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Control of Virus Diseases of Plants 159(TNV) protected the <strong>plant</strong> systemically aga<strong>in</strong>st disease caused by the fungusColletotrichum lagenarium (Jenns and Kuc, 1977). SAR expessed <strong>in</strong> Vigna <strong>plant</strong>sfollow<strong>in</strong>g <strong>in</strong>oculation with TNV aga<strong>in</strong>st challenge by TNV was not expressedaga<strong>in</strong>st challenge by a CMV which <strong>in</strong>fects the host systemically (Pennazzioand Roggero, 1991). Associated with SAR was the stimulation of ethyleneform<strong>in</strong>genzyme activity. Infection of ecotype Dijon of Arabidopsis thalianawith turnip cr<strong>in</strong>kle virus (TCV) leads to the resistance aga<strong>in</strong>st further <strong>in</strong>fectionby TCV or Pseudomonas syr<strong>in</strong>gae (Uknes et al., 1993). SAR <strong>in</strong> cucumber<strong>plant</strong>s aga<strong>in</strong>st powdery mildew disease, caused by Sphaerotheca fulig<strong>in</strong>ea(Schlechtend Fr.) Pollacci, was <strong>in</strong>duced by localized <strong>in</strong>fection <strong>in</strong> cucumbercotyledons with TNV (Farrag et al., 2007).The SAR phenomenon is observed both <strong>in</strong> dicotyledonous and monocotyledonous<strong>plant</strong>s; it provides the third and f<strong>in</strong>al l<strong>in</strong>e of defence aga<strong>in</strong>stpathogens. The first l<strong>in</strong>e of defence consists of genetically <strong>in</strong>herited resistancemechanisms that make <strong>plant</strong>s constitutively resistant to the majority ofpathogens present <strong>in</strong> the environment. The second l<strong>in</strong>e of defence is activated<strong>in</strong> the immediate vic<strong>in</strong>ity of the <strong>in</strong>fected or wounded site <strong>in</strong> an attemptto prevent the spread of pathogens throughout the <strong>plant</strong>. The local resistanceresponse develops more rapidly than SAR and <strong>in</strong>volves cell-wall and cuticlestrengthen<strong>in</strong>g, synthesis of tox<strong>in</strong>s, antifeedants and the production ofdefence-related prote<strong>in</strong>s <strong>in</strong>clud<strong>in</strong>g the PR prote<strong>in</strong>s. In addition to long-distancesignal molecules, local resistance may be partially mediated throughrelatively immobile endogenous elicitors, which <strong>in</strong>clude oligogalacturonidefragments of the <strong>plant</strong> cell wall (Lamb and Dixon, 1990). Several l<strong>in</strong>es of evidencesuggest that endogenous SA is a signal molecule <strong>in</strong> SAR. Involvementof SA <strong>in</strong> SAR came from the discovery that endogenous SA <strong>in</strong>creases by atleast 20-fold <strong>in</strong> the virus-<strong>in</strong>oculated leaves of tobacco (Malamy et al., 1990).The <strong>in</strong>crease co<strong>in</strong>cides with the appearance of hypersensitive response (HR)lesions on the <strong>in</strong>oculated leaves. Accumulation of SA <strong>in</strong>creased with the<strong>in</strong>tensity of HR and was proportional to the dose of virus <strong>in</strong>oculum (Yalpaniet al., 1991). Tissue accumulation of SA <strong>in</strong> TMV <strong>in</strong>oculated xanthi-nc tobaccoparalleled or preceded detectable <strong>in</strong>crease <strong>in</strong> the levels of PR-1 mRNA <strong>in</strong>both <strong>in</strong>oculated and un<strong>in</strong>oculated leaves (Malamy et al., 1990).7.4 SAR and the Role of PR Prote<strong>in</strong>sSAR strongly correlates with the coord<strong>in</strong>ate expression of at least n<strong>in</strong>e familiesof genes (sar genes), of which several encode the PR prote<strong>in</strong>s (Wardet al., 1991). Generally, tissues <strong>in</strong> which a significant amount of PR prote<strong>in</strong> hasbeen <strong>in</strong>duced are more resistant to <strong>in</strong>fection by pathogen than those that lackPR prote<strong>in</strong>s. PR prote<strong>in</strong>s were first discovered <strong>in</strong> 1970 <strong>in</strong> tobacco <strong>plant</strong>s react<strong>in</strong>ghypersensitively to TMV <strong>in</strong>fection (Gian<strong>in</strong>azzi et al., 1970; Van Loon andKammen, 1970). PR prote<strong>in</strong>s are produced <strong>in</strong> <strong>plant</strong>s <strong>in</strong> response to <strong>in</strong>fectionby viruses, bacteria (Metraux and Boller, 1986), fungi (Gian<strong>in</strong>azi et al., 1980)and viroids (Conejero et al., 1979) and are also synthesized <strong>in</strong> response tochemical treatments and specific physiological stresses (St<strong>in</strong>tzi et al., 1993).

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