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98 Figen Mert-Türk and Eric B. Holub a b c d e f Figure 1: The interaction phenotypes between the isolates of Peronospora parasitica and the lines of Arabidopsis thaliana. Nd-1 (a) exhibited PN (pitting with no pathogen sporulation phenotypes) after inoculation with the Emoy2 isolate. However, Col-pad3 was more susceptible than wild-type accession Col-0 following inoculation with the same isolate and exhibited medium sporulation (b). The response of Nd-pad3 to Emoy2 was same as the Nd-1 parent (c). Flecking host response with no sporulation was observed in Nd-1 (d) following inoculation with any RPP13-avirulent isolate (d). Col-pad3 was fully susceptible as it does not have a functional resistance gene to recognise the isolates (e). The Nd-pad3, however, exhibited the same interaction phenotype observed in Nd-1 parent (f). These results indicate that PAD3 is not required for resistance mediated by either RPP1 or RPP13 genes against to the corresponding isolates of P. parasitica, therefore, camalexin is not an essential defence component for either source of downy mildew resistance. Therefore, phytoalexin deficient Nd-1 has only camalexin disability but still keeps other known defence molecules for protecting itself from pathogen. Because the Col-pad3 mutant was more susceptible than wild-type accession Col-0 following inoculation with the isolate Emoy2, the result might suggest that camalexin accumulation is important but not enough in protection Col-0 from Emoy2. Col-0 recognises the avirulence product of P. parasitica with RPP4 resistance gene. Therefore RPP4 is likely camalexin-dependent. Thomma et al. (1999) reported that resistance of the Col-pad3 mutant to Botrytis cinerea is not altered relative to that of wild-type. When Col-pad3 inoculated with Alternaria brassicicola, however, it exhibited enhanced susceptibility compared to the wild-type suggesting a role for PAD3 in resistance to A. brassicicola. This finding proved that camalexin is an important defence components. The same research group also reported that PR1, therfore SA, and also jasmonate/ethylene-dependent defence responses were not impaired in the Col-pad3 mutant following inoculation with A. brassicicola. Therefore camalexin production appears to be controlled by a pathway that exhibits little or no cross-talk with salicylate-, ethylene-, and jasmonate-dependent signalling events. Acknowledgement This research was supported financially by Turkish Higher Educational Council (YOK) and Çanakkale Onsekiz Mart University, Turkey. This article is dedicated to first author’s parents for their ever-lasting help and support during her education. References Aarts N, Metz M, Holub E, Staskawicz BJ,Daniels MJ and Parker J. Different requirements for EDS1 and NDR1 by disease resistance genes define at least two R gene-mediated signalling pathways in Arabidopsis. Proc of the Nat Ac of Sci USA. 96: 10307-10311, 1998. Bittner-Eddy P, Can C, Gunn N, Pinel M, Tör M, Crute I, Holub E and Beynon J. Genetic and physical mapping of the RPP13 locus in Arabidopsis responsible for specific recognition of several Peronospora parasitica (downy mildew) isolates. Mol Plant-Microbe Int. 12: 792-802, 1999. Bittner-Eddy PD, Crute IR, Holub EB and Beynon JL. RPP13 is a simple locus in Arabidopsis thaliana for alleles that specify downy mildew resistance to different avirulence determinants in Peronospora parasitica. The Plant Journal 21: 189-214, 2000. Botella MA, Parker JE, Frost LN, Bittner-Eddy PD, Beynon JL, Daniels MJ, Holub EB and Jones JDG. Three genes of Arabidopsis RPP1 complex resistance
