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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

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