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Plant basal resistance - Universiteit Utrecht

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Chapter 1<br />

constitutively produced in blackcurrant leaves, but is pathogen-inducible in rice leaves after<br />

infection (Kodama et al., 1988).<br />

Glucosinolates are N- and S- containing indolic phytoanticipins that are<br />

exclusively found in Brassicaceae. Upon tissue damage, glucosinolates are hydrolysed by<br />

endogenous β-thioglucoside glucohydrolases, also known as myrosinases, which results in<br />

the accumulation of toxic metabolites, such as isothiocyanates, thiocyanates and nitriles<br />

(Halkier and Gershenzon, 2006). Glucosinolates can be directly toxic, but their activity is<br />

mostly based on deterrence of plant attackers, including mammals, birds, insects, mollusks,<br />

nematodes, bacteria and fungi (Halkier and Gershenzon, 2006).<br />

The most abundant class of phytoanticipins in Poaceae are benzoxazinones<br />

(BXs). These phenolic compounds have broad-spectrum defence activity against insects,<br />

nematodes, bacteria and insects (Niemeyer, 1988, 2009). Their biosynthesis originates from<br />

indole and is mostly under developmental control, which leads to accumulation of less<br />

inactive BX-glucosides in the vacuole (Frey et al., 2009). BX-glucosides are hydrolysed by<br />

β-glucosidases upon tissue disruption, which leads to the release of biocidal aglycone BXs<br />

(Nikus and Jonsson, 1999).<br />

Semiochemicals<br />

Semiochemicals are naturally produced low molecular weight compounds used as signals<br />

in communication between organisms. Based on their effects, they can be divided into:<br />

pheromones (chemical cues used for intra-species communication), kairomones (chemcials<br />

used for host identification and location), allomones (defence secretions which only<br />

serve the producing organism itself) and allelochemicals (signalling chemicals used for<br />

communication between individuals of different species). Semiochemicals can be volatile<br />

or non-volatile. Volatile semiochemicals can act over long distances, while non-volatile<br />

semiochemicals more likely act over shorter ranges (Romeis and Zebitz, 1997). <strong>Plant</strong>-<br />

derived semiochemicals can originate from wide range of biosynthetic pathways, but<br />

predominantly come from the lipoxygenase and isoprenoid pathways. Well know examples<br />

of above-ground semiochemicals are monoterpenes, such as (E)-ocimene, sesquiterpenes,<br />

such as germacrene D, (E)-β-farnesene, and the aromatic compounds methyl salicylate. The<br />

emission of volatile semio-chemicals signals is not restricted to above-ground plant parts,<br />

the sesquiterpene (E)-β-caryophyllene was found to be released from maize root upon<br />

feeding by the Western Corn Rootworm, which can attract entomapathogenic nematodes<br />

(Rasmann et al., 2005).<br />

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