The costs and benefits of resistance and tolerance behaviors against Varroa mite (Varroa destructor Anderson and Trueman) in honey bee (Apis mellifera L.)
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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.