Pesticides, poor nutrition, parasites and diseases work synergistically to contribute to the decline of the honey bee. Heritable sub-lethal behavior/immune effects may also contribute to the decline. Maternal stress is a common source of heritable immune/behavior deficits in many species. A stressed honey bee queen has the potential to pass such deficits on to worker bees. Using a repeated measures design, this study will determine whether the health of worker bee is reduced by a cold stress on the queen by analyzing egg hatch rate and protein content, emergence rate, and adult aggression and immune function for offspring laid before and after the stressor. Results show that queen stress influences egg hatching rate and emergence rate but does not impact egg protein content, adult offspring immune function or aggressive behavior.

The inadvertent spread of exotic pests and pathogens has resulted in devastating losses for bees. The vast majority of bee disease research has focused on a single species of managed bee, the European honey bee (Apis mellifera). More recently, pathogen spillover from managed bees is implicated in the decline of several bumble bee species (Bombus spp.) demonstrating a need to better understand the mechanisms driving disease prevalence in bees, transmission routes, and spillover events. RNA viruses, once considered specific to honey bees, are suspected of spilling over from managed honey bees into wild bumble bee populations. To test this, I collected bees and flowers in the field from areas with and without honey bee apiaries nearby. Prevalence of deformed wing virus (DWV) and black queen cell virus (BQCV) as well as replicating DWV infections in Bombus vagans and B. bimaculatus were highest in bumble bees collected near honey bee apiaries (χ 12 < 6.531, P < 0.05). My results suggest that honey bees are significant contributors of viruses to bumble bees. Flowers have been suspected as bridges in virus transmission among bees. I detected bee viruses on 18% of the flowers collected within honey bee apiaries and detected no virus on flowers in areas without apiaries, thus providing evidence that viruses are transmitted at flowers from infected honey bees. In controlled experiments using captive colonies in flight cages, I found that honey bees leave viruses on flowers but not equally across plant species. My results suggest that there are differences in virus ecology mediated by floral morphology and/or pollinator behavior. No bumble bees became infected in controlled experiments, indicating that virus transmission through plants is a rare event that is likely to require repeated exposure. The few studies examining viruses in bumble bees are generally limited to virus detection, resulting in little understanding of the conditions affecting virus titers. In honeybees, infections may remain latent, capable of replicating under certain conditions, such as immunosuppression induced by pesticide exposure. I tested whether exposure to imidacloprid, a neonicotinoid pesticide, affects virus titers in bumble bees. In previous honey bee studies, imidacloprid exposure increased virus titers. In contrast, I found that bumble bee exposure to imidacloprid decreased BQCV and DWV titers (χ42 < 20.873, p < 0.02). My findings suggest that virus-pesticide interactions are species-specific and results from honey bee studies should not be generalized across other bee species. Having found that honey bees are significant contributors of viruses to wild bees and flowers, I investigated how honey bee management practices affect disease spread and developed recommendations and tools to lesson the risk of spillover events. Honey bee disease may be exacerbated by migratory beekeeping which increases stress and opportunities for disease transmission. I experimentally tested whether migratory conditions contribute to disease spread in honey bees and found negative yet varying effects on bees suggesting that the effects of migratory practices may be ameliorated with rest time between pollination events. State apiary inspection programs are critical to controlling disease spread and reducing the risk of spillover. However, these programs are often resource constrained. I developed and deployed a toolkit that enables state programs to prioritize inspections and provide a platform for beekeeper education. Using novel data collected in Vermont, I discovered several promising avenues for future research and provided realistic recommendations to improve bee health.

