees (Apis mellifera carnica Poll.). Alcohol (Fayetteville, N.Y.), (Alcohol). [Print ed.], 2006, vol. 38, spp. 179-183.
BOŽIČ, Janko, DICESARE, John, WELLS, Harrington, ABRAMSON, Charles I. Ethanol levels in honeybee hemolymph resulting from alcohol ingestion. Alcohol (Fayetteville, N.Y.), (Alcohol). [Print ed.], 2007, vol. 40, in press.
11. ABRAMSON, Charles I., WELLS, Harrington, BOŽIČ, Janko. A social insect model for the study of ethanol induced behavior: the honey bee. V: YOSHIDA, Rin (ur.). Trends in alcohol abuse and alcoholism research. Hauppauge, New York: Nova Science Publishers, 2007, pp. 197-218.


Beekeeping interest

I’m also interested in different aspects of beekeeping technology. Beekeepers in Slovenia use their own hive type - Alberti-Znidersic or simply AZ hive. This standard enable us to keep hives in beehouse or even beehouse on the wheels (no upload and download when moving hives). Disadvantage of this technology is more expensive hive and more hive tools then standard LR hives, what makes beekeeping less profitable. I’m looking for technological solutions which would be more profitable and still friendly for the beekeepers.


I’m a teaching assistant for Ethology Lab. Over the years, before I started this job, several different practical exercises were developed using fishes, mousses, rats and also some other animals. I’m trying to add few exercises with honeybees. Since 2000 I teach also Insect Biology course. Since 2002 I teach graduate course Comparative endocrinology and since 2004 I teach Beekeeping  course at undergraduate and graduate level.

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Related Links

  • Beekeeping in Slovenia
  • International Bee Research Association
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    Janko Bozic was born on December 21, 1963, to Anton and Cecilija Bozic in Cretez near Krsko, Slovenia. He attended primary school in Krsko, completing high school in Brezice in 1982. In October of 1983 he entered Biology program at the University of Ljubljana where he achieved the Bachelor of Science degree in February 1988. For a year an a half he took a position in quality control division of paper mill factory in Krsko, Slovenia. In the October 1989 he got teaching assistant position at the University of Ljubljana and entered graduate study. He received Master degree from the Department of Biology, University Ljubljana, in September 1992. In January 1993 he entered the doctoral degree program in the Department of Zoology and Physiology at Louisiana State University. Dissertation was successfully defended on April 12, 1996. Currently Dr. Bozic is working as teaching assistant of Ethology at the Department of Biology, Biotechnical faculty at the University of Ljubljana. Mr. Bozic is married, has two sons and a daughter . They live in Ljubljana.
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    Published internet papers

    Queen mating behavior as an example of basic science observation in beekeeping technology development
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    Currently I'm involved in research in collaboration with Prof. C. Abramson (USA) studying effects of ethanol on honeybee behavior. I had also applicative project with company  Endovital Anton Fabjan s.p. on effects of feeding beet root syrup to honeybees.

    I’m also interested in:

  • regulation of foraging behavior expression with the emphasis on waggle dance communication
  • significance of dance signals and dance signal transmission
  • social grooming behavior
  • varroa resistance mechanisms
  • behavior patterns which are involved in comb building
  • swarming behavior, swarm formation, behavior process which enable new nest location, physiological changes during swarming
  • behavior and physiology of different ecotypes of carniolan bees
  • honeydew production and honeydew processing by bees
  • beekeeping automation and hive technology development


    Please, contact me if you are interested in cooperation.

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    University of Ljubljana
    Biotechnical faculty
    Department of Biology
    Vecna pot 111, p.p. 2995
    1001 Ljubljana, SLOVENIA
    tel. (386)1-423-33-88
    fax. (386)1-257-33-90
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