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====== XopY ======

Author: [[https://www.researchgate.net/profile/Irena_Macioniene|Irena Mačionienė]]\\
Internal reviewer: [[https://www.researchgate.net/profile/Lucas_Moriniere|Lucas Morinière]]\\
Expert reviewer: FIXME

Class: XopY\\
Family: XopY\\
Prototype: XOO1488 (//Xanthomonas oryzae// pv. //oryzae//; strain MAFF 311018)\\
RefSeq ID: [[https://www.ncbi.nlm.nih.gov/protein/AEQ97580.1|AEQ97580]] (276 aa)\\
3D structure: Unknown

===== Biological function =====

=== How discovered? ===

XopY was discovered by screening the genome of //X. oryzae //pv. //oryzae// MAFF 311018 for proteins displaying a N-terminal amino acid pattern associated with T3S substrates in //Pseudomonas syringae// (Furutani //et al.//, 2009). It has been primarily referred to as XOO1488, and then XopY (Song & Yang, 2010).
=== (Experimental) evidence for being a T3E ===

//X. oryzae //pv. //oryzae //transformants containing a plasmidic fusion of XopY (= XOO1488) with the Cya translocation reporter system were inoculated in tomato leaves. An increase of cAMP in the inflitrated areas was observed, thus revealing translocation of the fused protein into plant cells (Furutani //et al.//, 2009).
=== Regulation ===

XopY from //X. oryzae //pv. //oryzae// posseses a PIP and ‐10 box in the promoter region (TTCGB‐N<sub>15</sub> ‐TTCGB‐N<sub>30–32</sub> ‐YANNNT) (Yamaguchi //et al//., 2013a). Also, it was shown to be regulated by HrpX (Furutani //et al.//, 2009).

qRT-PCR revealed that transcript levels of 15 out of 18 tested non-TAL effector genes (as well as the regulatory genes //hrpG// and //hrpX//), including //xopY//, were significantly reduced in the //Xanthomonas oryzae// pv. //oryzae// Δ//xrvC// mutant compared with those in the wild-type strain PXO99<sup>A</sup>  (Liu //et al.//, 2016).
=== Phenotypes ===

Transgenic rice plants expressing XOO1488 (Xoo1488-OX) were generated and inoculated with the T3SS-deficient //hrpX// mutant of //Xoo//, which is incapable of type III effector delivery. The //Xoo hrpX// mutant did not cause lesions in wild-type plant, presumably because of strong induction of PTI. In contrast, Xoo1488-OX plants had severe disease symptoms following infection with the //Xoo hrpX// mutant. Bacterial populations of the //Xoo hrpX// mutant in Xoo1488-OX leaves were higher than in wild-type plants. The growth of wild-type isolate //Xoo// MAFF311018 in Xoo1488-OX plants also increased significantly over its growth in wild-type plants. Thus, it is likely that XopY inhibits PTI induced by infection of //Xoo hrpX// mutant. However, it was noted that XopY knockout strain did not exhibit any defect in virulence (Yamaguchi //et al//., 2013a). XopY was also shown to inhibit chitin-induced expression of defense-related genes (Yamaguchi //et al.//, 2013b). XopY of //X. oryzae// pv. //oryzicola// was also shown to trigger HR in non-host //Nicothiana benthamiana// plants (Li //et al.//, 2015).
=== Localization ===

Unknown.

=== Enzymatic function ===

Unknown.

=== Interaction partners ===

XopY was demonstrated to target OsRLCK185. Expression of XopY in rice cells compromises OsRLCK185-mediated immune responses, which is consistent with the fact that Xoo1488 inhibits trans-phosphorylation of the activation domain of OsRLCK185 by OsCERK1. Interestingly, XopY is phosphorylated by OsRLCK185, suggesting that modification of XopY in host cell may affect their virulent activity (Yamaguchi //et al//., 2013b).
===== Conservation =====

=== In xanthomonads ===

Yes (//e.g.//, //X. oryzae//, //X. translucens, X. vasicola//).
=== In other plant pathogens/symbionts ===

No.

===== References =====

Furutani A, Takaoka M, Sanada H, Noguchi Y, Oku T, Tsuno K, Ochiai H, Tsuge S (2009). Identification of novel type III secretion effectors in// Xanthomonas oryzae// pv. //oryzae//. Mol. Plant Microbe Interact. 22: 96-106. DOI: [[https://doi.org/10.1094/MPMI-22-1-0096|10.1094/MPMI-22-1-0096]]

Li S, Wang Y, Wang S, Fang A, Wang J, Liu L, Zhang K, Mao Y, Sun W (2015). The type III effector AvrBs2 in //Xanthomonas oryzae// pv. //oryzicola// suppresses rice immunity and promotes disease development. Mol. Plant Microbe Interact. 28: 869-880. DOI: [[https://doi.org/10.1094/MPMI-10-14-0314-R|10.1094/MPMI-10-14-0314-R]]

Liu Y, Long J, Shen D, Song C (2016). //Xanthomonas oryzae// pv. //oryzae// requires H-NS-family protein XrvC to regulate virulence during rice infection. FEMS Microbiol. Lett. 363: fnw067. DOI: [[https://doi.org/10.1093/femsle/fnw067|10.1093/femsle/fnw067]]

Song C, Yang B (2010). Mutagenesis of 18 type III effectors reveals virulence function of XopZ<sub>PXO99</sub> in //Xanthomonas oryzae //pv. //oryzae//. Mol. Plant Microbe Interact. 23: 893-902. DOI: [[https://doi.org/10.1094/MPMI-23-7-0893|10.1094/MPMI-23-7-0893]]

Yamaguchi K, Nakamura Y, Ishikawa K, Yoshimura Y, Tsuge S, Kawasaki T (2013a). Suppression of rice immunity by //Xanthomonas oryzae //type III effector Xoo2875. Biosci. Biotechnol. Biochem. 77: 796-801. DOI: [[https://doi.org/10.1271/bbb.120929|10.1271/bbb.120929]]

Yamaguchi K, Yamada K, Ishikawa K, Yoshimura S, Hayashi N, Uchihashi K, Ishihama N, Kishi-Kaboshi M, Takahashi A, Tsuge S, Ochiai H, Tada Y, Shimamoto K, Yoshioka H, Kawasaki T (2013b). A receptor-like cytoplasmic kinase targeted by a plant pathogen effector is directly phosphorylated by the chitin receptor and mediates rice immunity. Cell Host Microbe 13: 347-357. DOI: [[https://doi.org/10.1016/j.chom.2013.02.007|10.1016/j.chom.2013.02.007]]
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