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natural-products-in-plant-pest-management

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118 L.A. ShcherbakovaAt least 14 genes encod<strong>in</strong>g different protease <strong>in</strong>hibitors alone or <strong>in</strong>comb<strong>in</strong>ation with other heterologous genes have been reported to be transferred<strong>in</strong>to cultured <strong>plant</strong>s, which showed <strong>in</strong>creased resistance predom<strong>in</strong>antlyto <strong>in</strong>sects (Valuyeva and Mosolov, 2004). Plant protease <strong>in</strong>hibitorshave also been used to eng<strong>in</strong>eer resistance aga<strong>in</strong>st viruses <strong>in</strong> transgenic<strong>plant</strong>s. For example, expression of the gene encod<strong>in</strong>g the cyste<strong>in</strong>e prote<strong>in</strong>ase<strong>in</strong>hibitor from rice, oryzacystat<strong>in</strong>, by transgenic tobacco was found to conferresistance aga<strong>in</strong>st tobacco etch virus and potato Y virus, replication of whichdepends on cyste<strong>in</strong>e prote<strong>in</strong>ase activity (Valuyeva and Mosolov, 2004; Habiband Khalid, 2007).In the green consumerization context, it is important that some of theavailable prote<strong>in</strong>ase <strong>in</strong>hibitors are <strong>in</strong>active for non-pathogenic microorganismsand do not <strong>in</strong>hibit activity of prote<strong>in</strong>ases of animal orig<strong>in</strong>. Thus, chestnutcystat<strong>in</strong>, which strongly <strong>in</strong>hibits the protease activity and the growth ofpathogenic B. c<strong>in</strong>erea, Colletotrichum gram<strong>in</strong>icola, and Stagonospora nodorum,has no effect on the protease activity and the growth of the saprophyte Trichodermaviride (Pernas et al., 1999), and a prote<strong>in</strong>ase <strong>in</strong>hibitor extracted frombean seeds specifically suppresses ser<strong>in</strong>e prote<strong>in</strong>ase of pathogenic C. l<strong>in</strong>demuthianumbut does not <strong>in</strong>fluence animal tryps<strong>in</strong> and chymotryps<strong>in</strong> activity(Valuyeva and Mosolov, 2004). Non-specific antipathogenic activity of protease<strong>in</strong>hibitors suggests that transgenic crops produc<strong>in</strong>g <strong>in</strong>hibitors of <strong>in</strong>secticidalor nematicidal prote<strong>in</strong>ases may be <strong>in</strong>corporated <strong>in</strong>to <strong>in</strong>tegratedsystems of <strong>plant</strong> protection aga<strong>in</strong>st <strong>pest</strong>s and pathogens. A further advantageof this approach is the possibility that <strong>in</strong>hibitory activity aga<strong>in</strong>st prote<strong>in</strong>asescould be comb<strong>in</strong>ed with the <strong>in</strong>secticidal activity of lect<strong>in</strong>s, result<strong>in</strong>g <strong>in</strong> asynergistic antipathogenic effect.PR prote<strong>in</strong>sPlants have numerous defence mechanisms that are activated <strong>in</strong> response topathogen attacks, abiotic stresses and chemicals that mimic pathogenchallenge. Production of prote<strong>in</strong>s related to pathogenesis (PR prote<strong>in</strong>s) is onesuch <strong>in</strong>ducible mechanism (Van Loon et al., 1994; Van Loon et al., 2006). Theseprote<strong>in</strong>s are not detectable at all or are present only at basal concentrations<strong>in</strong> healthy <strong>plant</strong> tissues. S<strong>in</strong>ce the term PR prote<strong>in</strong>s based on the abovecharacteristics was often used for all constitutive <strong>plant</strong> prote<strong>in</strong>s for whichcontent or <strong>in</strong>creas<strong>in</strong>g activity was <strong>in</strong>duced by microorganisms, the new term‘<strong>in</strong>ducible defence-related prote<strong>in</strong>s’ was recently <strong>in</strong>troduced (Van Loon et al.,2006).PR prote<strong>in</strong>s consist of a large variety of families with members that differ<strong>in</strong> occurrence, expression and biological activities; they are divided <strong>in</strong>to 17classes (Sels et al., 2008). A range of the above-mentioned <strong>plant</strong> prote<strong>in</strong>s andpeptides such as chit<strong>in</strong>ases, β-1,3-glucanases, peroxidase, defens<strong>in</strong>s and prote<strong>in</strong>ase<strong>in</strong>hibitors are PR prote<strong>in</strong>s and vice versa; some typical PR prote<strong>in</strong>sare antimicrobial and <strong>in</strong>hibit pathogen growth <strong>in</strong> vitro. For <strong>in</strong>stance, tobaccoPR-1a is antifungal, and tomato prote<strong>in</strong>s of the PR-1 family <strong>in</strong>hibit zoosporegerm<strong>in</strong>ation and reduce pathogenicity of P. <strong>in</strong>festans. The ability to disrupt

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