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bacteria:t3e:avrbs2 [2020/07/03 08:59] – rkoebnikbacteria:t3e:avrbs2 [2025/02/21 11:40] (current) – joana_costa
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-====== AvrBs2 ======+====== The Type III Effector AvrBs2 from //Xanthomonas// ======
  
 Author: [[https://www.researchgate.net/profile/Spela_Alic|Špela Alič]]\\ Author: [[https://www.researchgate.net/profile/Spela_Alic|Špela Alič]]\\
-Internal reviewer: [[https://www.researchgate.net/profile/Ralf_Koebnik|Ralf Koebnik]]\\ +Internal reviewer: [[https://www.researchgate.net/profile/Ralf_Koebnik|Ralf Koebnik]]
-Expert reviewer: FIXME+
  
 Class: AvrBs2\\ Class: AvrBs2\\
 Protein family: AvrBs2\\ Protein family: AvrBs2\\
-Prototype: AvrBs2 (//Xanthomonas euvesicatoria// pv. //euvesicatoria// aka //Xanthomonas campestris// pv. //vescicatoria//; strain 85-10)\\ +Prototype: AvrBs2 (//Xanthomonas euvesicatoria// pv. //euvesicatoria//, ex //Xanthomonas campestris// pv. //vesicatoria//; strain 75-3)\\ 
-RefSeq ID: [[https://www.ncbi.nlm.nih.gov/protein/WP_011345810.1|WP_011345810.1]] (714 aa)\\+GenBank ID: [[https://www.ncbi.nlm.nih.gov/protein/AAD11434.1|AAD11434.1]] (714 aa)\\ 
 +RefSeq ID: [[https://www.ncbi.nlm.nih.gov/protein/WP_039418336.1|WP_039418336.1]] (729 aa)\\ 
 +Synonym: AvrRxc1/3 (Ignatov //et al.//, 2002)\\
 3D structure: Unknown 3D structure: Unknown
  
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 === (Experimental) evidence for being a T3E === === (Experimental) evidence for being a T3E ===
  
-AvrBs2 fused to the calmodulin-activated adenylate cyclase domain was shown to translocate into plant cells (cytosol), detected through rise of cAMP levels inside the plant tissue. The //hrpF// <sup>-</sup>  mutant was used as a negative control to prove the translocation process. Further it was shown that AvrBs2 contains two N-terminal secretion and translocation signals: first for secretion and the second for enhancing translocation (Casper-Lindley //et al//., 2002).+Mary Beth Mudgett and coworkers provided the first evidence that AvrBs2 is secreted from //Xanthomonas campestis// pv. //vesicatoria// (//Xcv//) and that secretion is type III (//hrp//) dependent (Mudgett //et al.//, 2000). N- and C-terminal deletion analyses of AvrBs2 identified the effector domain of AvrBs2 that is recognized by //Bs2// pepper plants. By using a truncated //Pseudomonas syringae// AvrRpt2 effector reporter devoid of type III signal sequences, they localized the minimal region of AvrBs2 that is required for type III secretion in //Xcv.// Furthermore, they identified the region of AvrBs2 that is required for both type III secretion and translocation to host plants (Mudgett //et al.//, 2000). The mapping of AvrBs2 sequences sufficient for type III delivery also revealed the presence of a potential mRNA secretion signal (Mudgett //et al.//, 2000), a hypothesis that was first put forward by the lab of Olaf Schneewind (Anderson & Schneewind, 1997) and that provoked controversies over the years to come (Ghosh, 2004; Habyarimana & Ahmer, 2013). 
 + 
 +Type III-dependent translocation of AvrBs2 was later confirmed using the calmodulin-dependent adenylate cyclase domain (Cya) of the //Bordetella pertussis// cyclolysin as a reporter (Casper-Lindley //et al//., 2002). Effector translocation into plant cells (cytosol) was detected through rise of cAMP levels inside the plant tissue of pepper plants. Mutants and //hrcV// and //hrpF// were used as a negative controls to prove that the secretion and translocation process depends on the type III secretion system (Casper-Lindley //et al//., 2002). Further it was shown that AvrBs2 contains two N-terminal secretion and translocation signals: the first for secretion and the second for enhancing translocation (Casper-Lindley //et al//., 2002). 
 + 
 +Once the effector domain of AvrBs2 that is recognized by //Bs2// pepper plants was identified (Mudgett //et al.//, 2000), this knowledge was used to construct a Tn//5//-based reporter transposon, which was sucessfully used in genetic screens to isolate type III effectors from //Xanthomonas// (Roden //et al.//, 2004).
 === Regulation === === Regulation ===
  
