水稻细菌性条斑病菌致病与水稻抗病机制研究进展

doi:10.11983/CBB25098

植物学报 ›› 2025, Vol. 60 ›› Issue (5): 759-772.DOI: 10.11983/CBB25098 cstr: 32102.14.CBB25098

水稻细菌性条斑病菌致病与水稻抗病机制研究进展

吴艾安, 陶一菲, 方思棋, 许欣悦, 朱珊珊, 陈诗颖, 王廷超, 郭威^,^*()

浙江师范大学生命科学学院, 金华 321004

收稿日期:2025-05-30 接受日期:2025-09-02 出版日期:2025-09-10 发布日期:2025-09-02
通讯作者: *郭威, 浙江师范大学生命科学学院副教授, 硕士生导师。主要从事植物-病原微生物互作研究, 聚焦大豆和水稻等农作物与黄单胞菌的分子互作机制。以第一/通讯作者发表学术论文20余篇。授权国家发明专利5项。主持完成国家自然科学基金和浙江省自然科学基金等科研项目9项。E-mail: weiguo817@zjnu.cn
基金资助:
国家重点研发计划(2024YFD1400702);国家级大学生创新创业训练计划(202510345016);国家级大学生创新创业训练计划(202510345082X);浙江省大学生科技创新活动计划(2025R404A039);浙江省大学生科技创新活动计划(2025R404A041)

Research Progress on Pathogenesis of Xanthomonas oryzae pv. oryzicola and Rice Resistance Mechanisms

Wu Aian, Tao Yifei, Fang Siqi, Xu Xinyue, Zhu Shanshan, Chen Shiying, Wang Tingchao, Guo Wei^,^*()

College of Life Sciences, Zhejiang Normal University, Jinhua 321004, China

Received:2025-05-30 Accepted:2025-09-02 Online:2025-09-10 Published:2025-09-02
Contact: *E-mail: weiguo817@zjnu.cn

摘要/Abstract

摘要： 水稻(Oryza sativa)细菌性条斑病(BLS)是由稻黄单胞菌稻生致病变种(Xoc)引起的一种重要检疫性病害。该病原菌兼具高度遗传多样性和强传播能力, 在种植集约化及气候变暖的双重驱动下, 在我国南方籼稻主产区持续扩散。该文从以下3个方面系统综述了Xoc-水稻互作机制研究进展。(1) 从病原层面解析了II型分泌系统(T2SS)、III型分泌系统(T3SS)及胞外多糖等关键毒性因子的致病机制, 揭示了致病小种的分化规律; (2) 从寄主层面阐明了PTI/ETI介导的抗病信号通路, 综述了抗病(R)基因克隆与感病(S)基因编辑研究进展; (3) 展望了未来的研究方向, 将致力于深度整合多组学技术系统解析Xoc致病信号网络, 依托泛基因组学规模化挖掘具持久广谱抗性的R基因, 创新构建S基因靶向编辑与植物免疫激活协同增效的绿色防控体系, 为BLS的可持续治理提供系统性解决方案。

关键词: 细菌性条斑病, 稻黄单胞菌稻生致病变种, 致病机制, 先天免疫, 抗/感病基因

Abstract: Rice bacterial leaf streak (BLS), caused by Xanthomonas oryzae pv. oryzicola (Xoc), is a significant quarantine disease. The pathogen exhibits both high genetic diversity and strong transmission capabilities. Driven by agricultural intensification and global warming, BLS has been progressively expanding across major indica rice-producing regions in southern China. This review systematically summarizes recent advances in Xoc-rice interaction mechanisms: (1) Pathogen perspective: elucidating pathogenic mechanisms of virulence factors (including T2SS, T3SS, and extracellular polysaccharides (EPS)) and pathovar differentiation patterns; (2) Host perspective: clarifying advances in PTI/ETI-mediated immunity signaling pathways, resistance (R) gene cloning, and susceptibility (S) gene editing; and (3) Future directions: proposing multi-omics approaches to decode Xoc pathogenicity networks, leveraging pan-genomics for large-scale mining of durable and broad-spectrum R genes, and constructing synergistic systems integrating S gene editing with immune activation to establish systematic solutions for sustainable BLS management.

Key words: bacterial leaf streak, Xanthomonas oryzae pv. oryzicola, pathogenic mechanism, innate immunity, resistance/susceptibility genes

吴艾安, 陶一菲, 方思棋, 许欣悦, 朱珊珊, 陈诗颖, 王廷超, 郭威. 水稻细菌性条斑病菌致病与水稻抗病机制研究进展. 植物学报, 2025, 60(5): 759-772.

