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--- bacteria:t3e:xopn [2020/07/03 18:48]
rkoebnik
+++ bacteria:t3e:xopn [2020/08/09 20:32] (current)
jvicente
@@ Line 7: / Line 7: @@
 Class: XopN\\
 Family: XopN\\
-Prototype: XopN (//Xanthomonas euvesicatoria// pv. //euvesicatoria// aka //Xanthomonas campestris// pv. //vescicatoria//; strain 85-10)\\
+Prototype: XopN (//Xanthomonas euvesicatoria// pv. //euvesicatoria//, ex //Xanthomonas campestris// pv. //vesicatoria//; strain 85-10)\\
 RefSeq ID: [[https://www.ncbi.nlm.nih.gov/protein/NP_643095|NP_643095]] (733 aa)\\
-D structure: unknown - similar to phosphatase 2a (pr65 / A) (Roden //et al//., 2004).
+D structure: unknown - similar to phosphatase 2a (pr65/A) (Roden //et al//., 2004).
 ===== Biological function =====
 === How discovered? ===
-XopN was identified in a genetic screen, using a Tn//5//-based transposon construct harboring the coding sequence for the HR-inducing domain of AvrBs2, but devoid of the effectors' T3SS signal, that was randomly inserted into the genome of //Xcv //strain 85-10. The XopN::AvrBs2 fusion protein triggered a //Bs2//-dependent hypersensitive response (HR) in pepper leaves (Roden //et al//., 2004).
+XopN was identified in a genetic screen, using a Tn//5//-based transposon construct harboring the coding sequence for the HR-inducing domain of AvrBs2, but devoid of the effectors' T3SS signal, that was randomly inserted into the genome of //X. campestris// pv. //vesicatoria// (//Xcv//) strain 85-10. The XopN::AvrBs2 fusion protein triggered a //Bs2//-dependent hypersensitive response (HR) in pepper leaves (Roden //et al//., 2004).
 === (Experimental) evidence for being a T3E ===
@@ Line 20: / Line 20: @@
 === Regulation ===
-Start codon of //xopN// was found downstream if a conserved cis-regulatory element, the plant-inducible promoter (PIP) box (TTCGG-N15-TTCTG). //xopN// is regulated by //hrpX// and //hrpG// genes (Cheong //et al//., 2013; Jiang //et al//., 2008).
+Start codon of //xopN// was found downstream of a conserved cis-regulatory element, the plant-inducible promoter (PIP) box (TTCGG-N15-TTCTG). //xopN// is regulated by //hrpX// and //hrpG// genes (Jiang //et al//., 2008; Cheong //et al//., 2013).
 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//) were significantly reduced in the //Xanthomonas oryzae// pv. //oryzae// Δ//xrvC// mutant compared with those in the wild-type strain PXO99<sup>A</sup>  , but this did not apply to //xopN// (Liu //et al.//, 2016).
@@ Line 28: / Line 28: @@
   * Its homolog XopN <sub>Xcc</sub>  was found as well to be required for full virulence on Chinese radish (Jiang //et al//., 2008).
   * XopN has been shown to play a role in host defence systems causing the reduction of PAMP-triggered immune responses and reduce the callose deposition in the host tissue. Moreover the deletion of //xopN//  open reading frame (ORF) reduced the //Xcv//  strain virulence exhibited by lower bacterial spot symptoms occurrence (Kim //et al//., 2009).
+  * The role of XopN in X. oryzae pv. oryzae is dependent on leaf stage (Cheong et al., 2013).
+  * XopN has been shown to be required for maximal pathogenicity of //X. axonopodis//  pv. //punicae//  (//Xap//) in pomegranate (Kumar and Mondal, 2013). The deletion of XopN from Xap caused higher accumulation of reactive oxygen species showing that XopN suppresses ROS-mediated defense responses during blight pathogenesis in pomegranate (Kumar //et al.//, 2016).
   * A Δ//xopN//–Δ//xopQ //double knock-out mutant in //X. phaseoli//  pv. //manihotis//  (//Xpm//) was less aggressive in the cassava host plant than its single mutation counterparts. In addition, //in planta //  bacterial growth was reduced at 5 dpi in the double mutant with respect to the wild-type strain CIO151 and individual knock-out strains. The phenotype of the double mutant could be complemented when transforming a plasmid containing //xopQ//. These results confirmed that //xopN //and// xopQ //are functionally redundant in //Xpm//  (Medina //et al.//, 2017).
