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186<br />

REN-SEN ZENG<br />

Glomus intraradices, Glomus mosseae, and Gigaspora rosea leads to the accumulation<br />

of similar cyclohexenone derivatives (Vierheilig et al., 2000). However, no fungusspecific<br />

induction of these compounds are known. Pathogens and endophyte did not<br />

induce the formation of cyclohexenone derivatives in barley roots (Maier et al., 1997).<br />

The role of cyclohexenone derivatives in disease resistance is unknown.<br />

In response to pathogen attack, plants activate an array of inducible defense<br />

reactions, many of which involve the transcriptional activation of the corresponding<br />

defense genes, including genes that encode enzymes involved in the synthesis of lignin<br />

and phytoalexins (Dixon et al., 1984; Dixon and Harrison 1990). Transcript levels of<br />

some pivotal enzymes of defense response of plants significantly increase after<br />

mycorrhizal fungal infection (Harrison and Dixon, 1993; 1994). Several inducible<br />

defence-related genes, including those encoding isoflavonoid phytoalexins such as<br />

phenylalanine ammonia lyase (PAL), chalcone synthase (CHS), chalcone isomerase<br />

(CHI) and for the cell wall structural protein HRGP, have been reported to be induced<br />

during mycorrhiz establishment (Tagu and Martin, 1996). Mycorrhization resulted<br />

in a local and systemic induction of plant defence-related enzymes chitinase,<br />

chitosanase and beta-1,3-glucanase, as well as superoxide dismutase in tomato plants<br />

(Pozo et al., 2002). Chalcone isomerase (CHI) and chitinase activities increased in<br />

inoculated roots prior to mycorrhizal colonization (Volpin et al., l994, Kapulink et<br />

al., 1996), whereas the increase in PAL activity coincided with colonization. Production<br />

of some new compounds, and increase in the activity of the enzymes peroxidase and<br />

polyphenol oxidase, was observed following inoculation with AM fungi (Charitha<br />

Devi and Reddy, 2002). Dumas-Gaudot et al. (1992a,b) found new chitinase isoforms<br />

that were specifically induced in several AM associations and were different from<br />

those elicited by root fungal pathogens, indicating a different pattern of plant response<br />

to pathogenic and mutualistic fungi. Glomus mosseae induced new chitinase isoforms<br />

in tomato roots (Pozo et al., 1996). Expression of genes encoding enzymes that<br />

synthesize phenolpropanoid compounds has been detected in mycorrhizal roots (Garcia-<br />

Garrido and Ocampo, 2002). Other defense related genes shown to be upregulated in<br />

mycorrhizal symbioses include: genes involved in metabolism of reactive oxygen<br />

species, chitinase and beta I ,3-glucanase, and genes involved in senescence, including<br />

glutathione-S-transferase. Mycorrhiza also induced changes in PR protein expression<br />

in tobacco leaves (Shaul et al., 1999).<br />

Colonization of barley, wheat and maize and rice roots by Glomus intraradices<br />

resulted in strong induction of transcript levels of the pivotal enzymes of<br />

methylerythritol phosphate pathway of isoprenoid biosynthes i.e., 1 -deoxy-D-xylulose<br />

5-phosphate synthase (DXS) and 1 -deoxy-D-xylulose 5-phosphate reductoisomerase<br />

(DXR) (Walter et al., 2000). At the same time six cyclohexenone derivatives were<br />

characterized from mycorrhizal wheat and maize roots. DXS2 transcript levels are<br />

low in most tissues but are strongly stimulated in roots upon colonization by<br />

mycorrhizal fungi, correlated with accumulation of carotenoids and apocarotenoids<br />

(Walter et al., 2002). Some reports show that the AM symbiosis may cause an increase,<br />

decrease, or no change in the plant defense reactions (Guenoune et al., 2001; Mohr et<br />

al., 1998).

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