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254<br />

L.P.S. VAN DER GEEST<br />

gl<strong>and</strong>s (Ongus, 2006). It was possible to isolate the virus <strong>and</strong> to determine the base<br />

sequence of the virus genome. The virus (Varroa destructor virus 1) is a single<br />

str<strong>and</strong>ed RNA genome <strong>and</strong>, based on the base sequence, it was decided that it belongs<br />

to the genus Iflavirus (Ongus et al., 2004). Viruses in this genus belong also to the<br />

picorna-like viruses. The virus is closely related to deformed wing virus, known from<br />

honey bees. The latter virus causes morphological anomalies in wings of bees.<br />

3. DISEASES CAUSED BY BACTERIA<br />

The most widely studied bacterium in invertebrate pathology is Bacillus<br />

thuringiensis. It was first described in 1915 <strong>by</strong> Berliner, who isolated it from soil<br />

samples in the Thuringian Forest in Germany. Simultaneously with spore formation,<br />

a crystalline body is formed in the bacterium. Upon ingestion <strong>by</strong> an insect, this<br />

crystal (δ-endotoxin) falls apart into toxic subunits that may cause paralysis of the<br />

alimentary tract, resulting in the death of the insect. Most varieties (serotypes) of B.<br />

thuringiensis show an effect on larvae of Lepidoptera, but some also on other groups<br />

of insects, e.g. Coleoptera <strong>and</strong> Diptera.<br />

Very comprehensive research has been carried out on B. thuringiensis that has<br />

resulted in the development of several commercial preparations that are mainly used<br />

against lepidopterous pests. Also, the gene encoding for the crystalline toxic body<br />

has been isolated <strong>and</strong> transferred into crop plants, e.g. corn <strong>and</strong> cotton, making these<br />

crops resistant towards a number of lepidopterous pests. Several serotypes of B.<br />

thuringiensis produce in addition an exotoxin, the β-exotoxin, named thuringiensin.<br />

This exotoxin is excreted <strong>by</strong> the bacterium into the culture medium. It has a<br />

nucleotide-like structure <strong>and</strong> inhibits DNA-dependent RNA polymerase. This results<br />

in a blockage of mitosis. When thuringiensin is applied to young holometabolous<br />

insects, morphological deformations may occur in the adult stage.<br />

Field applications of thuringiensin were successful against the citrus red mite P.<br />

citri (Hall, Hunter, & Arakawa, 1971) <strong>and</strong> Tetranychus pacificus (Hoy & Ouyang,<br />

1987). Later, Royalty, Hall, <strong>and</strong> Taylor (1990) conducted experiments <strong>by</strong> testing two<br />

different formulations of thuringiensin against the twospotted spider mite T. urticae.<br />

The results indicated that thuringiensin might be a potential acaricide. In particular<br />

young instars are susceptible, since these have a high growth rate. Various<br />

physiological processes in young <strong>org</strong>anisms require higher RNA synthesis than in<br />

the older slower growing stages. A major drawback is that thuringiensin is toxic for<br />

a wide range of <strong>org</strong>anisms. Not only are spider mites affected, but also beneficial<br />

mites, such as Phytoseiulus persimilis: oviposition starts to decline after 2 days <strong>and</strong><br />

ceases completely after 3–4 days in both predator <strong>and</strong> spider mite T. urticae (Guo,<br />

Zuo, Zhao, Wang, & Jiang, 1993). The chemical is apparently a nonselective<br />

acaricide that should not be used in combination with predatory mites.<br />

The spore-crystal complex of B. thuringiensis has been tested on spider mites<br />

<strong>by</strong> Krieg (1972), but no mortality was observed. However, Chapman <strong>and</strong> Hoy<br />

(1991) conducted experiments in which T. urticae <strong>and</strong> Metaseiulus occidentalis<br />

were treated with a commercial preparation of B. thuringiensis var. tenebrionis. This<br />

variety of B. thuringiensis shows an effect on beetles <strong>and</strong> is recommended for use<br />

against the Colorado Potato Beetle, Leptinotarsa decemlineata. No effect was noted

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