Wu Aian, Tao Yifei, Fang Siqi, Xu Xinyue, Zhu Shanshan, Chen Shiying, Wang Tingchao, Guo Wei. Research Progress on Pathogenesis of Xanthomonas oryzae pv. oryzicola and Rice Resistance Mechanisms. Chinese Bulletin of Botany, 2025, 60(5): 759-772.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB25098

https://www.chinbullbotany.com/CN/Y2025/V60/I5/759

图/表 6

表1 中国水稻生态种植区与细菌性条斑病(BLS)流行区的空间耦合关系

Table 1 Spatial coupling relationship between rice planting ecoregions and bacterial leaf streak (BLS) epidemic areas in China

中国稻作区	BLS主要发病区	BLS严重发病区
华南双季稻稻作区	海南; 广东和广西北部; 云南和福建南部	广东和广西南部; 福建北部
华中双单季稻稻作区	四川东北部; 安徽和江苏北部; 湖南和江西南部	湖北; 浙江; 上海; 湖南东北部; 江西北部; 安徽和江苏南部
华北单季稻稻作区	几乎无	几乎无
东北早熟单季稻稻作区	几乎无	几乎无
西南高原单双季稻稻作区	贵州; 云南西部和南部	湖南西北部; 四川西南部
西北干燥区单季稻稻作区	几乎无	几乎无

图1 稻黄单胞菌稻生致病变种(Xoc)侵染循环与水稻细菌性条斑病(BLS)传播途径示意图

Figure 1 Schematic diagram of Xanthomonas oryzae pv. oryzicola (Xoc) infection cycle and rice bacterial leaf streak (BLS) transmission pathways

图2 Xoc与寄主水稻互作示意图 Xoc利用T2SS分泌CWDEs水解水稻细胞壁组分, 促进病原侵染与定殖; 利用T3SS向水稻细胞中注入T3SEs (TALEs和non-TALEs), TALEs通过核定位靶向寄主基因启动子EBE区, 特异性激活R/E/S基因表达, 触发ETI或ETS, non-TALEs协同抑制调控激素合成的PTI。Xoc分泌EPS包裹菌体形成黏质层, 逃避ROS迸发、胼胝质沉积等寄主免疫识别。Xoc: 稻黄单胞菌稻生致病变种; T2SS: II型分泌系统; CWDEs: 细胞壁降解酶; T3SS: III型分泌系统; T3SEs: III型分泌效应蛋白; TALEs: 转录激活类效应子; non-TALEs: 非转录激活类效应子; EBE: 效应蛋白结合元件; R gene: 抗性基因; E gene: 执行者基因; S gene: 感性基因; ETI: 效应蛋白触发的免疫; ETS: 效应蛋白触发的感病; PTI: 病原体相关分子模式触发的免疫; EPS: 胞外多糖; ROS: 活性氧; Xops: 黄单胞菌外泌蛋白; MAPK: 丝裂原活化蛋白激酶; SA: 水杨酸; JA: 茉莉酸; ET: 乙烯; RVDs: 可变重复区; NLS: 核定位信号; AD: 酸性转录激活域。箭头表示激活作用, 钝线表示抑制作用。

Figure 2 Schematic diagram of the interaction between Xoc and host rice Xoc employs the T2SS to secrete CWDEs that hydrolyze rice cell wall components, facilitating its invasion and colonization; concurrently, it utilizes the T3SS to inject T3SEs (TALEs and non-TALEs) into rice cells. TALEs target the EBE region of host gene promoters through nuclear localization, specifically activating R/E/S gene expression to trigger ETI or ETS, while non-TALEs cooperatively suppress PTI that regulates hormone synthesis. Xoc secretes EPS to form a mucoid layer encapsulating bacterial cells, thereby evading host immune recognition including ROS burst and callose deposition. Xoc: Xanthomonas oryzae pv. oryzicola; T2SS: Type II secretion system; CWDEs: Cell wall-degrading enzymes; T3SS: Type III secretion system; T3SEs: Type III secretion effectors; TALEs: Transcription activator-like effectors; non-TALEs: Non transcription activator-like effectors; EBE: Effector-binding element; R gene: Resistance gene; E gene: Executor gene; S gene: Susceptibility gene; ETI: Effector-triggered immunity; ETS: Effector-triggered susceptibility; PTI: Pattern-triggered immunity; EPS: Extracellular polysaccharides; ROS: Reactive oxygen species; Xops: Xanthomonas outer proteins; MAPK: Mitogen-activated protein kinase; SA: Salicylic acid; JA: Jasmonic acid; ET: Ethylene; RVDs: Repeat-variable diresidues; NLS: Nuclear localization signal; AD: Acidic activation domain. Lines with arrows mean activation while blunt lines mean repression.