Unlike other rosaceous fruit crops such as apple and cherry, commercial raspberry cultivars are largely self-fertile and can set fruit in the absence of pollinators. However, their floral morphology often prevents complete self-pollaintion. Incomplete pollination yields unmarketable small or crumbly fruits. Insect visitation is therefore essential to maximizing raspberry yield. Honey bees are typically used to pollinate commercial raspberry; however, escalating prices for hive rentals coupled with increasing acreage encourage evaluation of other manageable pollinators. Bumble bees (Bombus spp.) and several mason bees (Osmia spp.) are promising raspberry pollinators. Five bee species were evaluated and compared for their single-visit pollination efficacies on raspberry. From this a pollinator effectiveness index was created and an estimation of the minimum number of visits required to maximize fruit set was calculated. This estimation was then experimentally verified. Finally, in an attempt to synchronize their brief activity period with raspberry bloom, winter management options aimed at delaying the emergence of the mason bee, O. lignaria, were investigated. All five bee species proved excellent pollinators of raspberry. None of the alternative manageable species greatly outperformed honey bees. For this reason honey bees remain the most economical and practical option for open-field raspberry pollination. The adoption of alternative manageable bees could still be justified in other production systems, such as high-tunnel or greenhouse grown raspberry, which hamper honey bees’ ability to forage effectively. The pollinator effectiveness score for honey bees suggested that as few as two visits can achieve maximum fruit set. This estimate was confirmed through experimentation on three different red raspberry cultivars. For two of these cultivars, just one visit yielded drupelet counts similar to openly-pollinated flowers. This information can be used to help refine stocking density estimates for honey bees on raspberry. Wintering bees at 0° or -3° C rather than 4° C effectively delayed emergence of O. lignaria by more than a month without any impact on post-winter performance. These results suggest winter storage at near freezing temperatures is a viable management option for the use of O. lignaria with later-blooming crops.

In the honey bee, responsiveness to sucrose correlates with many behavioural parameters such as age of first foraging, foraging role and learning. Sucrose responsiveness can be measured using the proboscis extension response (PER) by applying sucrose solutions of increasing concentrations to the antenna of a bee. We tested whether the biogenic amines octopamine, tyramine and dopamine, and the dopamine receptor agonist 2-amino-6,7-dihydroxy-1,2,3,4-tetrahydronaphthalene (6,7-ADTN) can modulate sucrose responsiveness. The compounds were either injected into the thorax or fed in sucrose solution to compare different methods of application. Injection and feeding of tyramine or octopamine significantly increased sucrose responsiveness. Dopamine decreased sucrose responsiveness when injected into the thorax. Feeding of dopamine had no effect. Injection of 6,7-ADTN into the thorax and feeding of 6,7-ADTN reduced sucrose responsiveness significantly. These data demonstrate that sucrose responsiveness in honey bees can be modulated by biogenic amines, which has far reaching consequences for other types of behaviour in this insect. (C) 2002 Elsevier Science B.V. All rights reserved.

In an earlier project completed in our laboratory a seasonal fluctuation in Phosphoglucomutase (PGM) phenotype frequencies was found, so that the winter bees were almost all heterozygotes and long lived than the summer bees among which homozygotes were significantly at high frequencies at Pgm locus. Same results were obtained in populations of three subspecies, A. m. meda, A. m. caucasica, and A. m. carnica from different climatic regions. In the current study environmental cues related with seasonal change in PGM phenotype frequency was examined along with the correlation between PGM heterozygosity and overwintering success. Cessation of food influx was found to be effective by itself as an environmental cue that causes a sudden and sharp increase in PGM heterozygosity. In addition to that, PGM heterozygosity of the colonies with greater overwintering success was found to be significantly higher than the ones with intermediate or low overwintering success. Benefiting from the previous studies and the results of current study, ethyl oleate was suggested as a chemical signal that functions in the regulation of PGM heterozygosity.

Pollinering är avgörande för ett antal kommersiella grödor samt för olika växtarter i inhemsk vegetation. Honungsbi (Apis mellifera) är en av de viktigaste pollinerarna. I regionen Taranaki i Nya Zeeland finns två aspekter av kommersiell pollinering, dels betalar många biodlare för tillgång till manuka för de antibakteriella egenskaperna honungen får av den, dels betalar frukt- och grönsaksodlare för pollinering av sina grödor. Markägare kan ha ett intresse av att spara manuka på sina marker och få en utkomst från biodlarna genom att dessa betalar för åtkomsten. Studien syftar till att klarlägga de effekter som kommersiell pollinering har i Taranaki-regionen idag och för framtida användning och har utgått från intervjuer med biodlare, frukt- och grönsaksodlare och personer som representerar myndigheterna. Resultaten visar att de ekonomiska effekterna av kommersiell pollinering i dag i Taranaki-regionen är relativt små. Även den kommersiella pollineringens roll för skydd och förvaltning av inhemsk natur undersöks. Kommersiell pollinering kan vara en viktig del inom naturskydd men är samtidigt beroende på om interaktionen med andra bin kan vara skadlig. Pollineringstjänster kan utvecklas dels genom information till markägare och jordbrukare, dels genom marknadsföring från biodlare. / Pollination is crucial for a number of commercial crops as well as for native vegetation. The honey bee (Apis mellifera) is one of the most important pollinators. In the Taranaki region in New Zealand there are two aspects of commercial pollination; beekeepers pay for access to manuka due to the antibacterial quality it gives the honey and orchardists pay for pollination of their crops. Landowners could have an interest in managing the manuka and having beekeepers paying for the access. This study seeks to elucidate the effects that commercial pollination has in Taranaki region, today as well as for future use, through interviews with beekeepers, orchardists and authority personnel. The results show that the effects of commercial pollination in Taranaki are of minor extent. Also the role commercial pollination plays for environmental management is highlighted. Commercial pollination may be a part of nature protection and play a particular role in effort to increase biodiversity, but it is much depending on the interaction with native bees. Pollination services may be developed in different ways including informing landowners and farmers and marketing of beekeepers.

