Progress 10/01/00 to 09/30/05
Outputs RNA knockdown of vitellogenin was shown to affect the age of onset of foraging and foraging behavior of honey bees. They were also more responsive to low concentrations of sugar when tested with the proboscis extention response protocols. These results confirmed our hypothesis that the onset of foraging and foraging behavioral decisions are linked to reproductive regulatory networks of honey bees. Studies with wild type bees gave further support. We found that worker bees with more ovarioles per ovary foraged earlier in life, were more likely to collect pollen, and when they foraged for nectar collected nectar with lower sugar concentrations.
Impacts This work will lead to a better understanding of the evolution of division of labor in social insects and may lead to better methods to manipulate honey bees for pollination and honey production.
Publications
- Amdam, G., A. Csondes, M. K. Fondrk, and R. E. Page. 2006. Complex social behaviour derived from maternal reproductive traits. Nature 439: 76-78.
- Amdam, G. V., K. Norberg, R. E. Page, J. Erber, R. Scheiner. In press. 2006. Downregulation of vitellogenin gene activitiy increases the gustatory responsiveness of honey bee workers (Apis mellifera L.). Behavioral Brain Research.
- Rueppell, O., S. Chandra, T. Pankiw, M. K. Fondrk, M. Beye, G. Hunt and R. E. Page. 2006. The genetic architecture of sucrose responsiveness in the honey bee (Apis mellifera L.). Genetics 172: 243-251.
- Rueppell, O., R. E. Page, and M. K. Fondrk. 2005. Male maturation rate respond to selection on pollen hoarding in honeybees. Animal Behaviour 71: 227- 234.
- Humphries, M. A., M. K. Fondrk, and R. E. Page. 2005. Locomotion and the pollen hoarding behavioral syndrome of the honey bee (Apis mellifera L.). Journal of Comparative Physiology A 191: 669-674.
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Progress 01/01/04 to 12/31/04
Outputs We continue to maintain strains of bees selected for their pollen storage behavior. These strains have been extremely useful tools for many different studies we have done over the years. This year they were instrumental in determining the underlying physiological basis of many traits that are correlated with pollen foraging behavior. This has been an enigma for years. We found that the high strain bees, and wild type pollen foragers, have orvaries with more ovarioles and are in a higher state of reproductive activation.
Impacts This work will lead to a better understanding of the evolution of division of labor in social insects and may lead to better methods to manipulate honey bees for pollination and honey production.
Publications
- Amdam, G.V., K. Norberg, M.K. Fondrk, and R.E. Page. 2004. Reproductive ground plan may mediate colony-level effects on individual foraging behavior in honey bees. Proceedings of the National Academy of Science USA 101: 11350-11355.
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Progress 01/01/03 to 12/31/03
Outputs We completed 5 QTL maps of the honey bee. We mapped economically important traits including the age of initiation of foraging behavior, foraging behavioral traits, and sensitivity of bees to sugar. The maps are being used to assist in the assembly of the now complete honey bee genome sequence. We are looking for candidate genes in the regions of the QTL. We also isolated and characterized the complementary sex determining (csd) gene for the honey bee.
Impacts The maps we have constructed will greatly aid in assembly of the honey bee genome and the identification of major genes of economic importance. The csd gene is the single most important gene for honey bee breeding. Identification will lead to better methods of breeding.
Publications
- Beye, M, M. Hasselmann, M. K. Fondrk, R. E. Page, and S. W. Omholt. 2003. The gene csd is the primary signal for sexual development in the honeybee and encodes an SR-type protein. Cell 114: 419-429.
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Progress 01/01/02 to 12/31/02
Outputs We have continued to test the efficacy of our patented synthetic brood pheromone on pollen foraging behavior. We tested new formulations in commercial hives and demonstrated an increase in pollen collection in the treated colonies for two days. We also tested the effects of long term exposure to brood pheromone. Commercial colonies were treated continuously for 10 days and were then examined for areas of stored pollen, brood, and honey. We found no obvious negative effects. We completed 5 new saturated genomic maps of the honey bee. We found at least 6 new statistically significant quantitative trait loci (QTL) that affect foraging behavior.
Impacts We presented the results of the brood pheromone studies to almond growers. We will do additional tests in commercial colonies this next year to determine the efficacy of using brood pheromone as an enhancer of almond pollination.
