蜜蜂蜂王质量及其性状表现影响因素研究进展

作为重要的传粉昆虫,蜜蜂蜂群损失现象受到广泛关注。研究表明,蜂王问题是导致蜂群损失的主要因素之一。蜂王是蜂群中唯一雌性生殖器官发育完全的个体,是维持蜂群存续的关键。蜂王质量决定了蜂群的群势以及生产性能,对蜂群的发展和存活至关重要。本文详细介绍了表征蜂王质量的相关指标及其在蜂王选育过程中的应用,深入论述了幼虫日龄、营养、交尾、环境温度、病虫害以及农药等相关因素对蜂王质量及其性状表现的影响,以期为蜂王的培育和使用以及蜜蜂资源的保护和利用提供参考。     

Maternal influence on the acceptance of virgin queens introduced into Africanized honey bee (Apis mellifera) colonies

The oviposition potential of honey bee queens decreases with age, therefore it is important to replace old queens with younger ones on a periodic basis. However, queen replacement is problematic, especially in Africanized honey bee colonies, since many introduced queens are not accepted, and virgin queens are less easily accepted than are mated queens. We assessed the influence of genetic origin (queen mother) on the acceptance of queens, when they were introduced as virgins into Africanized honey bee colonies. For this purpose, 12 daughter queens from each of 11 mother queens with no degree of kinship among themselves were introduced. Introductions were made monthly, for 12 months, though the winter months of June and July were not included, as there is little brood and drones are rare in winter. There was some seasonal variation in the acceptance rates; generally there was greater acceptance in months with good honey flows. However, the acceptance of introduced queens was influenced by their origin. The rate of acceptance of daughter queens from the 11 different mother queens varied significantly, ranging from 33 to 75%. There appears to be a genetic influence of the mother queen on the introduced queen acceptance rate.     

The effects of honey bee (Apis mellifera L.) queen reproductive potential on colony growth

Reproduction in species of eusocial insects is monopolized by one or a few individuals, while the remaining colony tasks are performed by the worker caste. This reproductive division of labor is exemplified by honey bees (Apis mellifera L.), in which a single, polyandrous queen is the sole colony member that lays fertilized eggs. Previous work has revealed that the developmental fate of honey bee queens is highly plastic, with queens raised from younger worker larvae exhibiting higher measures in several aspects of reproductive potential compared to queens raised from older worker larvae. Here, we investigated the effects of queen reproductive potential (“quality”) on the growth and winter survival of newly established honey bee colonies. We did so by comparing the growth of colonies headed by “high-quality” queens (i.e., those raised from young worker larvae, which are more queen-like morphologically) to those headed by “low-quality” queens (i.e., those raised from older worker larvae, which are more worker-like morphologically). We confirmed that queens reared from young worker larvae were significantly larger in size than queens reared from old worker larvae. We also found a significant positive effect of queen grafting age on a colony’s production of worker comb, drone comb, and stored food (honey and pollen), although we did not find a statistically significant difference in the production of worker and drone brood, worker population, and colony weight. Our results provide evidence that in honey bees, queen developmental plasticity influences several important measures of colony fitness. Thus, the present study supports the idea that a honey bee colony can be viewed (at least in part) as the expanded phenotype of its queen, and thus selection acting predominantly at the colony level can be congruent with that at the individual level.     

Low natural levels of Nosema ceranae in Apis mellifera queens

Queens are the primary female reproductive individuals in honey bee colonies and, while they are generally free from Nosema ceranae infection, they are nevertheless susceptible. We sought to determine whether queens are naturally infected by N. ceranae, as these infections could be a factor in the rapid spread of this parasite. Queens were analyzed using real-time PCR and included larval queens, newly emerged, and older mated queens. Overall, we found that all tissues we examined were infected with N. ceranae at low levels but no samples were infected with Nosema apis. The infection of the ovaries and spermatheca suggests the possibility of vertical transmission of N. ceranae.     

Pathological effects of the microsporidium Nosema ceranae on honey bee queen physiology (Apis mellifera)

Nosema ceranae, a microsporidian parasite originally described in the Asian honey bee Apis cerana, has recently been found to be cross-infective and to also parasitize the European honey bee Apis mellifera. Since this discovery, many studies have attempted to characterize the impact of this parasite in A. mellifera honey bees. Nosema species can infect all colony members, workers, drones and queens, but the pathological effects of this microsporidium has been mainly investigated in workers, despite the prime importance of the queen, who monopolizes the reproduction and regulates the cohesion of the society via pheromones. We therefore analyzed the impact of N. ceranae on queen physiology. We found that infection by N. ceranae did not affect the fat body content (an indicator of energy stores) but did alter the vitellogenin titer (an indicator of fertility and longevity), the total antioxidant capacity and the queen mandibular pheromones, which surprisingly were all significantly increased in Nosema-infected queens. Thus, such physiological changes may impact queen health, leading to changes in pheromone production, that could explain Nosema-induced supersedure (queen replacement).     