+  * //Agrobacterium//  mediated transient transfer of the gene for XopN resulted in suppression of rice innate immune responses induced by LipA, a hydrolitic enzyme secreted by //X. oryzae//  pv. //oryzae//  (Xoo), but a //xopN// <sup>//-// </sup>   mutant of //Xoo//  retains the ability to suppress these innate immune responses indicating other functionally redundant proteins; XopQ, XopX and XopZ were shown to be suppressors of LipA induced innate immune responses; mutation in any one of the //xopN, xopQ, xopX or xopZ//  genes causes partial virulence deficiency (Sinha et al., 2013). XopN was shown to contribute significantly to //X. oryzae//  pv. //oryzae//  (Xoo) virulence on a susceptible rice variety Nipponbare. XopN was shown to be highly translocated to suppress rice defense responses (Mo //et al.//, 2020).
+  * XopN and AvrBS2 were shown to significantly contribute to virulence of //X. oryzae//  pv. //oryzicola//  (Xoc GX01) (Liao //et al.//, 2020).
 === Localization ===
-XopN was localized by confocal microscopy using fluorescent tagged fusion (yellow fluorescent protein [YFP]-XopN). [YFP]-XopN was localized throughout the plant cytoplasm and also associated with the plant plasma membrane (PM) (Kim //et al//., 2009).
+XopN was localized by confocal microscopy using fluorescent tagged fusion (yellow fluorescent protein [YFP]-XopN). [YFP]-XopN was localized throughout the plant cytoplasm and also associated with the plant plasma membrane (PM) (Kim //et al//., 2009). Kumar et al. (2016) demonstrated that XopN is localized in the pasma membrane of //N. benthamiana//, pomegranate and onion cells.
 === Enzymatic function ===
-Unknown – Kim //et al//. (2009) did not confirm that XopN is an enzyme (Kim //et al//., 2009).
+XopN binds TARK1, a tomato atypical receptor kinase required for PTI. Taylor //et al.//  (2012) showed that XopN promotes TARK1/TFT1 complex formation //in vitro//  and //in planta//  by functioning as a molecular scaffold.TFT proteins are involved in immune signaling during //X. euvesicatoria//  infection and can interact with multiple effectors including XopN (Dubrow //et al.//, 2018). TARK1 was shown to interact with proteins predicted to be associated with stomatal closure (Guzman et al., 2020).
+Three effectors (XopZ, XopN and XopV) were shown to be able to supress the peptidoglycan-triggered MAPK activation and a triple mutant of Xoo lacking these genes showed additively reduced virulence (Long et al., 2018).
 === Interaction partners ===
-XopN interact with two types of proteins in tomato: Tomato Atypical Receptor-like Kinase1 (TARK1) and four Tomato Fourteen-Three-Three isoforms (TFT1, TFT3, TFT5, and TFT6) (Kim //et al//., 2009).
+XopN interact with two types of proteins in tomato: Tomato Atypical Receptor-like Kinase1 (TARK1) and four Tomato Fourteen-Three-Three isoforms (TFT1, TFT3, TFT5, and TFT6) (Kim //et al//., 2009). XopN interacts with the tomato 14-3-3 isoform TFT1 that functions in PTI and is a XopN virulence target (Taylor //et al.//, 2012).
+Two rice proteins, OsVOZ2 and a putative thiamine synthase (OsXNP) were identified as targets of XopN<sub>KXO85</sub>  by yeast two-hybrid screening (Cheong et al., 2012).
 ===== Conservation =====
@@ Line 46: / Line 54: @@
 === In xanthomonads ===
-Yes (//e.g.//, //X//. //campestris//, //X//. //citri//,// X//. //oryzae//). Since the G+C content of the //xopN//  gene is similar to that of the //Xcv//  //hrp gene//  cluster, it may be a member of a “core” group of //Xanthomonas//  spp. effectors (Roden et al., 2004).
+Yes (//e.g.//, //X. axonopodis//, //X//. //campestris//, //X//. //citri//,// X//. //oryzae//). Since the G+C content of the //xopN//  gene is similar to that of the //Xcv////hrp gene//  cluster, it may be a member of a “core” group of //Xanthomonas//  spp. effectors (Roden et al., 2004).
 === In other plant pathogens/symbionts ===
@@ Line 55: / Line 63: @@
 Cheong H, Kim CY, Jeon JS, Lee BM, Sun Moon J, Hwang I (2013). //Xanthomonas oryzae//  pv. //oryzae//  type III effector XopN targets OsVOZ2 and a putative thiamine synthase as a virulence factor in rice. PloS ONE 8: e73346. DOI: [[https://doi.org/10.1371/journal.pone.0073346|10.1371/journal.pone.0073346]].
