猕猴桃细菌性溃疡病菌T3SS关键效应蛋白基因致病功能

doi:10.3864/j.issn.0578-1752.2022.03.007

Abstract

Abstract: 【Objective】 Pseudomonas syringae pv. actinidiae (Psa), the causal agent of bacterial canker of kiwifruit, is the most devastating pathogen in global kiwifruit production. The pathogenic bacteria secrete a series of effectors (T3SEs) into host cell to promote infection and pathogenesis by the type III secretion system (T3SS). The objective of this study is to analyze the T3SEs information in Psa genome and systematically evaluate the pathogenicity function of T3SS and T3SEs, and to provide the basis evidence for the research of the pathogenic mechanism and the establishment of the control strategies.【Method】 By marker-free homologous recombination knockout technique, the M228 deficiency mutants of T3SS, ΔhrcS and ΔhrcC, were obtained for inoculating on host to evaluate the pathogenicity and injecting on Nicotiana benthamiana to observe the cell death response. Then, based on the T3SEs database downloaded from Pseudomonas-Plant Interaction, the T3SEs library of strong pathogenicity M228 and weak pathogenicity M227 was separately constructed against the database by local BLAST multiple sequence alignment program, and then the T3SEs information between them was compared. Moreover, 20 T3SE single- and poly-genetic mutants from M228 and 2 HopR1-genetic complementing mutants were constructed, involving 19 T3SEs, and then the mutants were wound-inoculated on kiwifruit canes for assessing and statistical analyzing the pathogenicity.【Result】 T3SS was proved to be essential for Psa pathogenicity on host and hypersensitive response (HR) on non-host by the hrcS and hrcC mutants, separately. Further the BLAST results against database showed there were almost 31 complete T3SE genes and their sequences were displayed 100% similarity between the strong pathogenicity strain and attenuated strain. Then, some T3SE genes were selected for deletion mutants. The results showed that hopM1/avrE1 and hopR1 genes were essential for Psa pathogenicity and had no function redundant with each other. In addition, the avrPto5- and avrRpm1-deletion mutant could in turn increase the Psa pathogenicity. Based on avrPto- and T3SE group (cluster A, E and F) deletion mutant, single- or poly-genetic mutant of hopR1 and hopM1/avrE1 could still separately lead to a significant decrease in Psa pathogenicity. However, simultaneous deletion of hopM1/avrE1, hopR1, avrPto5 and A-F-E cluster resulted in complete loss of pathogenicity.【Conclusion】 HopR1 and its homologous family HopM1/AvrE1, which don’t have a redundant function independent with others, are the unique key pathogenicity factors in Psa, but AvrPto5-and AvrRpm1-deletion can enhance Psa pathogenicity.

Key words: bacterial canker of kiwifruit, Pseudomonas syringae pv. actinidiae (Psa), effector, T3SS, pathogenicity

ZHANG JinLong,ZHAO ZhiBo,LIU Wei,HUANG LiLi. The Function of Key T3SS Effectors in Pseudomonas syringae pv. actinidiae[J].Scientia Agricultura Sinica, 2022, 55(3): 503-513.

Figures/Tables 7

Table 1

The list of the detail information of deletion and complement mutants"

