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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