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 === Phenotypes === === Phenotypes ===
  
-  * AvrBs2 has been demonstrated to be required for full virulence of //X. euvesicatoria//  pv. //euvesicatoria//  (aka //X. campestris//  pv. //vesicatoria//), //X. oryzae//  pv. //oryzicola//, //X. phaseoli //pv. //manihotis//  (aka //X. axonopodis//  pv. //manihotis//) (Zhao //et al//., 2011; Li //et al//., 2015; Mutka //et al.//, 2016; Medina //et al//., 2018).+  * The loss of a functional //avrBs2//  gene was found to affect the fitness of //Xcv//  and revealed fitness costs for three additional, plasmid-borne effector genes (//avrBs1//, //avrBs3//, //avrBs4//) in //Xcv//, indicating that complex functional interactions exist among effector genes (Wichmann & Bergelson, 2004). 
 +  * AvrBs2 has been demonstrated to be required for full virulence of //Xcv//, //X. oryzae//  pv. //oryzicola//, //X. phaseoli//  pv. //manihotis//  (aka //X. axonopodis//  pv. //manihotis//) (Zhao //et al//., 2011; Li //et al//., 2015; Mutka //et al.//, 2016; Medina //et al//., 2018).
   * Recognition of //AvrBs2//  by OsHRL makes rice more resistant against //X. oryzae//  pv. //oryzicola//  (Park //et al//., 2010).   * Recognition of //AvrBs2//  by OsHRL makes rice more resistant against //X. oryzae//  pv. //oryzicola//  (Park //et al//., 2010).
 +  * It was shown in pepper and tomato lines without //Bs2//  that mutations of catalytic residues in the glycerolphosphodiesterase did not interfere with the ability of the plant to recognize AvrBs2 through the cognate R gene //Bs2//  and trigger disease resistance. This finding suggests that recognition of AvrBs2 is independent of its glycerolphosphodiesterase enzyme activity (Zhao //et al//., 2011).
 +  * AvrBs2 contributes to //X. oryzae//  pv. //oryzicola//  virulence by suppressing PAMP-triggered defense responses in rice (Li //et al//., 2015).
 +  * AvrBs2 transiently expressed in //Arabidopsis//  protoplasts suppressed flg22-induced NHO1 expression (Li //et al//., 2015).
 +  * Induced expression of AvrBs2 in transgenic cell cultures was shown to dramatically suppress flg22-induced and chitin-induced immune responses, such as ROS burst and PR gene expression (Li //et al//., 2015).
 +  * A ∆//xopK//  mutant strain of //Xanthomonas phaseoli//  pv. //manihotis//  showed reduced growth in planta and delayed spread through the vasculature system of cassava. Moreover, the ∆//avrBs2//  mutant strain exhibited reduced water-soaking symptoms at the site of inoculation (Mutka //et al.//, 2016).
 +  * XopN and AvrBS2 were shown to significantly contribute to virulence of //X. oryzae//  pv. //oryzicola//  (Xoc GX01) (Liao //et al.//, 2020).
 +
 +=== Localization ===
 +
 +The //avrBs2//  gene is chromosomal (Coplin, 1989). The AvrBs2 protein is translocated from bacterial cells into the plant cytosol. Subcellular localization of AvrBs2 was demonstrated using recombinant AvrBs2::GFP reporter fusions transiently expressed in rice protoplasts. Green fluorescence of AvrBs2::GFP was detected across the entire cell. Similar subcellular localization was observed in //Nicotiana benthamiana//  (Li //et al//., 2015).
 +
 +=== Enzymatic function ===
 +
 +Shown activity of glycerolphosphodiesterase catalytic site //in-vitro//; agrocinopine synthase activity predicted but has not been experimentally confirmed (Zhao //et al//., 2011).
 +
 +=== Interaction partners ===
 +
 +Gene-for-gene relationship with corresponding resistance gene //Bs2//  (Minsavage //et al//., 1990). Furthermore, interaction between AvrBs2 and OsHRL was experimentaly shown by yeast two-hybrid screening (Park //et al//., 2010).
 +
 +===== Conservation =====
 +
 +=== In xanthomonads ===
 +
 +Yes (//e.g.//, //X//. //arboricola//, //X//. //campestris//, //X//. //citri//, //X. euvesicatoria//, //X//. //fuscans//, //X. oryzae//, //X//. //phaseoli//).
 +
 +=== In other plant pathogens/symbionts ===
 +
 +No report.
 +
 +===== Biological function =====
 +
 +=== How discovered? ===
 +
 +Indirectly – the pathovars that induced //Bs2//-mediated hypersensitivity were classified as having AvrBs2 activity (Kearney & Staskawicz, 1990).
 +
 +=== (Experimental) evidence for being a T3E ===
 +