This thesis examines three factors that may influence the change in protein content and size of the brood food glands in honey bees. Effects on the mandibular gland, involved in the production of brood food and in royal jelly, have not been examined in relation to primer pheromones while effects on the hypopharyngeal glands, also involved in the production of brood food, have not been examined in relation to queen mandibular pheromone. This thesis provides preliminary insight into how these pheromones affect the extractable protein content of brood food glands. The first study in this thesis assessed the effects of brood pheromone (BP), queen mandibular pheromone (QMP), and pollen presence on the protein content of hypopharyngeal and mandibular glands of the honey bee. In this study, newly emerged bees were caged for 12 days in one of eight treatments: Queenless state: 1) control (no pollen + no pheromone), 2) pollen, 3) BP, 4) BP + pollen; Queenright state: 1) QMP, 2) QMP + pollen, 3) BP + QMP, 4) BP + QMP + pollen. This study indicated that regardless of pheromone treatment, the most influential factor on gland protein content and size was pollen. The second experiment examined effects of varying pollen dilution on hypopharyngeal and mandibular gland protein content, bee mass, and lipid content of the honey bee. In this experiment, newly emerged bees were caged for 7 days and fed one of five treatments: pollen, 1:1 pollen: cellulose (vol:vol), 1:2 pollen: cellulose (vol:vol); 1:3 pollen: cellulose (vol:vol), and cellulose. This study indicated that bees on the pollen diet were significantly greater than all other diluted diets in measurements of hypopharyngeal gland protein content, lipid content, and mass with significantly less consumption. However, mandibular gland protein content of bees on the pollen diet was significantly greater only from pure cellulose.

During spring, it has been observed that several thousand bees leave their hive, and settle on some object such as a tree branch. Some of the scout bees search for a suitable place where a new hive can be set up, while the rest collect together to form a swarm. Heinrich (J. of Exp. Biology 91 (1981) 25; Science 212 (1981) 565; Scientific American 244:6 (1981) 147) has done some experiments with free and captive swarms. His observations are as follows. (1)The core (centre) temperature is around 35°C irrespective of the ambient temperature. (2)The mantle (outer surface) temperature exceeds the ambient temperature by 2- 3°C, provided the ambient temperature is greater than 20°C. Otherwise the mantle temperature is maintained around 17°C. (3) The temperature gradient vanishes just before take-off of the swarm. The present work attempts to predict temperature profiles in swarms and compare them with the data of Heinrich. A continuum model involving unsteady heat conduction and heat generation within the swarm is used. Heat loss from the outer surface of the swarm by free convection and radiation is accounted for approximately. To simplify the analysis, internal convection within the swarm is neglected. The energy balance equation is solved using the finite element method. The effective thermal conductivity (k) is determined by comparing model predictions with data for a swarm of dead bees. The estimated value of k is 0.20 W/m-K. Both spherical and a non-spherical axisymmetric shapes are considered. Considering axisymmetric swarms of live bees, temperature profiles are obtained using various heat generation functions which are available in literature. The effective thermal conductivity is assumed to be the same as that for the swarm of dead bees. Results based on a modified version of Southwick's heat generation function (The Behavior and Physiology of Bees, pp. 28-47, 1991) are qualitatively in accord with the data. The predicted maximum temperature within the swarm and the temperature at the lower surface of the swarm at the ambient temperature of 5°C are 34°C and 17-20°C, respectively. These are comparable to the measured values of 36°C and 19°C. The predicted maximum temperature within the swarm and the temperature at the lower surface of the swarm at the ambient temperature of 9°C are 36.5°C and 17-22°C, respectively. These are comparable to the measured values of 35°C and 19°C. The predicted oxygen consumption rates are 2.55 ml/g/hr for a swarm of 5284 bees at an ambient temperature Ta = 5°C and 1.15 ml/g/hr for 16,600 bees at Ta = 9°C. These are of the same order as the measured values (2 ml/g/hr for 5284 bees at Ta = 4.4DC and 0.45-0.55 ml/g/hr for 5284 bees at Ta = 10°C). Omholt and Lanvik (J. of Theoretical Biology, 120 (1986) 447) assumed a non-uniform steady state profile and used it to estimate the heat generation function. Using this function in the transient energy balance, it is found that their steady state profile is unstable.