The impact of caging the queens during the flow season on some biological activities of honeybee colonies

This study was achieved in a private apiary located in a banana farm in Sa El Hagar, Basioun, Gharbia, Egypt from August 15, 2019 to May 25, 2020, including the banana (Musa sp., Musaceae) flow season (August and September) and extend to Egyptian clover (Trifolium alexandrinum L., Fabaceae) flow season (May). The study aimed to evaluate the effect of confining the queen during the banana flow season on the brood rearing, honey yield, and activation of worker's ovaries. Also, we determined the negative impact of caging the queen during the banana flow season on the activity of the colony in brood rearing, storing pollen, and honey yield after releasing the queen on 5 October, extending to the next flow season in May. The obtained results showed that the honeybee colonies with the caged queen produced significantly more honey yield and less brood production than the free queen ones during the banana flow season. Also, the caging of the queen did not affect the colony strength after releasing the queen despite the partial development of the ovaries of some workers, but they did not lay eggs. In addition, releasing the queens suppressed the ovaries of the laying workers. It can be concluded that caging the queen during the banana flow season helps the colonies to produce more honey yield without effect on the colony strength after releasing the queen despite the ovaries development of few workers without egg-laying.     

Maternity of emergency queens in the Cape honey bee,Apis mellifera capensis

During reproductive swarming, some workers of the Cape honey bee, Apis mellifera capensis, lay eggs in queen cells, many of which are reared to maturity. However, it is unknown if workers are able to lay in queen cells immediately after queen loss during an episode of emergency queen rearing. In this study we experimentally de‐queened colonies and determined the maternity of larvae and pupae that were reared as queens. This allowed us to determine how soon after queen loss workers contribute to the production of new queens. We were further interested to see if workers would preferentially raise new queens from queen‐laid brood if this was introduced later. We performed our manipulations in two different settings: an apiary setting where colonies were situated close together and a more natural situation in which the colonies were well separated. This allowed us to determine how the vicinity of other colonies affects the presence of parasites. We found that workers do indeed contribute to queen cell production immediately after the loss of their queen, thus demonstrating that some workers either have activated ovaries even when their colony has a queen or are able to activate their ovaries extremely rapidly. Queen‐laid brood introduced days after queen loss was ignored, showing that workers do not prefer to raise new queens from queen brood when given a choice. We also detected non‐natal parasitism of queen cells in both settings. We therefore conclude that some A. m. capensis genotypes specialize in parasitizing queen cells.     

Evidence for emerging parasites and pathogens influencing outbreaks of stress-related diseases like chalkbrood

In agriculture, honey bees play a critical role as commercial pollinators of crop monocultures which depend on insect pollination. Hence, the demise of honey bee colonies in Europe, USA, and Asia caused much concern and initiated many studies and research programmes aiming at elucidating the factors negatively affecting honey bee health and survival. Most of these studies look at individual factors related to colony losses. In contrast, we here present our data on the interaction of pathogens and parasites in honey bee colonies. We performed a longitudinal cohort study over 6 years by closely monitoring 220 honey bee colonies kept in 22 apiaries (ten randomly selected colonies per apiary). Observed winter colony losses varied between 4.8% and 22.4%; lost colonies were replaced to ensure a constant number of monitored colonies over the study period. Data on mite infestation levels, infection with viruses, Nosema apis and Nosema ceranae, and recorded outbreaks of chalkbrood were continuously collected. We now provide statistical evidence (i) that Varroa destructor infestation in summer is related to DWV infections in autumn, (ii) that V. destructor infestation in autumn is related to N. apis infection in the following spring, and most importantly (iii) that chalkbrood outbreaks in summer are related to N. ceranae infection in the preceding spring and to V. destructor infestation in the same season. These highly significant links between emerging parasites/pathogens and established pathogens need further experimental proof but they already illustrate the complexity of the host–pathogen-interactions in honey bee colonies.     