 +AvrBs2 fused to the calmodulin-activated adenylate cyclase domain was shown to translocate into plant cells (cytosol), detected through rise of cAMP levels inside the plant tissue. The //hrpF//  <sup>-</sup>   mutant was used as a negative control to prove the translocation process. Further it was shown that AvrBs2 contains two N-terminal secretion and translocation signals: first for secretion and the second for enhancing translocation (Casper-Lindley //et al//., 2002).
 +
 +=== Regulation ===
 +
 +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 //avrBs2//, 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).
 +
 +Transcriptome analysis (RNA-seq) and qRT-PCR have shown that //avrBs2//  gene expression is downregulated in a //X. citri//  pv. //citri//  Δ//phoP//  mutant, indicating that PhoP is a positive regulator of //avrBs2//  expression (Wei //et al//., 2019).
 +
 +=== Phenotypes ===
 +
 +  * AvrBs2 has been demonstrated to be required for full virulence of //X. euvesicatoria//  pv. //euvesicatoria//  (aka //X. campestris//  pv. //vesicatoria//), //X. oryzae//  pv. //oryzicola//, //X. phaseoli//  pv. //manihotis//  (aka //X. axonopodis//  pv. //manihotis//) (Zhao //et al//., 2011; Li //et al//., 2015; Mutka //et al.//, 2016; Medina //et al//., 2018).
 +  * Recognition of AvrBs2 by OsHRL makes rice more resistant against //X. oryzae//  pv. //oryzicola//  (Park //et al//., 2010).
   * It was shown in pepper and tomato lines without //Bs2 //that mutations of catalytic residues in the glycerolphosphodiesterase did not interfere with the ability of the plant to recognize AvrBs2 through the cognate R gene //Bs2//  and trigger disease resistance. This finding suggests that recognition of AvrBs2 is independent of its glycerolphosphodiesterase enzyme activity (Zhao //et al//., 2011).   * It was shown in pepper and tomato lines without //Bs2 //that mutations of catalytic residues in the glycerolphosphodiesterase did not interfere with the ability of the plant to recognize AvrBs2 through the cognate R gene //Bs2//  and trigger disease resistance. This finding suggests that recognition of AvrBs2 is independent of its glycerolphosphodiesterase enzyme activity (Zhao //et al//., 2011).
   * AvrBs2 contributes to //X. oryzae//  pv. //oryzicola//  virulence by suppressing PAMP-triggered defense responses in rice (Li //et al//., 2015).   * AvrBs2 contributes to //X. oryzae//  pv. //oryzicola//  virulence by suppressing PAMP-triggered defense responses in rice (Li //et al//., 2015).
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   * Induced expression of AvrBs2 in transgenic cell cultures was shown to dramatically suppress flg22-induced and chitin-induced immune responses, such as ROS burst and PR gene expression (Li //et al//., 2015).   * Induced expression of AvrBs2 in transgenic cell cultures was shown to dramatically suppress flg22-induced and chitin-induced immune responses, such as ROS burst and PR gene expression (Li //et al//., 2015).
   * A ∆//xopK//  mutant strain of //Xanthomonas phaseoli//  pv. //manihotis//  (aka //Xanthomonas axonopodis//  pv. //manihotis//) showed reduced growth in planta and delayed spread through the vasculature system of cassava. Moreover, the ∆avrBs2 mutant strain exhibited reduced water-soaking symptoms at the site of inoculation (Mutka //et al.//, 2016).   * A ∆//xopK//  mutant strain of //Xanthomonas phaseoli//  pv. //manihotis//  (aka //Xanthomonas axonopodis//  pv. //manihotis//) showed reduced growth in planta and delayed spread through the vasculature system of cassava. Moreover, the ∆avrBs2 mutant strain exhibited reduced water-soaking symptoms at the site of inoculation (Mutka //et al.//, 2016).
 +  * //Agrobacterium//-mediated transient expression of both XopQ and XopX in rice cells resulted in induction of rice immune responses, which were not observed when either protein was individually expressed. A screen for //Xanthomonas//  effectors which can suppress XopQ-XopX induced rice immune responses, led to the identification of five effectors, namely XopU, XopV, XopP, XopG and AvrBs2, that could individually suppress these immune responses. These results suggest a complex interplay of //Xanthomonas//  T3SS effectors in suppression of both pathogen-triggered immunity and effector-triggered immunity to promote virulence on rice (Deb //et al.//, 2020).
 === Localization === === Localization ===
  