突变体名称 The name of mutants	详细信息 Related detail information
ΔhrcC	T3SS结构基因hrcC功能缺陷突变体 hrcC-deletion generating the T3SS-deficient mutant
ΔhrcS	T3SS结构基因hrcS功能缺陷突变体hrcS-deletion generating the T3SS-deficient mutant
ΔO	T3SE hopM1/avrE1缺失突变体T3SE hopM1/avrE1-deletion mutant
ΔM	T3SE hopR1缺失突变体T3SE hopR1-deletion mutant
ΔOM	T3SE hopM1/avrE1+hopR1缺失突变体T3SE hopR1-deletion mutant based on ΔO background
ΔMO	T3SE hopR1+hopM1/avrE1缺失突变体T3SE hopM1/avrE1-deletion mutant based on ΔM background
ΔM-C-hopR1	T3SE hopR1突变体基础上回补hopR1的回补菌株T3SE hopR1-complement mutant based on ΔM background
ΔOM-C-hopR1	T3SE hopR1+hopM1/AvrE1突变体基础上回补hopR1的回补菌株T3SE hopR1-complement mutant based on ΔOM background
ΔA	表2中对应T3SE基因簇A缺失突变体T3SE cluster A-deletion mutant listing on Table 2
ΔF	表2中对应T3SE基因簇F缺失突变体T3SE cluster F-deletion mutant listing on Table 2
ΔE	表2中对应T3SE基因簇E缺失突变体T3SE cluster E-deletion mutant listing on Table 2
ΔAF	表2中对应T3SE基因簇A和F缺失突变体T3SE cluster A- and F-deletion mutant listing on Table 2
ΔAFE	表2中对应T3SE基因簇A、F和E全敲除缺失突变体T3SE cluster A-, F- and E-deletion mutant listing on Table 2
ΔAFEL	在ΔAFE突变体的背景下敲除avrPto5获得突变体avrPto5-deletion mutant based on ΔAFE background
ΔAFELM	在ΔAFEL突变体的背景下敲除hopR1获得突变体hopR1-deletion mutant based on ΔAFEL background
ΔAFELO	在ΔAFEL突变体的背景下敲除hopM1/avrE1获得突变体hopM1/avrE1-deletion mutant based on ΔAFEL background
ΔAFELOM	在ΔAFEL突变体的背景下同时敲除hopM1/avrE1和hopR1获得突变体 hopM1/avrE1- and hopR1-deletion mutant based on ΔAFEL background
ΔOMN	在ΔOM突变体基础上敲除avrRpm1获得突变体avrRpm1-deletion mutant based on ΔOM background

