Synthetic bee pollen foraging pheromone and uses thereof

Details Synthetic bee pollen foraging pheromone and uses thereof Synthetic bee pollen foraging pheromone and uses thereof

2002-02-04

2003-07-22

Swiatek, Robert P. (Department: 3643)

Bee culture

Method

Honeybee

Reexamination Certificate

active

06595828

ABSTRACT:

BACKGROUND OF THE INVENTION
Technical Field
The invention relates to a synthetic pheromone composition relating to the foraging behavior of bees, to its composition and to its uses.
Introduction
Social insects perform all the tasks of colony growth and maintenance concurrently by division of labor. One view is that division of labor is an emergent property that is self-organized through the interactions of individual workers with stimuli in their environment (Page R. E., Jr. et al.
PSA
2:28-298 (1990)). A division of labor emerges when individuals have different response thresholds for stimuli that release behavioral responses. The evolution of specific patterns of organization results as a consequence of selection acting on genetic variability affecting response threshold distributions of colonies.
Bees are insects of the order Hymenoptera, which feed on pollen and nectar. They constitute a group of about 20 000 species throughout the world, known taxonomically as the Superfamily Apoidea. Honey bees of the genus Apis belong to the family Apidae, a sub-group of this superfamily. Although the question of how many honey bee species exist is still debated among taxonomists, at least four species are commonly recognized: the dwarf, or midget, bee
Apis florea,
the giant, or rock, bee
Apis dorsata,
the oriental (Indian, Chinese, Japanese, etc.) bee
Apis cerana,
and the common (European, African, etc.) honey bee
Apis mellifera.
The existence of another giant bee,
Apis laboriosa,
has recently been confirmed.
Bees of the family Apoidea are eusocial insects, that is to say that they engage in favorable social activity. Depending on the species, a colony of bees consists of a queen, dozens to thousands of workers, and at certain seasons of the year anywhere from a few to a few-hundred drones. Among the members of the colony there is division of labor and specialization in the performance of biological functions.
Bees construct variously shaped cells to rear their brood and to store their food. In the genus Apis, general utilization of comb space is similar among the species, with honey stored in the upper part of the comb, with, beneath it, rows of pollen-storage cells, worker-brood cells, and drone-brood cells, in that order. The groundnut-shaped queen cells are normally built at the lower edge of the comb.
As an inherited behavior characteristic, all bee colonies tend to store a certain amount of honey and pollen as their food reserve. The quantity of food stored depends upon several factors, including the seasonal availability of forage, the worker population of the colony and its rate of reproduction, the capacity of the nest, etc. Another important inherited behavior characteristic lies in the colony's natural site of comb construction. For instance, some Apis species build single comb nests in the open, while others build multiple-comb nests in dark cavities.
In honey bees, studies suggest that colony environment modulates foraging behavior. The amount of pollen stored in the comb affects the proportion of pollen foragers. More stored pollen results in less pollen foraging (Allen M. D. et al.,
Ann Applied Biol
44:649-656 (1956); Fewell, J. H. et al.,
Behav Ecol Sociobiol
30:387-393 (1992); Fewell, J. H. et al.,
Experientia
49:1106-1112 (1993)). The amount of young larvae also affects the proportion of foragers collecting pollen: more larvae and empty space result in more pollen foraging (Al-Tikrity W. S. et al., J Apic Res 11:9-12 (1972); Barker, R. J.,
J Apic Res
10:23-26 (1971); Dreller C. et al.,
Behav. Ecol. Sociobiol.
45:227-233 (1999); Eckert C. D. et al.,
Oecologia
97:248-255 (1994); Fewell, J. H. et al.,
Behav Ecol Sociobiol
30:387-393 (1992); Free, J. B.
Anim Behav
15:134-144 (1967); Free, J. B.
App Anim Ethol
5:173-178 (1979); Jaycox 1970; Todd F. E. et al.,
J Econ Ent