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 === In xanthomonads === === In xanthomonads ===
  
-Yes (//e.g.//, //X//. //arboricola//, //X//. //campestris//, //X//. //citri//, //X. euvesicatoria//, //X//. //fuscans//, X. oryzae, //X//. //phaseoli//).+Yes (//e.g.//, //X//. //arboricola//, //X//. //campestris//, //X//. //citri//, //X. euvesicatoria//, //X//. //fuscans//, //X. oryzae//, //X//. //phaseoli//). 
 + 
 +Field strains of //X. euvesicatoria//  pv. //euvesicatoria//  and //X//. //campestris//  pv. //campestris//  were found to accumulate mutations in the //avrBs2/////avrRxc1/3//  gene in order to overcome //Bs2/////Rxc1/////Rxc3//-mediated resistance (Swords //et al.//, 1996; Gassmann //et al.//, 2000; Ignatov //et al.//, 2002). Yet, the global //Xcv//  population was found to be extremely clonal, with very little genetic variation throughout the chromosome, including //avrBs2//  and the plasmid-borne //avrBs1//, a finding that is consistent with recent evolution or population expansion of the species (Wichmann //et al.//, 2005).
  
 === In other plant pathogens/symbionts === === In other plant pathogens/symbionts ===
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 ===== References ===== ===== References =====
 +
 +Anderson DM, Schneewind O (1997). A mRNA signal for the type III secretion of Yop proteins by //Yersinia enterocolitica//. Science 278: 1140-1143. DOI: [[https://doi.org/10.1126/science.278.5340.1140|10.1126/science.278.5340.1140]]
  
 Casper-Lindley C. Dahlbeck D, Clark ET, Staskawicz BJ (2002). Direct biochemical evidence for type III secretion-dependent translocation of the AvrBs2 effector protein into plant cells. Proc. Natl. Acad. Sci. USA 99: 8336-8341. DOI: [[https://doi.org/10.1073/pnas.122220299|10.1073/pnas.122220299]] Casper-Lindley C. Dahlbeck D, Clark ET, Staskawicz BJ (2002). Direct biochemical evidence for type III secretion-dependent translocation of the AvrBs2 effector protein into plant cells. Proc. Natl. Acad. Sci. USA 99: 8336-8341. DOI: [[https://doi.org/10.1073/pnas.122220299|10.1073/pnas.122220299]]
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 Coplin DL (1989). Plasmids and their role in the evolution of plant pathogenic bacteria. Ann. Rev. Phytopathol. 27: 187-212. DOI: [[https://doi.org/10.1146/annurev.py.27.090189.001155|10.1146/annurev.py.27.090189.001155]] Coplin DL (1989). Plasmids and their role in the evolution of plant pathogenic bacteria. Ann. Rev. Phytopathol. 27: 187-212. DOI: [[https://doi.org/10.1146/annurev.py.27.090189.001155|10.1146/annurev.py.27.090189.001155]]
  