locus recognise distinct Peronospora parasitica avirulence determinants. The Plant Cell. 10: 1847- 1860, 1998. Browne LM, Conn KL, Ayer WA and Tewari JP. The camalexins: new phytoalexins produced in the leaves of Camelina sativa (Cruciferae). Tetrehedron. 47: 3909- 3914, 1991. Dixon RA, Lamb CJ, Sewalt VJH and Paiva NL. Metabolic engineering: Prospects for crop improvement through the genetic manipulation of phenylpraponoid biosynthesis and defence responses- a review. Gene. 179: 61-71, 1996. Glazebrook J. and Ausubel FM. Isolation of phytoalexin-deficient mutants of Arabidopsis thaliana and characterisation of their interactions with bacterial pathogens. Proc of the Nat Ac of Sci USA. 91: 8955- 8959, 1994. Glazebrook J, Zook M, Mert F, Kagan I, Rogers EE, Crute IR, Holub EB, Hammerschmidt R and Ausubel FM. Phytoalexin-deficient mutants of Arabidopsis reveal that PAD4 encodes a regulatory factor and that four PAD genes contribute to downy mildew resistance. Genetics. 146: 381-392, 1997. Holub EB, Beynon JL and Crute IR. Phenotypic and genotypic characterisation of interactions between isolates of Peronospora parasitica and accessions of Arabidopsis thaliana. Mol Plant Mic Int. 7: 223-239, 1994. Holub EH. Organisation of resistance genes in Arabidopsis. In: The Gene-For-Gene Relationship in Plant-Parasite Interactions. Crute IR, Holub EB and BurdonJJ (Eds). CAB International. 5-26, 1997. Holub EB and Beynon JL. Symbiology of mouse-ear cress (Arabidopsis thaliana) and Oomycetes. Adv in Bot Res. 24: 227-272, 1997. Kúc J. Phytoalexins, stress metabolism, and disease resistance in plants. Annu Rev of Phytopathol. 33: 275- 297, 1995. Mansfield JW. Role of phytoalexins in disease resistance. In: Phytoalexins. Mansfield JW and Bailey J (Eds). Blackie, Glasgow. 1982. Mansfield JW. Antimicrobial compounds and resistance: the role of phytoalexins and antianticipins. In: Mechanisms of Resistance to Plant Diseases. Slusarenko AJ, Fraser RSS and VanLoon LC (Eds). Kluwer, Amsterdam. 1999. Mert-Türk F, Bennett MH, Glazebrook J, Mansfield J and Holub E. Biotic and abiotic elicitation of camalexin in Arabidopsis thaliana. 7 th International Congress of Plant Pathology. Edinburgh, Scotland, UK.1998. Mert-Türk F. Quantification of enhanced downy mildew susceptibility and camalexin accumulation in the mutants of Arabidopsis thaliana. PhD thesis. University of London, Imperial College of Science, Technology and Medicine, UK. 2001. Camalexin and RPP1 and RPP13 resistance genes 99 Nam HG, Giraudat J, Den Boer B, Moonan F, Loos WDB, Hauge BM and Goodman HM. Restriction fragment polymorphism lincage map of Arabidopsis thaliana. Plant Cell. 1: 699-705, 1989. Oppenheimer DG, Herman PL, Sivakumarin S, Esch J and Marks MD. A myb gene required for leaf trichome differentiation in Arabidopsis is expressed in stipules. Cell. 67: 483-493, 1991. Thomma BPHJ, Nelissen I, Eggermont K and Broekaert F. Deficiency in phytoalexin production causes enhanced susceptibility of Arabidopsis thaliana to the fungus Alternaria brassicola. The Plant Journal. 19: 163-171, 1999. Tör M, Holub EB, Brose E, Musker R, Gunn N, Can C, Crute IR and Beynon JL. Map position of three loci in Arabidopsis thaliana associated with isolate-specific recognition of Peronospora parasitica (downy mildew). Mol Plant Mic Int. 7: 214-222, 1994. Tsuji J, Jackson EP, Gage DA, Hammerschmidt R and Somerville SC. Phytoalexin accumulation in Arabidopsis thaliana during the hypersensitive reaction to Pseudomonas syringae pv. syringae. Plant Physiol. 98: 1304-1309, 1992.
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