Table 1

Table 2

Fig. 1

Fig. 2

Fig. 3

Fig. 4

Fig. 5

References 36

[1]	秦虎强, 高小宁, 赵志博, 朱穗层, 李建民, 黄丽丽. 陕西猕猴桃细菌性溃疡病田间发生动态和规律. 植物保护学报, 2013, 40(3): 225-230.
	QIN H Q, GAO X N, ZHAO Z B, ZHU H C, LI J M, HUANG L L.The prevalence dynamics and rules of bacterial canker of kiwifruit in Shaanxi. Acta Phytophylacica Sinica, 2013, 40(3): 225-230. (in Chinese)
[2]	高小宁, 赵志博, 黄其玲, 秦虎强, 黄丽丽. 猕猴桃细菌性溃疡病研究进展. 果树学报, 2012, 29(2): 262-268.
	GAO X N, ZHAO Z B, HUANG Q L, QIN H Q, HUANG L L.Advances in research on bacterial canker of kiwifruit. Journal of Fruit Science, 2012, 29(2): 262-268. (in Chinese)
[3]	ZHAO Z, CHEN J, GAO X, ZHANG D, ZHANG J, WEN J, QIN H, GUO M, HUANG L.Comparative genomics reveal pathogenicity- related loci in Pseudomonas syringae pv. actinidiae biovar 3. Molecular Plant Pathology, 2019, 20(7): 923-942. doi: 10.1111/mpp.2019.20.issue-7
[4]	孙思, 牛建军, 王岱. 细菌三型分泌系统效应蛋白转运的研究进展. 微生物学报, 2017, 57(10): 1452-1460.
	SUN S, NIU J J, WANG D.Advances in studies of translocation of effector by bacterial type 3 secretion system. Acta Microbiologica Sinica, 2017, 57(10): 1452-1460. (in Chinese)
[5]	LINDEBERG M, CUNNAC S, COLLMER A.Pseudomonas syringae type III effector repertoires: Last words in endless arguments. Trends in Microbiology, 2012, 20(4): 199-208. doi: 10.1016/j.tim.2012.01.003
[6]	DOS SANTOS A M P,FERRARI R G, CONTE-JUNIOR C A. Type three secretion system in Salmonella typhimurium: The key to infection. Genes and Genomics, 2020, 42(5): 495-506. doi: 10.1007/s13258-020-00918-8
[7]	朱秀秀, 高必达, 赵廷昌, 张月娟. 植物病原细菌Ⅲ型分泌系统及Pseudomonas syringae pv. tomato的信号分子分泌研究进展. 湖南农业科学, 2009(2): 19-22.
	ZHU X X, GAO B D, ZHAO Y C, ZHANG Y J.Research progress of type III secretory system of plant pathogenic bacteria and signal molecule secretion of Pseudomonas syringae pv. tomato. Hunan Agricultural Sciences, 2009(2): 19-22. (in Chinese)
[8]	MARLOVITS T C, KUBORI T, LARA-TEJERO M, THOMAS D, UNGER V M, GALAN J E.Assembly of the inner rod determines needle length in the type III secretion injectisome. Nature, 2006, 441(7093): 637-640. doi: 10.1038/nature04822
[9]	MACHO A P, ZIPFEL C.Plant PRRs and the activation of innate immune signaling. Molecular Cell, 2014, 54(2): 263-272. doi: 10.1016/j.molcel.2014.03.028
[10]	温晶. 猕猴桃溃疡病菌Ⅲ型效应蛋白的筛选及效应蛋白HopX3功能的初步研究[D]. 杨凌: 西北农林科技大学, 2016.
	WEN J.Identification of Psa type III effectors and preliminary analysis of effector HopX3 in pathogenicity[D]. Yangling: Northwest A&F University, 2016. (in Chinese)
[11]	BALTRUS D A, NISHIMURA M T, ROMANCHUK A, CHANG J H,MUKHTAR M S, CHERKIS K, ROACH J, GRANT S R, JONES C D, DANGL J L.Dynamic evolution of pathogenicity revealed by sequencing and comparative genomics of 19 Pseudomonas syringae isolates. PLoS Pathogens, 2011, 7(7): e1002132.
[12]	CUNNAC S, CHAKRAVARTHY S, KVITKO B H, RUSSELL A B, MARTIN G B, COLLMER A.Genetic disassembly and combinatorial reassembly identify a minimal functional repertoire of type III effectors in Pseudomonas syringae. Proceedings of the National Academy of Sciences of the United States of America, 2011, 108(7): 2975-2980.
[13]	KVITKO B H, PARK D H, VELASQUEZ A C, WEI C F, RUSSELL A B, MARTIN G B, SCHNEIDER D J, COLLMER A.Deletions in the repertoire of Pseudomonas syringae pv. tomato DC3000 type III secretion effector genes reveal functional overlap among effectors. PLoS Pathogens, 2009, 5(4): e1000388. doi: 10.1371/journal.ppat.1000388
[14]	TAMPAKAKI A P, SKANDILIS N, GAZI A D, BASTAKI M N, PANAGIOTIS F S, CHAROVA S N, KOKKINIDIS M, PANOPOULOS N J.Playing the “Harp”: Evolution of our understanding of hrp/hrc genes 1. Annual Review of Phytopathology, 2010, 48: 347-370. doi: 10.1146/phyto.2010.48.issue-1
[15]	CHOI S, JAYARAMAN J, SEGONZAC C, PARK H J, PARK H, HAN S W, SOHN K H.Pseudomonas syringae pv. actinidiae type III effectors localized at multiple cellular compartments activate or suppress innate immune responses in Nicotiana benthamiana. Frontiers in Plant Science, 2017, 8: 2157. doi: 10.3389/fpls.2017.02157
[16]	JAYARAMAN J, CHOI S, PROKCHORCHIK M, CHOI D S, SPIANDORE A, RIKKERINK E H, TEMPLETON M D, SEGONZAC C, SOHN K H.A bacterial acetyltransferase triggers immunity in Arabidopsis thaliana independent of hypersensitive response. Scientific Reports, 2017, 7(1): 3557. doi: 10.1038/s41598-017-03704-x
[17]	CHOI S, JAYARAMAN J, SOHN K H.Arabidopsis thaliana SOBER1 (SUPPRESSOR OF AVRBST-ELICITED RESISTANCE 1) suppresses plant immunity triggered by multiple bacterial acetyltransferase effectors. New Phytologist, 2018, 219(1): 324-335. doi: 10.1111/nph.2018.219.issue-1
[18]	YOON M, RIKKERINK E H A. Rpa1 mediates an immune response to avrRpm1_Psa and confers resistance against Pseudomonas syringae pv. actinidiae. The Plant Journal, 2020, 102(4): 688-702.