-Gassmann W, Dahlbeck D, Chesnokova O, Minsavage GV, Jones JB, Staskawicz BJ (2000). Molecular evolution of virulence in natural field strains of //Xanthomonas campestris//  pv. //vesicatoria//. J. Bacteriol. 182: 7053-7059. DOI: [[https://doi.org/10.1128/jb.182.24.7053-7059.2000|10.1128/jb.182.24.7053-7059.2000]] FIXME+Deb S, Ghosh P, Patel HK, Sonti RV (2020). Interaction of the //Xanthomonas//  effectors XopQ and XopX results in induction of rice immune responses. Plant J. 104: 332-350. DOI: [[https://doi.org/10.1111/tpj.14924|10.1111/tpj.14924]] 
 + 
 +Gassmann W, Dahlbeck D, Chesnokova O, Minsavage GV, Jones JB, Staskawicz BJ (2000). Molecular evolution of virulence in natural field strains of //Xanthomonas campestris//  pv. //vesicatoria//. J. Bacteriol. 182: 7053-7059. DOI: [[https://doi.org/10.1128/jb.182.24.7053-7059.2000|10.1128/jb.182.24.7053-7059.2000]] 
 + 
 +Ghosh P (2004). Process of protein transport by the type III secretion system. Microbiol. Mol. Biol. Rev. 68: 771-795. DOI: [[https://doi.org/10.1128/MMBR.68.4.771-795.2004|10.1128/MMBR.68.4.771-795.2004]] 
 + 
 +Habyarimana F, Ahmer BM (2013). More evidence for secretion signals within the mRNA of type 3 secreted effectors. J. Bacteriol. 195: 2117-2118. DOI: [[https://doi.org/10.1128/JB.00303-13|10.1128/JB.00303-13]] 
 + 
 +Ignatov AN, Monakhos GF, Dzhalilov FS, Pozmogova GV (2002). Avirulence gene from //Xanthomonas campestris//  pv. //campestris//  homologous to the //avrBs2//  locus is recognized in race-specific reaction by two different resistance genes in Brassicas. Genetika 38: 1656-1662 [Article in Russian] / Russian J. Genet. 38: 1404-1410. DOI: [[https://doi.org/10.1023/A:1021643907032|10.1023/A:1021643907032]]
  
 Kearney B, Staskawicz BJ (1990). Widespread distribution and fitness contribution of //Xanthomonas campestris//  avirulence gene //avrBs2//. Nature 346: 385-386. DOI: [[https://doi.org/10.1038/346385a0|10.1038/346385a0]] Kearney B, Staskawicz BJ (1990). Widespread distribution and fitness contribution of //Xanthomonas campestris//  avirulence gene //avrBs2//. Nature 346: 385-386. DOI: [[https://doi.org/10.1038/346385a0|10.1038/346385a0]]
  
 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]] 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]]
 +
 +Liao ZX, Li JY, Mo XY, Ni Z, Jiang W, He YQ, Huang S (2020). Type III effectors //xopN//  and //avrBS2//  contribute to the virulence of //Xanthomonas oryzae//  pv. //oryzicola//  strain GX01. Res. Microbiol. 171: 102-106. DOI: [[https://doi.org/10.1016/j.resmic.2019.10.002|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]] 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]]
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 Minsavage GV, Dahlbeck D, Whalen MC, Kearney B, Bonas U, Staskawicz BJ, Stall RE (1990). Gene-for-gene relationships specifying disease resistance in //Xanthomonas campestris//  pv. //vesicatoria//-pepper interactions. Mol. Plant Microbe Interact. 3: 41-47. DOI: [[https://doi.org/10.1094/MPMI-3-041|10.1094/MPMI-3-041]] Minsavage GV, Dahlbeck D, Whalen MC, Kearney B, Bonas U, Staskawicz BJ, Stall RE (1990). Gene-for-gene relationships specifying disease resistance in //Xanthomonas campestris//  pv. //vesicatoria//-pepper interactions. Mol. Plant Microbe Interact. 3: 41-47. DOI: [[https://doi.org/10.1094/MPMI-3-041|10.1094/MPMI-3-041]]
  
