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158 H.N. Verma and V.K. Baranwaltobacco and tomato <strong>plant</strong>s express<strong>in</strong>g the pokeweed antiviral gene are foundto be resistant to a broad spectrum of <strong>plant</strong> viruses (Lodge et al., 1993).A problem often encountered <strong>in</strong> us<strong>in</strong>g type-I RIPs is the fact that theycannot <strong>in</strong>hibit prote<strong>in</strong> synthesis <strong>in</strong> <strong>in</strong>tact cells. However, when coupled totype-II RIP, they can be used effectively. The toxicity of <strong>plant</strong> materialsconta<strong>in</strong><strong>in</strong>g type-II RIPs have long been known and has great medic<strong>in</strong>alpotential. Recently, apoptosis was described <strong>in</strong> both lymphoid tissue and <strong>in</strong>the <strong>in</strong>test<strong>in</strong>e of abr<strong>in</strong>- and ric<strong>in</strong>-poisoned rats. Apoptosis was also observed<strong>in</strong> tissue culture of cancer cells treated with ric<strong>in</strong>.The antiviral activity of the RIP could be due to <strong>in</strong>activation of ribosomeof the <strong>in</strong>fected <strong>plant</strong> cell. As compared to the usefulness <strong>in</strong> the <strong>plant</strong>s, the roleof RIPs <strong>in</strong> human and animal systems has been much more widely documented.In spite of all these efforts, the biological significance of RIPs <strong>in</strong>nature is not yet known and moreover, their antiviral action does not seem todepend upon the <strong>in</strong>hibition of host ribosomes (Chen et al., 1993).Great potential exists today <strong>in</strong> elucidat<strong>in</strong>g the possible significance ofRIPs and their exact antiviral role which may not always depend upon <strong>in</strong>hibitionof host ribosomes. Their co-action with another antiviral mechanismalso needs to be explored, especially their place <strong>in</strong> the cascade of events follow<strong>in</strong>gviral <strong>in</strong>fection up to establishment of resistance, both systemic orlocalized.7.3 Pathogen-<strong>in</strong>duced Systemic Acquired Resistance (SAR)The <strong>in</strong>fection of <strong>plant</strong>s by necrotiz<strong>in</strong>g pathogens, <strong>in</strong>clud<strong>in</strong>g fungi, bacteriaand viruses, <strong>in</strong>duces systemic resistance to subsequent attack by the pathogens.This resistance is called systemic acquired resistance (SAR) (Kessmanet al., 1994; Ryals et al., 1994; Prasad et al., 2001). Also, it can be activated <strong>in</strong>numerous <strong>plant</strong>s by pre-<strong>in</strong>oculation with biotic <strong>in</strong>ducers <strong>in</strong>clud<strong>in</strong>g pathogens(Sticher et al., 1997). One of the prom<strong>in</strong>ent features of SAR is that resistanceis expressed aga<strong>in</strong>st pathogen which can be widely different from the<strong>in</strong>itial <strong>in</strong>fect<strong>in</strong>g pathogens.In a range of <strong>plant</strong> species, the development of necrotic lesions <strong>in</strong> responseto pathogen <strong>in</strong>fection leads to <strong>in</strong>duction of generalized disease resistance <strong>in</strong>un<strong>in</strong>fected tissue. Thus, TMV <strong>in</strong>oculated hypersensitive tobacco cultivardevelops systemic resistance aga<strong>in</strong>st TMV (virus), Phytophthora parasitica var.nicot<strong>in</strong>ae (fungi) and Pseudomonas tabaci (bacteria). TMV also <strong>in</strong>duced resistanceaga<strong>in</strong>st Peronospora tabac<strong>in</strong>a and reduced reproduction of the aphidMyzus persicae (McIntyre et al., 1981). Thus a s<strong>in</strong>gle viral agent <strong>in</strong>duced resistance<strong>in</strong> tobacco aga<strong>in</strong>st diverse challenges.The first report of virus <strong>in</strong>duced SAR came <strong>in</strong> 1952. Primary <strong>in</strong>oculationof the lower leaves of D. barbatus with carnation mosaic virus (CarMV)resulted <strong>in</strong> the development of fewer lesions on the upper leaves upon challenge<strong>in</strong>oculation with CarMV (Gilpatrick and We<strong>in</strong>traub, 1952). Virus<strong>in</strong>ducedresistance was further substantiated by Ross (1961a, 1961b) andLoebenste<strong>in</strong> (1963). Cucumber <strong>plant</strong>s <strong>in</strong>fected with tobacco necrosis virus

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