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MITE PATHOGENS IN IPM<br />
253<br />
Zenner, 2005). It is not within the scope of this article to describe these viruses<br />
comprehensively, as they are considered to be plant, resp. vertebrate viruses.<br />
Several viruses are known from the honeybee (Apis mellifera), some of which<br />
are associated with the varroa mite Varroa jacobsoni <strong>and</strong> the honeybee tracheal mite<br />
(HBTM) Acarapis woodi (Sammataro, Gerson, & Needham 2000). These viruses<br />
may always be present in the bee, either in a latent or in an unapparent form.<br />
Wounds inflicted <strong>by</strong> the mites may activate the viruses in the bee. These viruses are<br />
probably not capable to infect the varroa mite or the HBTM, but the presence of<br />
these mites affects the incidence of virus disease in honey bees. The varroa mite may<br />
also play a role in the transmission of Bee Kashmir Virus (KBV), although the virus<br />
was already known from bee colonies before Apis mellifera colonies were infested<br />
<strong>by</strong> V. destructor. The presence of the virus in the mite has been demonstrated <strong>by</strong><br />
several authors (e.g. Chen, Pettis, Evans, Kramer, & Feldlaufer, 2004). Virus-free<br />
mites may become infected <strong>by</strong> coinhabiting in the same cell as virus-infected mites.<br />
Whether transmission to honey bees occurs mechanically or biologically is a<br />
question that has not yet been conclusively solved. Shen, Yang, Cox-Foster, <strong>and</strong> Cui<br />
(2005) suggested that varroa mites cause suppression of the immune system of the<br />
honey been which leads to activation of latent virus infections.<br />
Liu (1991) found virus-like particles in a sample of HBTM that originated from<br />
Scotl<strong>and</strong>, but no such particles were found in HBTM samples from California.<br />
Tissues of affected mites showed extensive lysis, while most cells were tightly<br />
packed with virus-like particles. The virions are 27–30 nm in size <strong>and</strong> are arranged<br />
in paracrystalline arrays forming hexagonal patterns. The ultrastructural morphology<br />
of the particles indicates that we may be dealing with a picorna-like virus. Such<br />
viruses resemble picornaviruses that are found in vertebrates. Picorna-like viruses<br />
are also known from honeybees, but on the basis of histopathological studies it was<br />
assumed that the virus found in HBTM is not derived from honeybees, but actually<br />
multiplies inside the mite.<br />
Kleespies, Radtke, <strong>and</strong> Bienefield (2000) performed a search for diseases in<br />
varroa mites in parasitized bee colonies. They found mites with characteristic<br />
internal black-colored changes of the gut <strong>and</strong> the fat body. On living adult bees,<br />
3.6% of the mites showed this anomaly, in brood cells, even 8% of the juvenile<br />
mites were affected. Disease incidence <strong>and</strong> intensity of the symptoms can be<br />
enhanced <strong>by</strong> changes in environmental conditions, such as deficiency of bee brood,<br />
deficiency of pollen, abnormal brood temperature <strong>and</strong> death of the host. The authors<br />
found that longevity of black-colored mites was reduced <strong>by</strong> 43%. Cytopathological<br />
studies showed the presence of a large number of spherical virus-like particles,<br />
especially in the nuclei of fatbody <strong>and</strong> muscle tissue. The particles measure<br />
approximately 27–60 nm in diameter <strong>and</strong> were very similar to the particles found <strong>by</strong><br />
Liu (1991) in HBTM. Per os infection experiments with extracts of fatbody tissue<br />
derived from symptomatic mites were unsuccessful.<br />
Ongus et al. (2004) detected virus-like particles in varroa mites that were<br />
collected in bee hives in The Netherl<strong>and</strong>s. The virions were mainly present in the<br />
cytoplasm of mite tissue <strong>and</strong> resemble the virus-like particles found <strong>by</strong> Kleespies<br />
et al. (2000). Immunochemical studies revealed that the virus was localized in the<br />
abdominal part of the alimentary tract <strong>and</strong> in the gastric caeca, but not in the salivary