Queen replacement in African and European honey bee colonies with and without afterswarms

We examined the dynamics of the queen replacement process in African and European colonies that did and did not produce afterswarms. In colonies without afterswarms, the queen replacement process was completed in 24–48 hours, the first-emerging virgin queen (VQ) typically inherited the natal nest even if multiple queens emerged, workers performed few vibration signals on emerged queens, and all signaling activity was directed toward early emerging VQs. In contrast, if colonies did produce afterswarms, the queen replacement process required 5–6 days, there was no advantage for first-emerging queens, vibration rates on emerged queens were 25 times greater, and signaling activity was directed toward all VQs. Although vibration signal activity was more pronounced in colonies with afterswarms, the signal was consistently associated with increased VQ survival under all conditions. These trends were exhibited similarly in the African and European colonies, suggesting that they have broad applicability to queen-replacement decisions over a range of environmental and racial conditions. However, the African and European colonies differed in the total number of queens involved in the elimination process and the relative importance of queen duels and pre-emergence destruction under the different reproductive strategies. Taken together, our results suggest that worker behavior is a major determinant for the outcome of queen replacement, either through reduced interactions that allow first-emerged queens to rapidly eliminate rivals, or through increased use of interactions such as the vibration signal, which may allow workers to influence the ultimate fate of each emerged VQ. We discuss the possibility that these behavior patterns may reflect the roles of cooperation and conflict in shaping honey bee reproductive decisions. Received 8 May 2007; revised 7 November 2007; accepted 20 November 2007.     

Effects of insemination quantity on honey bee queen physiology

Mating has profound effects on the physiology and behavior of female insects, and in honey bee (Apis mellifera) queens, these changes are permanent. Queens mate with multiple males during a brief period in their early adult lives, and shortly thereafter they initiate egg-laying. Furthermore, the pheromone profiles of mated queens differ from those of virgins, and these pheromones regulate many different aspects of worker behavior and colony organization. While it is clear that mating causes dramatic changes in queens, it is unclear if mating number has more subtle effects on queen physiology or queen-worker interactions; indeed, the effect of multiple matings on female insect physiology has not been broadly addressed. Because it is not possible to control the natural mating behavior of queens, we used instrumental insemination and compared queens inseminated with semen from either a single drone (single-drone inseminated, or SDI) or 10 drones (multi-drone inseminated, or MDI). We used observation hives to monitor attraction of workers to SDI or MDI queens in colonies, and cage studies to monitor the attraction of workers to virgin, SDI, and MDI queen mandibular gland extracts (the main source of queen pheromone). The chemical profiles of the mandibular glands of virgin, SDI, and MDI queens were characterized using GC-MS. Finally, we measured brain expression levels in SDI and MDI queens of a gene associated with phototaxis in worker honey bees (Amfor). Here, we demonstrate for the first time that insemination quantity significantly affects mandibular gland chemical profiles, queen-worker interactions, and brain gene expression. Further research will be necessary to elucidate the mechanistic bases for these effects: insemination volume, sperm and seminal protein quantity, and genetic diversity of the sperm may all be important factors contributing to this profound change in honey bee queen physiology, queen behavior, and social interactions in the colony.     

Patriline differences in emergency queen rearing in the honey bee, <Emphasis Type="Italic">Apis mellifera</Emphasis>

Summary In the polyandrous honey bee, Apis mellifera, workers can potentially increase their inclusive fitness by rearing full-sister queens. If the mother queen dies suddenly, workers feed a few larvae in worker cells with royal jelly and rear them into queens (emergency queen rearing). Using DNA microsatellite markers we determined the patriline of emergency queens reared in two colonies headed by naturally-mated queens before being made queenless. We found that some patrilines were reared more than others in one colony, but not in the other. These differences between colonies suggest that selective rearing is not always present and this might explain the mixed results of previous nepotism studies in the honey bee.Received 10 February 2003; revised 7 March 2003; accepted 17 March 2003.     

Three QTL in the honey bee Apis mellifera L. suppress reproduction of the parasitic mite Varroa destructor

Varroa destructor is a highly virulent ectoparasitic mite of the honey bee Apis mellifera and a major cause of colony losses for global apiculture. Typically, chemical treatment is essential to control the parasite population in the honey bee colony. Nevertheless a few honey bee populations survive mite infestation without any treatment. We used one such Varroa mite tolerant honey bee lineage from the island of Gotland, Sweden, to identify quantitative trait loci (QTL) controlling reduced mite reproduction. We crossed a queen from this tolerant population with drones from susceptible colonies to rear hybrid queens. Two hybrid queens were used to produce a mapping population of haploid drones. We discriminated drone pupae with and without mite reproduction, and screened the genome for potential QTL using a total of 216 heterozygous microsatellite markers in a bulk segregant analysis. Subsequently, we fine mapped three candidate target regions on chromosomes 4, 7, and 9. Although the individual effect of these three QTL was found to be relatively small, the set of all three had significant impact on suppression of V. destructor reproduction by epistasis. Although it is in principle possible to use these loci for marker-assisted selection, the strong epistatic effects between the three loci complicate selective breeding programs with the Gotland Varroa tolerant honey bee stock.     