[19]	JAYARAMAN J, YOON M, APPLEGATE E R, STROUD E A, TEMPLETON M D.AvrE1 and HopR1 from Pseudomonas syringae pv. actinidiae are additively required for full virulence on kiwifruit. Molecular Plant Pathology, 2020, 21(11): 1467-1480. doi: 10.1111/mpp.12989
[20]	赵志博. 猕猴桃细菌性溃疡病菌群体结构与致病机制研究[D]. 杨凌: 西北农林科技大学, 2016.
	ZHAO Z B.Population composition and pathogenetic mechanism in Psuesdomonas syringae pv. actinidiae[D]. Yangling: Northwest A&F University, 2016. (in Chinese)
[21]	WANG K, KANG L, ANAND A, LAZAROVITS G, MYSORE K S.Monitoring in planta bacterial infection at both cellular and whole-plant levels using the green fluorescent protein variant GFPuv. New Phytologist, 2007, 174(1): 212-223. doi: 10.1111/nph.2007.174.issue-1
[22]	KVITKO B H, COLLMER A.Construction of Pseudomonas syringae pv. tomato DC3000 mutant and polymutant strains//Plant Immunity. Methods and Protocols, 2011, 712: 109-128.
[23]	SAWADA H, FUJIKAWA T.Genetic diversity of Pseudomonas syringae pv. actinidiae, pathogen of kiwifruit bacterial canker. Plant Pathology, 2019, 68(7): 1235-1248. doi: 10.1111/ppa.v68.7
[24]	XIN X F, NOMURA K, AUNG K, VELASQUEZ A C, YAO J, BOUTROT F, CHANG J H, ZIPFEL C, HE S Y.Bacteria establish an aqueous living space in plants crucial for virulence. Nature, 2016, 539(7630): 524-529. doi: 10.1038/nature20166
[25]	JIN L, HAM J H, HAGE R, ZHAO W, SOTO-HERNANDEZ J, LEE S Y, PAEK S M, KIM M G, BOONE C, COPLIN D L, MACKEY D.Direct and indirect targeting of PP2A by conserved bacterial type-III effector proteins. PLoS Pathogens, 2016, 12(5): e1005609. doi: 10.1371/journal.ppat.1005609
[26]	DEGRAVE A, SIAMER S, BOUREAU T, BARNY M A.The AvrE superfamily: Ancestral type III effectors involved in suppression of pathogen-associated molecular pattern-triggered immunity. Molecular Plant Pathology, 2015, 16(8): 899-905. doi: 10.1111/mpp.2015.16.issue-8
[27]	PALACE S G, PROULX M K, SZABADY R L, GOGUEN J D.Gain-of-function analysis reveals important virulence roles for the Yersinia pestis type III secretion system effectors YopJ, YopT, and YpkA. Infection and Immunity, 2018, 86(9): e00318-18.
[28]	ÜSTÜN S, KÖNIG P, GUTTMAN D S, BÖRNKE F. HopZ4 from Pseudomonas syringae, a member of the HopZ type III effector family from the YopJ superfamily, inhibits the proteasome in plants. Molecular Plant-Microbe Interactions, 2014, 27(7): 611-623. doi: 10.1094/MPMI-12-13-0363-R
[29]	LEWIS J D, LEE A H Y,HASSAN J A,WAN J,HURLEY B,JHINGREE J R,WANG P W,LO T,YOUN J Y,GUTTMAN D S,DESVEAUX D. The Arabidopsis ZED1 pseudokinase is required for ZAR1-mediated immunity induced by the Pseudomonas syringae type III effector HopZ1a. Proceedings of the National Academy of Sciences of the United States of America, 2013, 110(46): 18722-18727.
[30]	LEWIS J D, LEE A, MA W B, ZHOU H B, GUTTMAN D S, DESVEAUX D.The YopJ superfamily in plant-associated bacteria. Molecular Plant Pathology, 2011, 12(9): 928-937. doi: 10.1111/j.1364-3703.2011.00719.x
[31]	ZHOU H B, LIN J A, JOHNSON A, MORGAN R L, ZHONG W W, MA W B.Pseudomonas syringae type III effector HopZ1 targets a host enzyme to suppress isoflavone biosynthesis and promote infection in soybean. Cell Host and Microbe, 2011, 9(3): 177-186. doi: 10.1016/j.chom.2011.02.007
[32]	MACHO A P, GUIDOT A, BARBERIS P, BEUZON C R, GENIN S.A competitive index assay identifies several Ralstonia solanacearum type III effector mutant strains with reduced fitness in host plants. Molecular Plant-Microbe Interactions, 2010, 23(9): 1197-1205. doi: 10.1094/MPMI-23-9-1197
[33]	BARTETZKO V, SONNEWALD S, VOGEL F, HARTNER K, STADLER R, HAMMES U Z, BORNKE F.The Xanthomonas campestris pv. vesicatoria type III effector protein XopJ inhibits protein secretion: Evidence for interference with cell wall-associated defense eesponses. Molecular Plant-Microbe Interactions, 2009, 22(6): 655-664. doi: 10.1094/MPMI-22-6-0655
[34]	CHEN H, HU Y, QIN K Y, YANG X Z, JIA Z J, LI Q, CHEN H B, YANG H.A serological approach for the identification of the effector hopz5 of Pseudomonas syringae pv. actinidiae: A tool for the rapid immunodetection of kiwifruit bacterial canker. Journal of Plant Pathology, 2018, 100(2): 171-177. doi: 10.1007/s42161-018-0041-y
[35]	KRAUS C M, MUNKVOLD K R, MARTIN G B.Natural variation in tomato reveals differences in the recognition of AvrPto and AvrPtoB effectors from Pseudomonas syringae. Molecular Plant, 2016, 9(5): 639-649. doi: 10.1016/j.molp.2016.03.001
[36]	KIM M G, GENG X, LEE S Y, MACKEY D.The Pseudomonas syringae type III effector AvrRpm1 induces significant defenses by activating the Arabidopsis nucleotide-binding leucine-rich repeat protein RPS2. The Plant Journal, 2009, 57(4): 645-653. doi: 10.1111/tpj.2009.57.issue-4