63:148-149 (1970)). Larval substances soluble in hexane releases pollen foraging (Pankiw T. et al.,
Behav Ecol Sociobiol
44:193-198 (1998)).
Foraging behavior and the mechanisms that regulate foraging activity are important components of social organization. Recently the honey bee proboscis extension response to sucrose has been identified as a “window” into a bee's perception of sugar (Page R. E., Jr. et al.
PSA
2:28-298 (1990)). The sucrose response threshold measured in the first week of adult life, prior to foraging age, predicts forage choice (Pankiw T. et al.,
Behav Ecol Sociobiol
47:265-267 (2000)). Bees with low response thresholds are more likely to be pollen foragers and bees with high response thresholds are more likely to forage for nectar. There is an associated genetic component to sucrose response thresholds and forage choice, such that bees selected to hoard high quantities of pollen have low response thresholds and bees selected to hoard low quantities of pollen have higher response thresholds.
The number of larvae in colonies also affects the number of bees foraging for pollen. Hexane extractable compounds from the surface of larvae (brood pheromone) significantly increase the number of pollen foragers (Pankiw T. et al.,
Behav Ecol Sociobiol
44:193-198 (1998); Pankiw T. et al.,
J Comp Physiol A
185:207-213 (1999); Pankiw T. et al.,
Behav Ecol Sociobiol.
49:206-213 (2001)). Brood pheromone decreases the sucrose response threshold of bees suggesting a pheromone-modulated sensory-physiological mechanism for regulating foraging division of labor. Honey bees respond reflexively to sucrose by extending the proboscis when a sufficiently concentrated solution touches the antennae (Bitterman, M. E. et al.,
J Comp Psychol
97:107-119 (1983)). Brood pheromone significantly decreased response thresholds as measured in the proboscis extension response assay (PER-RT assay), a response associated with pollen foraging (Page R. E. et al.,
Anim Behav
50:1617-1625 (1995)). Pollen foragers have lower sucrose response thresholds than do nectar foragers (Page R. E. et al.,
J Comp Physiol A
182:489-500 (1998)). Water foragers have the lowest response thresholds to sucrose as pre-foragers, pollen foragers have the next lowest response thresholds, followed by nectar foragers, and foragers returning empty have the highest response thresholds (Pankiw T. et al.,
Behav Ecol Sociobiol
47:265-267 (2000)).
Response thresholds to sucrose are also plastic, modulated by nutritional status and previous foraging experience. Response thresholds are lower in bees fed lower quality sucrose solutions for 24 hours than bees fed high quality solutions. Previous foraging experience with low or high quality sucrose solutions modulates response thresholds in the same way. Genotype places a constraint on phenotypic response threshold plasticity such that strains of bees that were selected to hoard high or low quantities of pollen (Page R. E. et al.,
Anim Behav
50:1617-1625 (1995)), when fed high or low quality sucrose solutions have response thresholds that are proportionately modulated, however strain differences are maintained at all concentrations fed.
Reports have suggested that nectar foraging may be induced by nurse bees by the secretion of a substance inhibiting pollen foraging in workers. (Camazine, S.
Behav Ecol Sociobiol.
32:265-272 (1993)). By this proposed mechanism, foraging behavior is controlled by the indirect effects of nurse bees producing a pollen-foraging inhibitor, with the amount of inhibitor available to feed to foragers depending on the amount of pollen stored and the number of larvae that must be fed. In a recent book on honey bee social organization, this explanation is supported as the way bees regulate pollen collection. (Seeley T. D.
The social physiology of honey bee colonies.
“Harvard University Press, Cambridge Mass.” (1995)).
The direct value of honey bees as pollinators is valued at more that $14 billion annually in the United States. However, commercial honey bee populations are continually dwindling due to recently imported parasites and pathogens, and due to the invasion of the Africanized Honey Bee.
According to Morse et al., the val

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