Twenty-five-year study of Nosema spp. in honey bees (Apis mellifera) in Serbia

A total of 7386 samples of adult honey bees from different areas of Serbia (fifteen regions and 79 municipalities) were selected for light microscopy analysis for Nosema species during 1992–2017. A selection of honey bee samples from colonies positive for microsporidian spores during 2009–2011, 2015 and 2017 were then subjected to molecular diagnosis by multiplex PCR using specific primers for a region of the 16S rRNA gene of Nosema species. The prevalence of microsporidian spore-positive bee colonies ranged between 14.4% in 2013 and 65.4% in 1992. PCR results show that Nosema ceranae is not the only Nosema species to infect honey bees in Serbia. Mixed N. apis/N. ceranae infections were detected in the two honey bee samples examined by mPCR during 2017. The beekeeping management of disease prevention, such as replacement of combs and queens and hygienic handling of colonies are useful in the prevention of Nosema infection.     

Management of Braula orientalis Örösi (Diptera: Braulidae) in honeybee colonies with tobacco smoke under semiarid conditions

In the present study, bee colonies were smoked with tobacco smoke in order to evaluate the monthly changes in the numbers of worker bees, the infestation rates of worker bees and queens with bee lice, and the annual average honey production per colony. In July of each year, 12 colonies were smoked with tobacco smoke; the remaining hives not smoked with tobacco smoke served as the control. The results indicated that the applications of tobacco smoke during July gave rise to an impressive reduction in the Braula infestation rate on workers (below 1.8%) and reduced the amount of bee lice on the queen to zero throughout the 2–3 months following smoke treatment. In the colonies not treated with smoke, the Braula infestation rates on worker bees started to increase in May and continued to increase constantly during the rest of year, reaching maximum infestation rates of 28.2% and 33.8% in December, with an average of 15 and 17 lice per queen in November in the first and second years, respectively. Worker bee populations peaked in April and July of each year in both treatments. The average honey production per colony was significantly higher in the colonies treated with smoke than those that were not for the first and second years. In conclusion, early summer months may be the crucial time to smoke the colonies with tobacco smoke in order to keep bee lice at low levels for the remaining seasons.     

Queen substances from the abdomen of the honey bee queen

Summary The secretion of the mandibular glands of a honey bee queen enables the worker bees to react to the presence of their queen. Extirpating the mandibular glands of the queen does not prevent that she is accepted by her colony. Hitherto this was attributed to contamination of the queen's body by mandibular gland substances during or preceding the extirpation. When, however, these glands are extirpated before they have secreted any material and the queens are inseminated artificially, the colonies still accept these queens. A normal-sized retinue, the absence of emergency cell building and the absence of activation of the worker's ovaries indicate that such a queen is still able to maintain her social position. This supports Verheijen-Voogd's (1959) conclusion that the queen's influence on her workers has a behavioural basis (chemoreception) rather than a biochemical one.Laboratory experiments reveal that apart from the mandibular gland substances other queen pheromones are produced in glands on the abdomen, most probably in the glands described by Renner and Baumann (1964).     

Effect of a lactic acid treatment during winter in temperate climate upon Varroa jacobsoni Oud. and the bee (Apis mellifera L.) colony

Three groups of bee colonies were treated with lactic acid, the pesticide Perizin or lactic acid and Perizin in order to validate the applicability of lactic acid in Varroa mite control. The lactic acid treatment was conducted during winter. Eight ml of lactic acid (15%) per comb side were applied with a dosage gun. The treatment was highly efficient and 94.2%–99.8% of the mites in a colony were killed. Due to precise dosage the lactic acid treatment caused less bee mortality than a treatment with the pesticide Perizin. A lactic acid treatment at-0.2°C caused bee mortality comparable to a Perizin treatment. The number of queen losses after lactic acid treatment and after Perizin treatment was comparable. The number of bees, the size of the brood area, the amount of stored honey and Nosema infestation rates were not significantly different in lactic acid treated colonies and Perizin treated colonies in spring after treatment.     