Publications
- Guzman-Novoa, E., G. J. Hunt, R. E. Page, and M. K. Fondrk. 2002. Genetic correlations among honey bee (Apis mellifera L.) behavioral characteristics and body size. Annals of the Entomological Society of America 95: 402-406.
- Gadau, J., C. U. Gerloffs, N. Krugers, H. Chan, P. Schmid-Hempel, A. Wille, and R. E. Page. 2001. A linkage analysis of sex determination in Bombus terrestris (L.) (Hymenoptera: Apidae). Heredity 87: 234-242.
- Pankiw, T., D. R. Tarpy, and R. E. Page. 2002. Genotype and rearing environment affect honeybee perception and foraging behaviour. Animal Behavior. In Press
- Gadau, J., R. E. Page, and J. H. Werren. 2002. The genetic basis of interspecific differences in wing size in Nasonia (Hymenoptera: Pteromalidae): major quantitative trait loci and epistasis. Genetics 161: 673-684.
- Tarpy, D. R. and R. E. Page. 2002. Sex determination and the evolution of polyandry in honey bees (Apis mellifera). Behavioral Ecology and Sociobiology 52: 143-150.
- DeGrandi-Hoffman, G., R. E. Page, J. Martin, and M. K. Fondrk. 2002. Can the frequency of reduced Varroa jacobsoni (oud.) fecundity in honey bee (Apis mellifera L.) pupae be increased by selection? Apidologie. In Press
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Progress 01/01/01 to 12/31/01
Outputs Honey bee larvae produce chemical compounds (pheromone)that stimulate pollen foraging in adults. We have produced a synthetic pheromone and demonstrated its efficacy under controlled conditions. This year we tested the synthetic pheromone under commercial-pollination conditions in almond orchards. The pheromone treated colonies collected more almond pollen than the controls. We have continued our genetic studies on honey bee pollen foraging. We have identified candidate genes and signal transduction pathways responsible for differences in lines of bees selected for their foraging behavior. The regulation of sex determination in honey bees involves a single major gene. We isolated, cloned, sequenced and characterized this gene.
Impacts We filed a patent for the synthetic brood pheromone.
Publications
- Pankiw, T., K.D. Waddington, and R.E. Page. 2001. Modulation of sucrose response thresholds in honey bees (Apis mellifera L.): influence of genotype, feeding, and foraging experinece. Journal of Comparative Physiology A 187:293-301.
- Page, r.E., M.K. Dondrk, G.J. Hunt, E. Guzman-Novao, M.A. Humphries, K. Nguyen, and A. S. Gren. 2001. Genetic dissection of honeybee (Apis mellifera L.) foraging behavior. Journal of Heredity 91:474-479.
- Pankiw, T. and R.E. Page. 2001. Brood pheromone modulates honey bee sucrose response thresholds (Apis mellifera L.). Behavioral Ecology and Sociobiology 49:206-213.
- Hasselmann, M., M.K. fondrk, R.E. Page, and M. Beye. 2001. fine scale mapping in the sex locus region of the honey bee (Apis mellifera). Insect Molecular Biology 10:605-608.
- Pankiw, T. and R.E. Page. 2001. Genotype and colony environment affect honey bee (Apis mellifera L.) development and foraging behavior. Behavioral Ecology and Sociobiology 51:87-94.
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Progress 01/01/00 to 12/31/00
Outputs Honey bee larvae produce chemical substances that release pollen foraging behavior in workers. We demonstrated that chemical extracts of larvae can more than double the number of foragers within an hour of application. We identified compounds produced by larvae and produced a synthetic pheromone that results in behavioral changes similar to those produced by larvae. Tests using the synthetic pheromone doubled the number of pollen foragers in colonies and lasted for about 1 hour. We continue to test these compounds and develop better release mechanisms.
Impacts (N/A)
Publications
- Pankiw, T. and Page, R. E. 2000. Response thresholds to sucrose predict foraging division of labor in honeybees. Behavioral Ecology and Sociobiology 47: 265-267.