Related Articles 15

[1]	DONG Yu, WU Qian, FENG Xuan, ZHENG YinYing, CUI BaiMing. A Novel Plasmid pEA60 of Erwinia amylovora Enhances the Pathogenicity of Strains by Regulating the Synthesis of Virulence Factors [J]. Scientia Agricultura Sinica, 2026, 59(5): 996-1007.
[2]	CONG QiQi, ZHANG JingYi, MENG XiangLong, DAI PengBo, LI Bo, HU TongLe, WANG ShuTong, CAO KeQiang, WANG YaNan. Identification of Hypovirus in Apple Ring Rot Fungus Botryosphaeria dothidea and Detection of Virus-Carrying Status in China [J]. Scientia Agricultura Sinica, 2025, 58(3): 478-492.
[3]	TONG ZhaoYang, LIU WenHua, ZHANG GuoXin, DONG ChunYan, ZHANG YanXia, XU XiaoWei, HE Dong, LIU HeChun, LI Yang, WANG FengTao, FENG Jing, YAO XiaoBo, LIU MeiJin, LIN RuiMing. The Relationship Between Occurrence of Hulless Barley Ear Rot and Population Migration of Grass Mite (Siteroptes spp.) [J]. Scientia Agricultura Sinica, 2025, 58(3): 493-506.
[4]	YANG WenJuan, GAO JiaCheng, WANG YanTing, LI Yan, GUO Ming, WANG JunCheng, MENG YaXiong, WANG HuaJun, SI ErJing. Function of Effector Pg00778 Regulation on the Pathogenicity of Pyrenophora graminea to Barley [J]. Scientia Agricultura Sinica, 2025, 58(15): 3020-3035.
[5]	DONG ZaiFang, DING TengTeng, SHAN YiXuan, LI HongLian, CHEN LinLin, XING XiaoPing. Autophagy-Related Gene FpAtg3 Involves in Growth and Pathogenicity of Fusarium pseudograminearum [J]. Scientia Agricultura Sinica, 2024, 57(6): 1080-1090.
[6]	ZHANG AiHong, YANG Fei, ZHAO YuanYe, ZHAO YiHan, DI DianPing, MIAO HongQin. Pathogenicity and Epidemic Risk of Barley Yellow Striate Mosaic Virus [J]. Scientia Agricultura Sinica, 2024, 57(23): 4686-4697.
[7]	WANG Yuan, DU MengDan, LI ZhengGang, SHE XiaoMan, YU Lin, LAN GuoBing, DING ShanWen, HE ZiFu, TANG YaFei. Identification of Pathogen Causing Tomato White Tip and Curl Leaf Disease and Its Pathogenicity in Guangdong Province [J]. Scientia Agricultura Sinica, 2024, 57(12): 2350-2363.
[8]	ZHANG Jian, ZHAO BinSen, FENG Hao, HUANG LiLi. Function and Mechanism Analysis of Vm-milRN7 Regulating the Pathogenicity of Valsa mali [J]. Scientia Agricultura Sinica, 2024, 57(10): 1930-1942.
[9]	GAO XiaoXiao, TU LiQin, YANG Liu, LIU YaNan, GAO DanNa, SUN Feng, LI Shuo, ZHANG SongBai, JI YingHua. Construction of an Infectious Clone of Tobacco Mild Green Mosaic Virus Isolate Infecting Pepper from Jiangsu Based on Genomic Clone [J]. Scientia Agricultura Sinica, 2023, 56(8): 1494-1502.
[10]	WANG JianFeng, CHENG JiaXin, SHU WeiXue, ZHANG YanRu, WANG XiaoJie, KANG ZhenSheng, TANG ChunLei. Functional Analysis of Effector Hasp83 in the Pathogenicity of Puccinia striiformis f. sp. tritici [J]. Scientia Agricultura Sinica, 2023, 56(5): 866-878.
[11]	GONG AnDong, LEI YinYu, WU NanNan, LIU JingRong, SONG MengGe, ZHANG YiMei, YANG Guang, YANG Peng. The Effect of 3-Oxyacyl ACP Reductase Gene FgOAR1 on the Growth, Development and Pathogenicity of Fusarium graminearum [J]. Scientia Agricultura Sinica, 2023, 56(24): 4854-4865.
[12]	LI HuiXin, SONG WenPing, HAN ZongXi, LIU ShengWang. Isolation and Pathogenicity of Fowl Adenovirus Serotype 8a Strain [J]. Scientia Agricultura Sinica, 2023, 56(16): 3226-3236.
[13]	HUANG JiaQuan,LI Li,WU FengNian,ZHENG Zheng,DENG XiaoLing. Proliferation of Two Types Prophage of ‘Candidatus Liberibacter asiaticus’ in Diaphorina citri and their Pathogenicity [J]. Scientia Agricultura Sinica, 2022, 55(4): 719-728.
[14]	YANG ShiMan, XU ChengZhi, XU BangFeng, WU YunPu, JIA YunHui, QIAO ChuanLing, CHEN HuaLan. Amino Acid of 225 in the HA Protein Affects the Pathogenicities of H1N1 Subtype Swine Influenza Viruses [J]. Scientia Agricultura Sinica, 2022, 55(4): 816-824.
[15]	LI ZhengGang,TANG YaFei,SHE XiaoMan,YU Lin,LAN GuoBing,HE ZiFu. Molecular Characteristics and Pathogenicity Analysis of Youcai Mosaic Virus Guangdong Isolate Infecting Radish [J]. Scientia Agricultura Sinica, 2022, 55(14): 2752-2761.