The inhibiting effect of the queen bee (Apis mellifera L.) foot-print pheromone on the construction of swarming queen cups

Experiments were conducted to determine the rôle of population density of queenright honey bee colonies, and that of the queen bee pheromonal secretions, on the induction and inhibition of swarming queen cup construction during swarming and non-swarming seasons. Construction of queen cups was induced experimentally in overcrowded queenright colonies, during winter, which is a non-swarming season. This construction was induced by high population density of bee workers: above a threshold of 2.3 bee workers/ml there was a relationship between the number of cups constructed and the colony density. During the swarming season a relationship was established between the free volume of a hive (population density) and the number of queen cups constructed: 1.5 cups in a colony that occupied 80,960 ml, compared with 77 cups in a colony hived within a volume of 20, 240 ml. Observations of the queen's movements upon combs in colonies of high and normal population densities showed that in an overcrowded colony the queen bee was almost absent from the bottom edges of the comb, where queen swarming cups and cells are constructed. The tarsal glands of queens are located in the fifth tarsomere and the glandular oily secretion is deposited by the foot-pads upon the combs surface. The rate of secretion by the queen's tarsal glands was about 13 times higher than by those of the workers. A bioassay for testing the inhibitory effects of the queen's glandular extracts on the construction of queen cups was developed. It was based on increasing worker bee population densities, and can be used effectively throughout the year in a subtropical climate.The application of tarsal and mandibular glands' secretion to comb bottom edges in overcrowded colonies (bioassay) caused the inhibition of queen cup construction. None of these two secretions affected construction of these cups when applied separately. We presume that due to colony overcrowding the queen bee is unable to deposit the non-volatile secretions from tarsal glands along the comb edges and that the deficiency of the foot-print pheromone triggers the construction of swarming cups along the non-inhibited areas.     

The Behavior of Honey Bees (Apis mellifera ligustica) during Queen Duels

Conflict is rare among the members of a highly cooperative society such as a honey bee colony. However, conflict within a colony increases drastically during colony reproduction ('swarming') when newly produced queens fight each other until only one queen remains in the nest. This study describes the behavior of queens and workers during naturally occurring queen combat. The duels of five pairs of queens were observed in three observation colonies. A typical duel is described qualitatively and the events of all five duels are described quantitatively. Several aspects of duels that are of particular interest are examined in detail, including the behavior of queens near capped queen cells, worker aggression toward queens, queen tooting, and the relation of queen and worker behavior to the outcome of the duel. The results of this investigation serve as a foundation for rigorous tests of hypotheses regarding the adaptive significance of queen and worker behavior during queen combat. The results presented suggest that: young queens patrol queen cells to kill rival queens while they are vulnerable; workers aggress queens to prevent them from destroying queen cells; queens toot to inhibit worker aggression; workers immobilize queens to make them easy targets for rival queens; and queens eject hind-gut contents to cause their rival to be immobilized by the workers.     

Effects of social conditions on Juvenile Hormone mediated reproductive development in Bombus terrestris workers

Abstract. During the annual life cycle of the bumble bee Bombus terrestris (L.) colony, there is a stage characterized by worker reproduction in the presence of the queen. It has been proposed that this is a result of a decrease in queen inhibition. This hypothesis was examined by studying the effects of queens taken from colonies at different stages of development on several aspects of worker physiology and behaviour: rates of Juvenile Hormone (JH) release in vitro , ovary development, and behaviour associated with reproduction. After optimizing and validating the radiochemical assay for JH release for bumble bee workers, we found that queenless workers had significantly more developed ovaries and higher rates of release of JH than did queenright workers, confirming and extending previous findings that suggest that bumblebee ovarian development is under JH control. Mated queens, separated from their colony and brood, can have the same inhibitory effect on the reproductive development of callow workers. In contrast, workers confined with virgin queens or in queenless groups demonstrated a significantly higher rate of release of JH, overt aggression and threatening behaviours. However, there were no differences in rates of release of JH between workers confined in groups in the laboratory with queens taken from colonies either before or after the onset of worker reproduction. Furthermore, overt aggression and threatening behaviours were similar and low in both types of groups. These results gave no support to the hypothesis that a decrease in queen inhibition is associated with the onset of worker reproduction. We also show that young workers reared in colonies either before or after worker reproduction occurs, or in queenless colonies, all demonstrated similar, low rates of release of JH. These results suggest that older workers may inhibit the corpora allata of younger workers in queenless colonies.