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ALLELOCHEMICALS : MANAGEMENT OF PLANT-PARASITIC NEMATODES 23<br />

group of glucosinolate compounds extracted from rapeseed exhibited various levels<br />

of nematicidal activity depending on concentration and length of exposure (Lazzeri<br />

et al., 1993).<br />

Later studies by Borek et al. (1995) investigated the persistence of glucosinolatederived<br />

allyl isothiocyanate and allylnitrile in six soils. They found that the two<br />

compounds differed with respect to the temperature, moisture conditions, and soil<br />

physical conditions that effected their transformation in soil, and that both compounds<br />

dissipated from soil at relatively rapid rates. Donkin et al. (1995) studied the toxicity<br />

of glucosinolates and their enzymatic breakdown products to Caenorhabditis elegans.<br />

They found that allyl isothyocyanate, one of the decomposition products of the<br />

glucosinolate sinigrin, was three times more toxic to the nematode C. elegans than<br />

corresponding glucosinolate itself.<br />

4.2. Benzaldehyde (benzoic aldehyde)<br />

Benzaldehyde occurs in seeds of bitter almond (Prunus dulcis). It is found naturally<br />

in several cyanogenic glucosides and is used in food and fragrances for its almondlike<br />

aroma and flavor (Harborne and Baxter, 1993). The value of benzaldehyde as a<br />

fungicide is well established (Flor, 1926), with the nematicidal activity more recently<br />

demonstrated. Benzaldehyde reduced populations of M. incognita in field microplots<br />

with no phytotoxicity to cotton at 0.18 -2.14 ml/kg soil (Bauske et al., 1994). The<br />

combination of chitin and benzaldehyde added to peat-based potting mix improved<br />

tomato transplant growth and reduced galling by M. incognita in greenhouse trials<br />

(Kokalis-Burelle et al., 2002). When tested in vitro against M. incognita eggs,<br />

benzaldehyde was 100% effective as an ovicide for this species of root-knot nematode<br />

(Kokalis-Burelle et al., 2002).<br />

The direct effect of benzaldehyde on C. elegans chemotaxis kinetics was analyzed<br />

by Nuttley et al. (2001). An initial attractive response to 100% benzaldehyde was<br />

reported, followed by a strong aversion to the chemical. They determined this behavior<br />

to be mediated by two genetically separable response pathways. Initially, upon<br />

exposure, the attraction response dominates but eventually gives way to a repulsive<br />

response. Oka (2001) found that with juveniles of M. javanica, immobilization and<br />

hatching inhibition in vitro were greater with benzaldehyde and furfural than with<br />

several other essential oils. Benzaldehyde and furfural also reduced galling on tomato<br />

in pot experiments where other aldehydes were not effective (Oka, 2001).<br />

The effects of benzaldehyde combined with several organic amendments on soil<br />

microbial populations and plant-parasitic and nonparasitic nematodes were investigated<br />

by Chavarría-Carvajal et al. (2001). They found that benzaldehyde combined with<br />

most organic amendments reduced damage from parasitic nematodes and selected for<br />

predominantly gram-positive rhizosphere bacteria. When benzaldehyde was combined<br />

with root-knot nematode egg parasitizing isolates of the fungus Fusarium solani,<br />

increasing rates of benzaldehyde in soil reduced nematode penetration and infection<br />

of the host plant, and resulted in increased parasitism of M. javanica females by the<br />

fungus (Siddiqui and Shaukat, 2003). However, the increasing rates of benzaldehyde

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