+Dubrow Z, Sunitha S, Kim JG, et al. Tomato 14-3-3 Proteins Are Required for Xv3 Disease Resistance and Interact with a Subset of Xanthomonas euvesicatoria Effectors. //Mol Plant Microbe Interact//. 2018;31(12):1301-1311. DOI:10.1094/MPMI-02-18-0048-R
+Guzman AR, Kim JG, Taylor KW, Lanver D, Mudgett MB. Tomato Atypical Receptor Kinase1 Is Involved in the Regulation of Preinvasion Defense. //Plant Physiol//. 2020;183(3):1306-1318. DOI:10.1104/pp.19.01400
 Jiang B, He Y, Cen W, Wei H, Jiang G, Jiang W, Hang X, Feng J, Lu G, Tang D, Tang J (2008). The type III secretion effector XopXccN of //Xanthomonas campestris//  pv. //campestris//  is required for full virulence. Res. Microbiol. 159: 216-220. DOI: [[https://doi.org/10.1016/j.resmic.2007.12.004|10.1016/j.resmic.2007.12.004]]
 Kim JG, Li X, Roden JA, Taylor KW, Aakre CD, Su B, Landone S, Kirik A, Chen Y, Baranage G, Martin BG, Mudgett BM, McLane H (2009). //Xanthomonas//  T3S effector XopN suppresses PAMP-triggered immunity and interacts with a tomato atypical receptor-like kinase and TFT1. Plant Cell 21: 1305-1323. DOI: [[https://doi.org/10.1105/tpc.108.063123|10.1105/tpc.108.063123]]
+Kumar R, Mondal KK (2013). XopN-T3SS effector modulates in planta growth of Xanthomonas axonopodis pv. punicae and cell-wall-associated immune response to induce bacterial blight in pomegranate. Physiological and Mol. Plant Pathol. 84: 36-43. DOI: 10.1016/j.pmpp.2013.06.002
+Kumar R, Soni M, Mondal KK (2016). XopN-T3SS effector of //Xanthomonas axonopodis//  pv. //punicae//  localizes to the plasma membrane and modulates ROS accumulation events during blight pathogenesis in pomegranate. Microbiol. Res. 193: 111-120. DOI: [[https://doi.org/10.1016/j.micres.2016.10.001|10.1016/j.micres.2016.10.001]]
+Liao ZX, Li JY, Mo XY, et al. Type III effectors xopN and avrBS2 contribute to the virulence of Xanthomonas oryzae pv. oryzicola strain GX01. //Res Microbiol//. 2020;171(2):102-106. DOI:10.1016/j.resmic.2019.10.002
 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]]
+Long J, Song C, Yan F, Zhou J, Zhou H, Yang B (2018). Non-TAL effectors from //Xanthomonas oryzae//  pv. //oryzae//  suppress peptidoglycan-triggered MAPK activation in rice. Front. Plant Sci. 9: 1857. doi: [[https://doi.org/10.3389/fpls.2018.01857|10.3389/fpls.2018.01857]]
 Medina CA, Reyes PA, Trujillo CA, Gonzalez JL, Bejarano DA, Montenegro NA, Jacobs JM, Joe A, Restrepo S, Alfano JR, Bernal A (2018). The role of type III effectors from //Xanthomonas axonopodis//  pv. //manihotis//  in virulence and suppression of plant immunity. Mol. Plant Pathol. 19: 593-606. DOI:[[https://doi.org/10.1111/mpp.12545|10.1111/mpp.12545]]
+Mo X, Zhang L, Liu Y, Wang X, Bai J, Lu K, Zou S, Dong H, Chen L (2020). Three proteins (Hpa2, HrpF and XopN) are concomitant type III translocators in bacterial blight pathogen of rice. Frontiers in Microbiology 11: 1601. DOI=10.3389/fmicb.2020.01601
 Roden JA, Belt B, Ross JB, Tachibana T, Vargas J, Mudgett MB (2004). A genetic screen to isolate type III effectors translocated into pepper cells during //Xanthomonas//  infection. Proc. Natl. Acad. Sci. USA 101: 16624-16629. DOI: [[https://doi.org/10.1073/pnas.0407383101|10.1073/pnas.0407383101]]
+Sinha D, Gupta MK, Patel HK, Ranjan A, Sonti RV (2013). Cell wall degrading enzyme induced rice innate immune responses are suppressed by the type 3 secretion system effectors XopN, XopQ, XopX and XopZ of //Xanthomonas oryzae//  pv. //oryzae//. PLoS One 8: e75867. DOI: [[https://doi.org/10.1371/journal.pone.0075867|10.1371/journal.pone.007586]]7
+Taylor KW, Kim JG, Su XB, Aakre CD, Roden JA, Adams CM, Mudgett MB (2012). Tomato TFT1 is required for PAMP-triggered immunity and mutations that prevent T3S effector XopN from binding to TFT1 attenuate //Xanthomonas//  virulence. PLoS Pathog. 8: e1002768. DOI: [[https://doi.org/10.1371/journal.ppat.1002768|10.1371/journal.ppat.1002768]]

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