- Nielsen, D. I., Ebert, P. R., Page, R. E., Hunt, G. J. and Guzman-Novoa, E. 2000. Improved polymerase chain reaction-based mitochondrial genotype assay for identification of the Africanized honey bee (Hymenoptera: Apidae). Annals of the Entomological Society of America 93: 1-6.
- Guzman-Novoa, G., Page, R. E., Spangler, H. G. and Erickson, E. H. 1999. A comparison of two assays to test the defensive behavior of honey bees (APIS MELLIFERA L.). Journal of Apicultural Research 38: 205-209.
- Nielsen, D. I., Ebert, P. R., Page, R. E., Hunt, G. J. and Guzman-Novoa, E. 2000. Improved polymerase chain reaction-based mitochondrial genotype assay for identification of the Africanized honey bee (Hymenoptera: Apidae). Annals of the Entomological Society of America 93: 1-6.
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Progress 01/01/99 to 12/31/99
Outputs Honey bee workers specialize on kinds of food materials they collect as foragers. Some specialize on collecting water, others pollen or nectar. Over the past 10 years we have been studying the underlying genetic basis for this behavioral variation. During the past year we determined that there is a close association between the age that a bee initiates foraging behavior and whether she comes from a strain selected for pollen or nectar collecting. These results are interesting because they now demonstrate a set of correlated traits that were previously thought to be independent. We have genetically mapped quantitative trait loci that affect the sensory physiological trait, responsiveness to sugar, foraging behavioral decisions (forage for pollen, nectar, or water) and affect the concentration of sugar in nectar collected by foragers. In addition we now can show that the foraging behavior correlates with the age of initiating foraging. Combined, these results suggest
that our three mapped QTLs are exhibiting broad pleiotropic effects. We have mapped 3 QTLs associated with the above traits. This year we have conclusively demonstrated that these QTLs are not independent in their action on these correlated traits. We have found strong interaction effects between two QTLs for different components of foraging behavior.
Impacts (N/A)
Publications
- Hunt, G. J., Collins, A.M., Rivera, R., Page, R.E., and Guzman-Novoa, E. 1999. Quantitative trait loci influencing honey bee alarm pheromone levels. Journal of Heredity 90: 585-589.
- Pankiw, T., and Page, R.E. 1999. The effect of genotype, age, sex, and caste on response thresholds to sucrose and foraging behavior of honey bees (APIS MELLIFERA L.). Journal of Comparative Physiology A 185: 207-213.
- Scheiner, R., Erber, J., and Page, R.E. 1999. Tactile learning and the individual evaluation of the reward in honey bees. Journal Comparative Physiology A 185: 1-10.
- Beye, M., Hunt, G.J., Page, R.E., Fondrk, M.K., Grohmann, L., and Moritz, R.F.A. 1999. Genetics 153: 1701-1708.
- Guzman-Novoa, E. and Page, R.E. 2000. Foraging behavior of European and European X Africanized Hybrid Bees (APIS MELLIFERA L.). Annals of the Entomological Society of America. In press.
- Nielsen, D. I., Ebert, P.R., Hunt, G.J., Guzman-Novoa, E., Kinnee, S.A., and Page, R.E. 1999. Identification of Africanized honey bees (Hymenoptera: Apidae) incorporating morphometrics and an improved PCR mitotyping procedure. Annals of the Entomological Society of America 92: 167-174.
- Dreller, C, Page, R.E., and Fondrk, M.K. 1999. Regulation of pollen foraging in honeybee colonies: effects of young brood, stored pollen, and empty space. Behavioral Ecology and Sociobiology 45: 227-233.
- Guzman-Novoa, E., Page, R.E., and Prieto-Merlos, D. 1998. Queen introduction , acceptance, and survival in honey bee (Hymenoptera: Apidae) colonies of a tropical, Africanized region. Journal of Economic Entomology 91: 1290-1294.