-Mudgett MB, Chesnokova O, Dahlbeck D, Clark ET, Rossier O, Bonas U, Staskawicz BJ (2000). Molecular signals required for type III secretion and translocation of the //Xanthomonas campestris//  AvrBs2 protein to pepper plants. Proc. Natl. Acad. Sci. USA 97: 13324-13329. DOI: [[https://doi.org/10.1073/pnas.230450797|10.1073/pnas.230450797]] FIXME+Mudgett MB, Chesnokova O, Dahlbeck D, Clark ET, Rossier O, Bonas U, Staskawicz BJ (2000). Molecular signals required for type III secretion and translocation of the //Xanthomonas campestris//  AvrBs2 protein to pepper plants. Proc. Natl. Acad. Sci. USA 97: 13324-13329. DOI: [[https://doi.org/10.1073/pnas.230450797|10.1073/pnas.230450797]]
  
 Mutka AM, Fentress SJ, Sher JW, Berry JC, Pretz C, Nusinow DA, Bart R (2016). Quantitative, image-based phenotyping methods provide insight into spatial and temporal dimensions of plant disease. Plant Physiol. 172: 650-660. DOI: [[https://doi.org/10.1104/pp.16.00984|10.1104/pp.16.00984]] Mutka AM, Fentress SJ, Sher JW, Berry JC, Pretz C, Nusinow DA, Bart R (2016). Quantitative, image-based phenotyping methods provide insight into spatial and temporal dimensions of plant disease. Plant Physiol. 172: 650-660. DOI: [[https://doi.org/10.1104/pp.16.00984|10.1104/pp.16.00984]]
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 Park SR, Moon SJ, Shin DJ, Kim MG, Hwang DJ, Bae SC, Kim JG , Yi BY, Byun MO (2010). Isolation and characterization of rice //OsHRL//  gene related to bacterial blight resistance. Plant Pathol. J. 26: 417-420. DOI: [[https://doi.org/10.5423/PPJ.2010.26.4.417|10.5423/PPJ.2010.26.4.417]] Park SR, Moon SJ, Shin DJ, Kim MG, Hwang DJ, Bae SC, Kim JG , Yi BY, Byun MO (2010). Isolation and characterization of rice //OsHRL//  gene related to bacterial blight resistance. Plant Pathol. J. 26: 417-420. DOI: [[https://doi.org/10.5423/PPJ.2010.26.4.417|10.5423/PPJ.2010.26.4.417]]
  
-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]] FIXME+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]]
  
-Swords KM, Dahlbeck D, Kearney B, Roy M, Staskawicz BJ (1996). Spontaneous and induced mutations in a single open reading frame alter both virulence and avirulence in //Xanthomonas campestris//  pv. //vesicatoria//  //avrBs2//. J. Bacteriol. 178: 4661-4669. DOI: [[https://doi.org/10.1128/jb.178.15.4661-4669.1996|10.1128/jb.178.15.4661-4669.1996]] FIXME +Swords KM, Dahlbeck D, Kearney B, Roy M, Staskawicz BJ (1996). Spontaneous and induced mutations in a single open reading frame alter both virulence and avirulence in //Xanthomonas campestris//  pv. //vesicatoria//  //avrBs2//. J. Bacteriol. 178: 4661-4669. DOI: [[https://doi.org/10.1128/jb.178.15.4661-4669.1996|10.1128/jb.178.15.4661-4669.1996]]
- +
-Timilsina S, Abrahamian P, Potnis N, Minsavage GV, White FF, Staskawicz BJ, Jones JB, Vallad GE, Goss EM (2016). Analysis of sequenced genomes of Xanthomonas perforans identifies candidate targets for resistance breeding in tomato. Phytopathology 106: 1097-1104. DOI: [[https://doi.org/10.1094/PHYTO-03-16-0119-FI|10.1094/PHYTO-03-16-0119-FI]] FIXME+
  
