In recent years, honeybees (Apis mellifera) have been strangely disappearing from their hives, and strong colonies have suddenly become weak and died. The precise aetiology underlying the disappearance of the bees remains a mystery. However, during the same period, Nosema ceranae, a microsporidium of the Asian bee Apis cerana, seems to have colonized A. mellifera, and it's now frequently detected all over the world in both healthy and weak honeybee colonies. For first time, we show that natural N. ceranae infection can cause the sudden collapse of bee colonies, establishing a direct correlation between N. ceranae infection and the death of honeybee colonies under field conditions. Signs of colony weakness were not evident until the queen could no longer replace the loss of the infected bees. The long asymptomatic incubation period can explain the absence of evident symptoms prior to colony collapse. Furthermore, our results demonstrate that healthy colonies near to an infected one can also become infected, and that N. ceranae infection can be controlled with a specific antibiotic, fumagillin. Moreover, the administration of 120 mg of fumagillin has proven to eliminate the infection, but it cannot avoid reinfection after 6 months. We provide Koch's postulates between N. ceranae infection and a syndrome with a long incubation period involving continuous death of adult bees, non-stop brood rearing by the bees and colony loss in winter or early spring despite the presence of sufficient remaining pollen and honey.

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How natural infection by Nosema ceranae causes honeybee colony collapse

The Western honey bee, Apis mellifera, is the most important animal pollinator in agriculture worldwide providing more than 90% of the commercial pollination services. Due to the development in agriculture the demands for honey bee pollination are steadily increasing stressing the pollination capacity of the global managed honey bee population. Hence, the long-term decline of managed honey bee hives in Europe and North-America is of great concern and stimulated intensive research into the possible factors presumably causing honey bee colony collapse. We here present a four-year study involving more than 1200 bee colonies from about 120 apiaries which were monitored for the entire study period. Bee samples were collected twice a year to analyze various pathogenic factors including the ectoparasitic mite Varroa destructor, fungi (Nosema spec., Ascosphaera apis), the bacterium Paenibacillus larvae, and several viruses. Data on environmental factors, beekeeping management practice, and pesticides were also collected. All data were statistically analyzed in respect to the overwintering mortality of the colonies. We can demonstrate for several factors that they are significantly related to the observed winter losses of the monitored honey bee colonies: (i) high varroa infestation level, (ii) infection with deformed wing virus (DWV) and acute bee paralysis virus (ABPV) in autumn, (iii) queen age, and (iv) weakness of the colonies in autumn. No effects could be observed for Nosema spec. or pesticides. The implications of these findings will be discussed.

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The German bee monitoring project: a long term study to understand periodically high winter losses of honey bee colonies

Viruses are significant threats to the health and well‐being of the honey bee, Apis mellifera . To alleviate the threats posed by these invasive organisms, a better understanding of bee viral infections will be of crucial importance in developing effective and environmentally benign disease control strategies. Although knowledge of honey bee viruses has been accumulated considerably in the past three decades, a comprehensive review to compile the various aspects of bee viruses at the molecular level has not been reported. This chapter summarizes recent progress in the understanding of the morphology, genome organization, transmission, epidemiology, and pathogenesis of honey bee viruses as well as their interactions with their honey bee hosts. The future prospects of research of honey bee viruses are also discussed in detail. The chapter has been designed to provide researchers in the field with updated information about honey bee viruses and to serve as a starting point for future research.

Honey bee viruses.

A survey of six bee viruses on a large geographic scale was undertaken by using seemingly healthy bee colonies and the PCR technique. Samples of adult bees and pupae were collected from 36 apiaries in the spring, summer, and autumn during 2002. Varroa destructor samples were collected at the end of summer following acaricide treatment. In adult bees, during the year deformed wing virus (DWV) was found at least once in 97% of the apiaries, sacbrood virus (SBV) was found in 86% of the apiaries, chronic bee paralysis virus (CBPV) was found in 28% of the apiaries, acute bee paralysis virus (ABPV) was found in 58% of the apiaries, black queen cell virus (BQCV) was found in 86% of the apiaries, and Kashmir bee virus (KBV) was found in 17% of the apiaries. For pupae, the following frequencies were obtained: DWV, 94% of the apiaries; SBV, 80% of the apiaries; CBPV, none of the apiaries; ABPV, 23% of the apiaries; BQCV, 23% of the apiaries; and KBV, 6% of the apiaries. In Varroa samples, the following four viruses were identified: DWV (100% of the apiaries), SBV (45% of the apiaries), ABPV (36% of the apiaries), and KBV (5% of the apiaries). The latter findings support the putative role of mites in transmitting these viruses. Taken together, these data indicate that bee virus infections occur persistently in bee populations despite the lack of clinical signs, suggesting that colony disease outbreaks might result from environmental factors that lead to activation of viral replication in bees.

Prevalence and Seasonal Variations of Six Bee Viruses in Apis mellifera L. and Varroa destructor Mite Populations in France

The hive of the honey bee is a suitable habitat for diverse mites (Acari), including nonparasitic, omnivorous, and pollen-feeding species, and para- sites. The biology and damage of the three main pest species Acarapis woodi, Varroa jacobsoni, and Tropilaelaps clareae is reviewed, along with detection and control methods. The hypothesis that Acarapis woodiis a recently evolved species is rejected. Mite-associated bee pathologies (mostly viral) also cause increasing losses to apiaries. Future studies on bee mites are beset by three main problems: (a) The recent discovery of several new honey bee species and new bee-parasitizing mite species (along with the probability that several species are masquerading under the name Varroa jacob- soni) may bring about new bee-mite associations and increase damage to beekeeping; (b) methods for studying bee pathologies caused by viruses are still largely lacking; (c) few bee- and consumer-friendly methods for controlling bee mites in large apiaries are available.

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Parasitic mites of honey bees: life history, implications, and impact.

SUMMARYMolecular protocols were applied to the diagnosis of Kashmir bee virus (KBV) infection in the honey bee, Apis mellifera. Procedures based on either serology (chemiluminescent Western blotting) or nucleic acid amplification (RT-PCR, reverse transcription-polymerase chain reaction) proved to be very effective, although it was possible in some circumstances to detect viral infection simply by Coomassie blue staining of SDS-PAGE gels. Under laboratory conditions, KBV infections were readily distinguished from those induced by acute paralysis virus, a closely related isolate, by using monospecific antisera raised against the putative picornavirus-like VP4 polypeptide. Since only a very small amount of material was required for either Western blotting or RT-PCR, a single bee could in theory be examined for the presence of a variety of different pathogens. PCR primers based on sequences from the viral RNA polymerase gene amplified a 417 bp amplicon from KBV-infected, but not healthy, pupae. The same prime...

Xue-Ren Shen

Papers

Molecular diagnosis of Kashmir bee virus infection