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--- bacteria:t3e:xopz [2020/07/09 12:13]
rkoebnik [XopZ]
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-====== XopZ ======
-Author: Marlène Lachaux\\
-Internal reviewer: FIXME \\
-Expert reviewer: FIXME
-Class: XopZ\\
-Family: XopZ\\
-Prototype: XopZ (//Xanthomonas oryzae// pv. o//ryzae//; strain PXO99<sup>A</sup> )\\
-RefSeq ID: 1,414-amino-acid protein 3D structure: The N-terminus of XopZ<sub>PXO99</sub>, containing two Nuclear Localization Signals (NLS) signals and several Nuclear Export Signals (NES) (Zhou //et al//., 2015).\\
-D structure: Unknown
-===== Biological function =====
-=== How discovered? ===
-In 2009, generation of PXO99 mutants for 18 non-TAL T3 effector genes allowed to investigate the function of several T3Es in //Xoo// strain PXO99<sup>A</sup>  . It was reported on XopZ that contributes to the full virulence of the strain PXO99<sup>A</sup>  (Ryan //et al//., 2009; Song //et al//., 2010).
-=== (Experimental) evidence for being a T3E ===
-With a PIP box upstream of the predicted translation start site //xopZ<sub>PXO99 </sub> //gene is certainly inducible in planta and regulated through the hypersensitive reaction and pathogenicity (//hrp//) regulatory network. PXO99<sup>A</sup>  and an //hrpG// mutant were grown in NB or //Xanthomonas hrp//-inducing medium (XOM2). The expression of //xopZ<sub>PXO99 </sub> //was only observed, by RT-PCR, in XOM2 medium and was //hrpG// dependent (Song //et al//., 2010).
-=== Regulation ===
-The //xopZ// gene was shown to be expressed in a //hrpG//-dependent manner. Presence of a PIP box (TTCTC-N15-TTCGC) 58 bp upstream of the predicted translation start site (Song //et al//., 2010).
-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 //xopZ//, 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 ===
-PXO99<sup>A</sup>  contains two identical copies of the gene due to a duplication of 212 kb in the genome. However, a deletion of one //xopZ// gene did not affect pathogenicity or bacterial growth in plants, while strains with mutations in both copies of //xopZ<sub>PXO99</sub> // displayed reduced virulence in terms of lesion length and bacterial multiplication compared with the wild type strain PXO99<sup>A</sup>  . The introduction of one genomic copy of //xopZ<sub>PXO99</sub> // restores the mutant to full virulence. To test whether XopZ<sub>PXO99</sub> inhibits the host cell-wall-associated defense responses (PTI), leaves of //Nicotiana benthamiana// were infiltrated with //Agrobacterium// cells with and without //xopZ<sub>PXO99</sub> // under the control of the Cauliflower mosaic virus 35S promoter 24 hours preceding inoculation of the same leaves with a T3SS mutant of PXO99<sup>A</sup>  (ME7). Twenty-four hours after inoculation, leaves inoculated with ME7 had more callose depositions than the leaves inoculated with //Agrobacterium //spp. expressing //xopZ<sub>PXO99</sub> //. This results suggesting a role for XopZ<sub>PXO99</sub> in interfering with host innate immunity (PTI) during //X. oryzae// pv. //oryzae// infection (Song //et al//., 2010). Besides, Western blot analysis with p44/42 MAP kinase antibody clearly showed that XopN, XopV and XopZ inhibited the peptidoglycan(PNG)-induced phosphorylation of OsMAPKs. Expression of all Xop effectors were verified by immunoblotting with anti-HA antibody. Thus, expression of three Xop effectors from PXO99<sup>A</sup>  in rice protoplasts results in compromised OsMAPK activation induced by PGN, highlighting their putative virulence functions during pathogenesis (Zhou //et al//., 2018).
-=== Localization ===
-XopZ<sub>PXO99</sub> localizes in the cytoplasm and nucleus of the plant cell (Zhou //et al//., 2015).
-=== Enzymatic function ===
-XopZ<sub>PXO99</sub> functions as a suppressor of LipA-induced innate immune responses since the mutation of //XopZ// partially compromises virulence while quadruple mutant of //xopN/xopQ/xopX/xopZ// induces calloses deposition just similarly to //Xoo// T3SS-mutant in rice leaves (Sinha //et al//., 2013). The function of XopZ is also to stabilize a putative host E3 ubiquitin ligase protein PBP (s-ribonuclease) in the nucleus and prevents its degradation-mediated by a cysteine protease (C1A) in plant cells. XopZ may function to interfere with the homeostatic state of the negative regulator (PBP) in immune system in rice, and subvert the plant immune response (Zhou //et al//., 2015).
-=== Interaction partners ===
-XopZ interacts with a putative host E3 ubiquitin ligase protein PBP (s-ribonuclease) //in vitro// and //in vivo//. Regions containing 193 aa - 225 aa of PBP is required for interacting with XopZ. PBP is a negative regulator of host immune response based on the disease phenotype in PBP-knockout rice plants. C1A directly interacts and strongly degrades PBP through its cysteine protease activity, leading to a homeostatic state of PBP in plant cells (Zhou //et al//., 2015).
-===== Conservation =====
-=== In xanthomonads ===
-Yes, is found to be conserved in all //Xanthomonas //spp. (whose genomes have been sequenced) with the exception of some clade-1 strains (//e.g.// //X. albilineans//) (Song //et al//., 2010; Sinha //et al//., 2013).
-=== In other plant pathogens/symbionts ===
-Related genes are also found in several //Pseudomonas syringae// pathovars (HopAs1 relatives), a few strains of //Ralstonia solanacearum// (AWR proteins), and the AAC00-1 strain of //Acidovorax avenae// subsp. //citrulli// (Song //et al//., 2010).
-===== 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]] FIXME  Information needs to be added to the profile.
-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]] FIXME  Information needs to be added to the profile.
-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]] FIXME  Information needs to be added to the profile.
-Potnis N, Krasileva K, Chow V, Almeida NF, Patil PB, Ryan RP, Sharlach M, Behlau F, Dow JM, Momol M, White FF, Preston JF, Vinatzer BA, Koebnik R, Setubal JC, Norman DJ, Staskawicz BJ, Jones JB (2011). Comparative genomics reveals diversity among xanthomonads infecting tomato and pepper. BMC Genomics 12: 146. DOI: [[https://doi.org/10.1186/1471-2164-12-146|10.1186/1471-2164-12-146]] FIXME  Information needs to be added to the profile (first description of XopZ2).
-Ryan RP, Koebnik R, Szurek B, Boureau T, Bernal A, Bogdanove A, Dow JM (2009). Passing GO (gene ontology) in plant pathogen biology: a report from the //Xanthomonas// Genomics Conference. Cell. Microbiol. 11: 1689-1696. DOI: [[https://doi.org/10.1111/j.1462-5822.2009.01387.x|10.1111/j.1462-5822.2009.01387.x]]
-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.0075867]]
-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]]
-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]]
-Zhou J (2015). Host target genes of the //Xanthomonas oryzae// pv. //oryzae// type III effectors for bacterial blight in rice. Doctoral Thesis, Iowa State University, USA. PDF: [[https://lib.dr.iastate.edu/etd/14469/|lib.dr.iastate.edu/etd/14469/]]

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