The use of bee natural product for enhancing pollination is especially valuable in problematic crops that are generally avoided by bees. In the present research we attempted to enhance bee visitation to Male Sterile (M-S) tomato flowers generally used in the production of hybrid seeds. These flowers that lack both pollen and nectar are unattractive to bees that learn rapidly to avoid them. The specific objects were to elucidate the chemical composition of the exocrine products of two bumble bee species the North American Bombus impatiens and the Israeli B. terrestris. Of these, to isolate and identify a bee attractant which when sprayed on M-S tomato flowers will enhance bee visitation, and to provide a procedure of the pheromone application regime. During the research we realized that our knowledge of B. impatiens is too little and we narrowed the objective to learning the basic social behavior of the bees and the pattern of foraging in a flight chamber and how it is affected by biogenic amines. Colonies of B. impatiens are characterized by a high number of workers and a relatively small number of queens. Size differences between queens and workers are pronounced and the queen seems to have full control over egg laying. Only about 9% of the workers in mature colonies had mature oocytes, and there were no signs of a "competition phase" as we know in B. terrestris. Queens and workers differ in their exocrine bouquet. Queen's Dufour's gland possesses a series of linear, saturated and unsaturated hydrocarbons whereas that of workers contains in addition a series of wax-type esters. Bees were trained to either visit or avoid artificially scented electronic flowers in a flight chamber. Since bee also learned to avoid scented non-rewarding flowers we attempted to interfere with this learning. We tested the effect of octopamine, a biogenic amine affecting bee behavior, on the choice behavior of free-flying bumblebees. Our results show that octopamine had no significant effect on the bees' equilibrium choice or on the overall rate of the behavioral change in response to the change in reward. Rather, octopamine significantly affected the time interval between the change in reward status and the initiation of behavioral change in the bee. In B. terrestris we studied the foraging pattern of the bees on tomato flowers in a semi commercial greenhouse in Yad Mordechai. Bee learned very quickly to avoid the non- rewarding M-S flowers, irrespective of their arrangement in the plot, i.e., their mixing with normal, pollen bearing flowers. However, bees seem to "forget" this information during the night since the foraging pattern repeats itself the next morning. Several exocrine products were tested as visitation enhancers. Among these, tarsal gland extracts are the most attractive. The compounds identified in the tarsal gland extract are mostly linear saturated hydrocarbons with small amounts of unsaturated ones. Application was performed every second day on leaves in selected inflorescences. Bee visitation increased significantly in the treated inflorescences as compared to the control, solvent treated. Treatment of the anthers cone was more effective than on the flower petals or the surrounding leaves. Methanol proved to be a non-flower-destructive solvent. We have shown that bumble bees (B. terrestris) can be manipulated by bee-borne attractants to visit non-rewarding flowers. We have further demonstrated that the bees learning ability can be manipulated by applying exogenously octopamine. Both methods can be additively applied in enhancing pollination of desired crops. Such manipulation will be especially useful in tomato cultivation for hybrid seed production.
Use of Bee-Borne Attractants for Pollination of Nonrewarding Flowers: Model System of Male-Sterile Tomato Flowers