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Progress 01/01/98 to 12/31/98
Outputs We continue to select and maintain two strains of honey bees that differ in the amount of pollen they store and in the foraging behavior of workers derived from them. This year we designed an experiment to explore the epistatic effects of three quantitative trait loci (QTLs) that we have mapped that affect differences in foraging behavior between our selected strains. We are now conducted analyses of molecular markers to determine the QTL alleles inherited by foragers that we have sampled. We continued our studies of the sensory physiology of bees derived from these strains. We previously showed that pollen foragers extend their tongues (probosci) with a higher probability than do nectar foragers when their antennae are stimulated with lower concentrations of sucrose solutions. We tested high and low strain worker bees of different ages and found differences in sucrose responses between them within a few days of emerging as adults. This suggests a genetic effect on
response thresholds. This past year we determined that the response threshold of worker honey bees that is 1 week old (2 weeks prior to actually foraging) correlates with the kinds of materials she collects when she initiates foraging. We also determined that response thresholds to sucrose correlate with associative learning in bees and that the high and low pollen hoarding strains differ dramatically with respect to different components of associative learning. We are continuing these studies.
Impacts (N/A)
Publications
- PANKIW, T., PAGE, R.E. AND FONDRK, M. Kim. 1998. Brood pheromone stimulates pollen foraging in honey bees (APIS MELLIFERA). Behav. Ecol. Sociobiol. 44: 193-198.
- PAGE, R.E., ERBER, J. AND FONDRK, M.K. 1998. The effect of genotype on response thresholds to sucrose and foraging behavior of honey bees (APIS MELLIFERA L.). J. Comp. Physiol. A 182:489-500.
- WADDINGTON, K.D., NELSON, C.M. AND PAGE, R.E. 1998. Effects of pollen quality and genotype on the dance of foraging honey bees. Animal
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Progress 01/01/97 to 12/01/97
Outputs We continue to select and maintain two strains of honey bees that differ in the amount of pollen they store and in the foraging behavior of workers. This year we were able to map and confirm a third quantitative trait locus that affects differences in foraging behavior. This locus, PLN3, was demonstrated to affect the quantities of pollen stored by colonies, the size of nectar and pollen loads collected by foragers, and the concentrations of sucrose in nectar that are found acceptable by nectar foragers. We continued our studies of the sensory physiology of bees derived from these strains. We previously showed that pollen foragers extend their tongues (probosci) with a higher probability than do nectar foragers when their antennae are stimulated with lower concentrations of sucrose solutions. We tested high and low strain worker bees of different ages and found differences in sucrose responses between them within a few days of emerging as adults. This suggests a
genetic effect on response thresholds. We also found that drones and queens differ in their responses in the same ways. This should allow us to select virgin queens and drones, mate them, and produce high and low sucrose responding strains. We have a working hypothesis that the sucrose response thresholds are directly related to whether bees forage for pollen or nectar. If true, then our high responding strain should be like our high pollen hoarding strain, while our low responding strain should be like our low pollen hoarding strain.
Impacts (N/A)
Publications
- Page, R.E. 1997. The evolution of insect societies. Endeavour 21:
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Progress 01/01/95 to 12/30/95
Outputs In 1990, we initiated a selection program designed to produce two strains of honey bees that differed with respect to the quantities of pollen that they stored. Colonies of our high strain stored an average of six times more pollen than the low strain colonies. Using 38 colonies derived from generation 2 parents, we then constructed a genetic linkage map of major quantitative trait loci that affect this pollen hoarding trait and that have a direct effect on pollen foraging behavior of individual worker honey bees. This year we completed a new linkage map based on generation 7 parents. This map is based on 153 colonies. We are currently performing analyses to determine if there are any new QTLs for pollen foraging behavior that we failed to detect with our first map.
Impacts (N/A)
Publications
- PAGE, R.E., WADDINGTON, K.D., HUNT, G.J. and FONDRK, M.K. 1995. Genetic determinants of honey bee foraging behavior. Animal Behavior 50:1617-1625.
- HUNT, G.J., PAGE, R.E., FONDRK, M.K. and DULLUM, C.J. 1995. Major quantitative trait loci affecting honey bee foraging behavior. Genetics 141:1537-1545.
- PAGE, R.E. and FONDRK, M.K. 1995. The effects of colony-level selection on the social organization of honey bee (APIS MELLIFERA L.) colonies: colony-level components of pollen hoarding. Behavioral Ecology and Sociobiology 36:135-144.
- PAGE, R.E., ROBINSON, G.E., FONDRK, M.K. and NASR, M.E. 1995. Effects of worker genotypic diversity on honey bee colony development and behavior (APIS MELLIFERA L.). Behavioral Ecology and Sociobiology 36:387-396.