Managed honey bee colonies face severe winter losses in northern climates. In my studies, interactions between genotypes of bees (genetically selected stock and unselected stock) with different levels of resistance and tolerance to varroa mites were assessed under a variety of treatment combinations to quantify effects of queen pheromone, acaricide treatment, wintering method, ventilation condition and pathogen infection on the costs and benefits associated with mite removal and mite-tolerance behaviors. In most of the experiments, mite-resistance caused greater varroa mite mortality within selected stock relative to unselected stock. Artificial and natural sources of queen pheromone caused greater varroa mite mortality within honey bee colonies relative to queenless colonies. While mite resistance had significant benefits, I showed that when producers selected colonies containing some mite resistance traits, it was traits associated with mite-tolerance and not mite-resistance were maintained and contributed to wintering success. Tolerance was effective at two levels of mites as obtained by late autumn treatment of colonies with oxalic but treatment did not improve wintering performance of either stock. Selected stock showed greater colony size, survival and resulted in more viable colonies in spring in comparison to unselected stock with similar initial mite levels (0.16 mites per bee). Selected stock showed greater relative wintering success than unselected stock when wintered indoors than when wintered outdoors but indoor wintering improved colony survival in both stocks relative to outdoor wintering. Carbon dioxide level increased within the winter bee cluster when colonies were maintained under restricted-ventilation (mean 3.82±0.031%, range 0.43-8.44%) and restricted ventilation increased mite mortality by 138% relative to standard-ventilation (mean 1.29±0.031%, range 0.09-5.26%), but restricted-ventilation did not affect bee mortality in comparison to standard-ventilation. In a laboratory study, I showed that Nosema inoculation (with co-infections of N. ceranae and N. apis) suppressed the effectiveness of mite removal behavior within selected bees relative to unselected bees. N. ceranae was more abundant than N. apis. Bees with greater mite removal capacities had higher costs associated with varroa-resistance as indicated by greater bee mortality rates when inoculated with varroa but bee mortality was not affected in Nosema inoculated bees.

ABSTRACT MORPHOMETRIC, MtDNA AND MICROSATELLITE ANALYSIS IN HONEYBEE POPULATIONS (Apis mellifera L.) OF NORTH AND NORTHWEST IRAN Morphometric measurements, mitochondrial DNA analyses and 5 microsatellite loci were used to investigate variation in the honeybee populations of Iran and comparing it with the Turkish populations. Five honeybee populations were sampled from North and west north of Iran. In morphometric aspect of the study 23 characters were measured from left forewings and hindlegs of honey bee samples. The data were analysed by multivariate statistical analyses. By using mtDNA analyses length polymorphism of the intergenic region COI-COII of mitochondrial DNA was studied. After amplification of this region by the polymerase chain reaction, DraI enzyme was used for restriction of amplified region. Results of mtDNA studies show no diversity between four populations and all of them exhibit the same C1 pattern. Five microsatellite loci (A7, A24, A28, A43 and A113) were used in this studies.A high level of average heterozygosity changing between 0.611 and 0.709 was detected in Iranian honey bee populations, and a significant degree of polymorphism was observed. Although Urmia, Sarein and Viladereg populations are similar, Amol population which has located in northern Iran shows a significant difference from other populations. Result obtained form morphometric studies are supporting microsatellite analyses. By comparing data obtained form Iranian honey bee populations with Turkish population (Hakkari), western populations (Urmia, Sarein and Viladereg) are more similar to Hakkari population. It is found Amol is significantly different form other populations and better represents Iranian honeybee.