图3 水稻免疫应答网络示意图 Xoc与水稻互作中, PRRs识别PAMPs激活RLCKs, 诱导Ca2+内流、MAPK激活及转录重编程等防御信号。RLCKs磷酸化RBOHs的N端, 被激活的RBOHs将O2转化为H2O2, 引起ROS迸发, 促进防御基因表达。MAPK级联信号调控SA、JA和ET等激素合成, 形成交叉调控网络以调节水稻抗性。NLRs识别T3SEs, 整合细胞内信号事件触发HR等防御反应。PTI与ETI协同互作, 共同增强免疫反应。Xoc: 稻黄单胞菌稻生致病变种; PRRs: 模式识别受体; PAMPs: 病原体相关分子模式; RLCKs: 受体样细胞质激酶; MAPK: 丝裂原活化蛋白激酶; RBOHs: 呼吸爆发氧化酶同源蛋白; ROS: 活性氧; SA: 水杨酸; JA: 茉莉酸; ET: 乙烯; NLRs: 核苷酸结合域-富含亮氨酸重复序列; T3SEs: III型分泌效应蛋白; HR: 过敏反应; PTI: 病原相关分子模式触发的免疫; ETI: 效应子触发的免疫; IAA: 生长素; GA: 赤霉素; ABA: 脱落酸。箭头表示直接(实线)或间接(虚线)激活作用, 钝线表示抑制作用。

Figure 3 Schematic diagram of rice innate immune system During Xoc-rice interactions, PRRs recognize PAMPs to activate RLCKs, inducing defense signals including Ca2+ influx, MAPK activation, and transcriptional reprogramming. RLCKs phosphorylate the N-terminus of RBOHs. The activated RBOHs convert O2 to H2O2, causing ROS burst and promoting defense gene expression. The MAPK cascade regulates the synthesis of plant hormones SA, JA, and ET, forming a hormone cross-regulatory network that modulates rice resistance. NLRs recognize T3SEs and integrate intracellular signaling events to trigger defense responses such as HR. PTI and ETI act synergistically to collectively enhance immune responses. Xoc: Xanthomonas oryzae pv. oryzicola; PRRs: Pattern recognition receptors; PAMPs: Pathogen-associated molecular patterns; RLCKs: Receptor-like cytoplasmic kinases; MAPK: Mitogen-activated protein kinase; RBOHs: Respiratory burst oxidase homologs; ROS: Reactive oxygen species; SA: Salicylic acid; JA: Jasmonic acid; ET: Ethylene; NLRs: Nucleotide-binding domain leucine-rich repeat containing receptors; T3SEs: Type III secretion effectors; HR: Hypersensitive response; PTI: Pattern-triggered immunity; ETI: Effector-triggered immunity; IAA: Auxin; GA: Gibberellin; ABA: Abscisic acid. Arrows indicate direct (solid) or indirect (dashed) activation, flathead arrows indicate inhibition.

表2 水稻细菌性条斑病(BLS)抗性(R)位点与基因

Table 2 Resistance (R) loci and genes for bacterial leaf streak (BLS) in rice

来源	位点与基因	参考文献
玉米	Rxo1	Zhao et al., 2005
Minghui 63	OsWRKY45-2	Tao et al., 2009
ZH11	OsMPK6	Shen et al., 2010
Acc8558	qBlsr5a: 5号染色体ID73-ID79 (30 kb)	Xie et al., 2014
Carolina Gold Select	Xo1: 1090 kb	Triplett et al., 2016
ZH11	AtNPR1	Xu et al., 2017
TP309	XCRK: 1号染色体上	张玉霞等, 2018
Acc8558	qBlsr3d: LOC_Os3g03570 (81 kb)	Wang et al., 2020a
DP3	bls1: 6号染色体上(21 kb)	Ma et al., 2021
DY19	bls2: 2号染色体RM13592-RM13599 (240 kb)	罗登杰等, 2021
X455	Xo2: 2号染色体RM12941-M6-1 (110 kb)	Chen et al., 2022b
WP1, 9311	qBLS4.1: 4号染色体上(521 kb)	韦敏益等, 2023

表3 水稻细菌性条斑病(BLS)感病(S)基因

Table 3 Susceptibility (S) genes for bacterial leaf streak (BLS) in rice

来源	基因	参考文献
Mudanjiang 8, Dongjin	OsWRKY45-1	Tao et al., 2009
Nipponbare	OsSULTR3;6	Cernadas et al., 2014
Nipponbare	Os09g29100	Cai et al., 2017
ZH11	OsTFIIAγ5, OsTFIIAγ5^V39E	Hui et al., 2019
Guihong 1, ZH11	OsSULTR3;6	Ni et al., 2021
IRBB10	OsSULRT3;6	Xu et al., 2021
ZH11	OsF3H_03g	Wu et al., 2022
ZH11	OsBLS6.2	Xie et al., 2023
Nipponbare, ZH11	OsHXK5, OsSULRT3;6	Wang et al., 2024

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水稻细菌性条斑病菌致病与水稻抗病机制研究进展

Research Progress on Pathogenesis of Xanthomonas oryzae pv. oryzicola and Rice Resistance Mechanisms

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