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Progress 01/01/94 to 12/30/94
Outputs We initiated in 1990 a breeding program designed to establish two strains of bees that collect and store large and small quantities of pollen, respectively. We were successful in this endeavor and have used these strains to study the genetic determinants of this behavior. This year we completed a genomic map for the honey bee that is based on DNA markers. We were able to locate on this map the major sex determining gene of honey bees. In addition, we localized two genes (quantitative trait loci, QTL) that have major effects on both the foraging behavior of individual worker honey bees and the amount of pollen stored by colonies. These two QTLs (designated PLN1 and PLN2) explain nearly 60 percent of the total phenotypic variance for stored pollen observed in a backcross population derived from our high and low pollen hoarding strains. This spring and summer we produced more than 200 new backcross colonies and measured them for stored pollen. These colonies will be used
to map potential, additional pollen hoarding QTLs that have smaller effects and to help localize PLN1 and PLN2 with greater precision.
Impacts (N/A)
Publications
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Progress 01/01/93 to 12/30/93
Outputs We initiated in 1990 a honey bee breeding program designed to establish a demonstration project for controlled breeding, and to develop basic management practices associated with selected bee breeding on a commercial level. We have selected honey bees that collect and store relatively high and low quantities of pollen. We are now in our sixth generation of selection. We crossed our high and low strains and tested the hybrids. We found that the hybrids performed like low-strain colonies for the amount of stored pollen, suggesting that pollen hoarding is a recessive trait. However, hybrid colonies stored significantly more honey, suggesting that honey production is over-dominant. This year we outcrossed queens from our high strain colonies to drones from commercial colonies that were preselected for high pollen stores. We are now crossing the outcrossed colonies to regenerate more genetic variability from which we can continue our selection.
Impacts (N/A)
Publications
- PAGE, R.E., JR., FONDRK, M.K. and ROBINSON, G.E. 1993. Selectable components of sex allocation in colonies of the honeybee (APIS MELLIFERA L.). Behav. Ecol. 4(3):239-245.
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Progress 01/01/92 to 12/30/92
Outputs We initiated a honey bee breeding program in 1990 that was designed with two objectives. The first objective was to establish a demonstration program of controlled breeding within genetically closed populations. From this program we hoped to solve the basic management problems and show California queen producers how they could raise commercially manageable, preferably European, queens in an Africanized environment. The second objective was to select for traits that are commercially important in order to make such a breeding program economically viable. We have succeeded in both objectives. We have selected four generations for pollen collecting. Colonies from our high pollen collecting lines store more than 5 times as much pollen and have about 40% more pollinators than our low lines. This year we continued our test program using these bees for pollination that involves queen producers, pollination service beekeepers, a pollination contract broker, and alfalfa and
almond producers. Our results from alfalfa pollination demonstrated that colonies with high line queens had significantly more pollen stored in the comb than unselected, commercial colonies. In almonds, we found that high line colonies stored more than twice as much pollen as commercial bees used for comparison. Individual high line colonies have about 1.8 times more pollen coming into the hive per unit time.
Impacts (N/A)
Publications
- No publications reported this period.
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Progress 01/01/91 to 12/30/91
Outputs Honey bee breeding program was initiated in 1990 that was designed with two objectives. The first objective was to establish a demonstration program of controlled breeding within genetically closed populations. From this program we hoped to solve the basic management problems and show California queen producers how they could raise commercially manageable, preferably European, queens in an Africanized environment. The second objective was to select for traits that are commercially important in order to make such breeding program economically viable. Both objectives were met. Four generations have been selected for pollen collecting. Colonies from high pollen collecting lines store more than five times as much pollen and have about 40% more pollinators than our low lines. This year a test program was implemented using these bees for pollination that involves queen producers, pollination service beekeepers, a pollination contract broker, and alfalfa and almond
producers. Results from alfalfa pollination demonstrated that colonies with high line queens had about twice the amount of pollen stored in the comb than low lines. In addition, high line colonies had significantly more brood. In almonds, we found that high line colonies stored more than twice as much pollen as commercial bees used for comparison. We have now initiated our 1992 alfalfa test with more than 2000 colonies.
Impacts (N/A)
Publications
- NO PUBLICATIONS REPORTED THIS PERIOD.
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