Metrics

Viewed

Full text

Abstract

Cited

Shared

Discussed

Comments

Recommended 0

No Suggested Reading articles found!

位置 Locus	T3SE	ICMP 18884		伴侣分子 Chaperon		基因敲除标识符 Symbol of gene deletion
	HopY1	IYO_000845
Cluster A	HopQ1	IYO_003525				A
	HopD1	IYO_003530
	AvrD1	IYO_003570
	AvrB4-l	IYO_003600
	HopAYl	IYO_003610
	HopX3	IYO_003635		ShcF
	HopAW1	IYO_003657
	HopBB1-2	IYO_003675		ShcF
	HopAF1	IYO_003680
	HopAO2	IYO_003720
	HopBB1-1	IYO_003727		ShcF
	HopS2	IYO_004052		ShcS2
	HopI1	IYO_005160
CEL/Cluster O	HopN1	IYO_006735		ShcN
	HopAA1-1	IYO_006745
	HopM1	IYO_006760		ShcM		O
	AvrE1	IYO_006770		ShcE		O
	AvrRpm1	IYO_008065				N
Cluster E	HopZ5	IYO_008282				E
Cluster E	HopH1	IYO_008285				E
	HopAM1-2	IYO_008385
	HopAE1	IYO_012225
	HopAZ1	IYO_018555
	AvrPto5	IYO_020425				L
	HonAM1-1	IYO_023205
Cluster F	HopAI1	IYO_023980				F
	HopAH1	IYO_023985
	HopAG1	IYO_023990
Cluster M	HopR1	IYO_024150				M
Cluster M	HopW1	IYO_024160
	HopF2	IYO_024217		ShcF
	HopAS1	IYO_027420
	HopA1	IYO_028375		ShcA
	HopZ3	IYO_029045		Tir
	AvrRpm2	IYO_029288		ShcF
Plasmid	HopAV1	IYO_029710
	HopAA1-2	IYO_029780-795
	HopAU1	IYO_029795
	存在的基因 Present		差异的基因 Variable		不完整的基因 Incomplete

The Function of Key T3SS Effectors in Pseudomonas syringae pv. actinidiae

RichHTML

PDF