 Wei C, Ding T, Chang C, Yu C, Li X, Liu Q (2019). Global regulator PhoP is necessary for motility, biofilm formation, exoenzyme production and virulence of //Xanthomonas citri//  subsp. //citri//  on citrus plants. Genes 10: 340. DOI: [[https://doi.org/10.3390/genes10050340|10.3390/genes10050340]] Wei C, Ding T, Chang C, Yu C, Li X, Liu Q (2019). Global regulator PhoP is necessary for motility, biofilm formation, exoenzyme production and virulence of //Xanthomonas citri//  subsp. //citri//  on citrus plants. Genes 10: 340. DOI: [[https://doi.org/10.3390/genes10050340|10.3390/genes10050340]]
  
-Wichmann G, Bergelson J (2004). Effector genes of //Xanthomonas axonopodis//  pv. //vesicatoria//  promote transmission and enhance other fitness traits in the field. Genetics 166: 693-706. DOI: [[https://doi.org/10.1534/genetics.166.2.693|10.1534/genetics.166.2.693]] FIXME+Wichmann G, Bergelson J (2004). Effector genes of //Xanthomonas axonopodis//  pv. //vesicatoria//  promote transmission and enhance other fitness traits in the field. Genetics 166: 693-706. DOI: [[https://doi.org/10.1534/genetics.166.2.693|10.1534/genetics.166.2.693]]
  
-Wichmann G, Ritchie D, Kousik CS, Bergelson J (2005). Reduced genetic variation occurs among genes of the highly clonal plant pathogen //Xanthomonas axonopodis//  pv. //vesicatoria//, including the effector gene //avrBs2//. Appl. Environ. Microbiol. 71: 2418-2432. DOI: [[https://doi.org/10.1128/AEM.71.5.2418-2432.2005|10.1128/AEM.71.5.2418-2432.2005]] FIXME+Wichmann G, Ritchie D, Kousik CS, Bergelson J (2005). Reduced genetic variation occurs among genes of the highly clonal plant pathogen //Xanthomonas axonopodis//  pv. //vesicatoria//, including the effector gene //avrBs2//. Appl. Environ. Microbiol. 71: 2418-2432. DOI: [[https://doi.org/10.1128/AEM.71.5.2418-2432.2005|10.1128/AEM.71.5.2418-2432.2005]]
  
 Zhao B, Dahlbeck D, Krasileva KV, Fong RW, Staskawicz BJ (2011). Computational and biochemical analysis of the //Xanthomonas//  effector AvrBs2 and its role in the modulation of //Xanthomonas//  type three effector delivery. PLoS Pathog. 7: e1002408. DOI: [[https://doi.org/10.1371/journal.ppat.1002408|10.1371/journal.ppat.1002408]] Zhao B, Dahlbeck D, Krasileva KV, Fong RW, Staskawicz BJ (2011). Computational and biochemical analysis of the //Xanthomonas//  effector AvrBs2 and its role in the modulation of //Xanthomonas//  type three effector delivery. PLoS Pathog. 7: e1002408. DOI: [[https://doi.org/10.1371/journal.ppat.1002408|10.1371/journal.ppat.1002408]]
 +
 +===== Further reading =====
 +
 +Timilsina S, Abrahamian P, Potnis N, Minsavage GV, White FF, Staskawicz BJ, Jones JB, Vallad GE, Goss EM (2016). Analysis of sequenced genomes of //Xanthomonas perforans//  identifies candidate targets for resistance breeding in tomato. Phytopathology 106: 1097-1104. DOI: [[https://doi.org/10.1094/PHYTO-03-16-0119-FI|10.1094/PHYTO-03-16-0119-FI]]. Corrected in: Phytopathology (2019) 109: 1820. DOI: [[https://doi.org/10.1094/PHYTO-03-16-0119.1-FI|10.1094/PHYTO-03-16-0119.1-FI]]
 +
 +===== Acknowledgements =====
 +
 +This fact sheet is based upon work from COST Action CA16107 EuroXanth, supported by COST (European Cooperation in Science and Technology).
  
bacteria/t3e/avrbs2.1593763153.txt.gz · Last modified: 2023/01/09 10:20 (external edit)

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