contain specific “avirulence” genes; races that are not suppressed lack a functional corresponding avirulence gene. In at least some cases, pathogen avirulence genes encode proteins that are required for infection of susceptible plant hosts (Kearney and Staskawicz 1990). The “gene-for-gene” concept was proposed to explain the interaction between a plant R gene and a pathogen avirulence gene, and this concept is used in agriculture to develop pest-protected crop varieties that are resistant to damage by pathogen races that have known virulence properties. A feature of race-specific R genes, and one of the major limitations associated with their use, is the occurrence of pathogen races that are unaffected by a given plant R gene; these can be pre-existing races that lack the corresponding avirulence genes or new races that have lost avirulence gene function. Study of numerous R genes isolated over the last few years has shown that many have a common evolutionary origin (Baker et al. 1997). Furthermore, race-specific R genes appear to function by triggering a cascade of molecular signaling and biochemical reactions that arrest pathogen spread at the initial site of infection, regardless of whether a particular R gene specifies resistance to a virus, fungus, or bacterium.

Several other types of disease-resistance genes that do not fit the gene-for-gene concept have also been identified. The HM1 gene of maize encodes a reductase that inactivates HC toxin, a cyclic tetrapeptide required for virulence of the fungus Cochliobolus carbonum race 1 (Johal and Briggs 1992). The recessive mlo gene in barley confers resistance to all races of the powdery mildew fungus, Erisyphe graminis f. sp. hordei, by priming the onset of several defense pathways (Buschges et al. 1997). Polygenic traits that confer quantitative pest-protection can also provide durable protection. Although the basis for this type of pest-protection is not entirely clear, cumulative effects of plant R genes that have been overcome by virulent pathogens might play a role in some systems (Li et al. 1999).

Genes for controlling insect and other invertebrate pests have also been identified and deployed, although they might be less common than plant R genes for viral, fungal, and bacterial pathogens. Some encode enzymes that catalyze synthesis of insecticidal or insect-deterrent compounds, whereas others trigger localized defense responses. Several nematode R genes are chemically related, or sequence-related, to race-specific pathogen R genes (Cai et al. 1997; Milligan et al. 1998); this suggests that the signaling mechanisms leading to resistance to nematode are similar to those for resistance to pathogens. The tomato Mi gene for resistance to the root-knot nematode, Meloidogyne incognita, also confers resistance to the potato aphid, Macrosiphum euphorbiae (Rossi et al. 1998; Vos et al. 1998); thus, some insect resistance genes could have broad specificity.



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