OsLOX10正调控水稻对稻瘟病和白叶枯病的抗性

doi:10.16819/j.1001-7216.2022.210604

中国水稻科学 ›› 2022, Vol. 36 ›› Issue (4): 348-356.DOI: 10.16819/j.1001-7216.2022.210604

OsLOX10正调控水稻对稻瘟病和白叶枯病的抗性

周永林¹^,², 申小磊¹^,², 周立帅¹^,², 林巧霞¹^,², 王朝露¹^,², 陈静¹^,², 冯慧捷¹, 张振文¹, 陈晓婷¹^,²^,³^,^*(), 鲁国东²

¹福建农林大学生命科学学院，福州 350002
²福建农林大学生物农药与化学生物学教育部重点实验室，福州 350002
³福建省作物有害生物监测与治理重点实验室，福州 350013

收稿日期:2021-06-10 修回日期:2021-09-30 出版日期:2022-07-10 发布日期:2022-07-12
通讯作者: 陈晓婷
基金资助:
国家自然科学基金资助项目(31972251);福建省作物有害生物监测与治理重点实验室开放课题(MIMCP-202001);福建农林大学科技创新专项(KFA17308A);福建农林大学科技创新专项(CXZX2020152D);福建农林大学生物学高原学科建设经费资助项目(71201800810)

OsLOX10 Positively Regulates Defense Responses of Rice to Rice Blast and Bacterial Blight

ZHOU Yonglin¹^,², SHEN Xiaolei¹^,², ZHOU Lishuai¹^,², LIN Qiaoxia¹^,², WANG Zhaolu¹^,², CHEN Jing¹^,², FENG Huijie¹, ZHANG Zhenwen¹, CHEN Xiaoting¹^,²^,³^,^*(), LU Guodong²

¹College of Life Sciences, Fujian Agriculture and Forestry University, Fuzhou 350002, China
²Key Laboratory of Biopesticide and Chemical Biology, Ministry of Education, Fujian Agriculture and Forestry University, Fuzhou 350002, China
³Fujian Key Laboratory for Monitoring and Integrated Management of Crop Pests, Fuzhou 350013, China

Received:2021-06-10 Revised:2021-09-30 Online:2022-07-10 Published:2022-07-12
Contact: CHEN Xiaoting

摘要/Abstract

摘要：

【目的】由稻瘟病菌（Magnaporthe oryzae）引起的稻瘟病和由水稻黄单胞菌（Xanthomonas oryzae pv. oryzae，Xoo）引起的白叶枯病严重影响水稻的产量和品质。创制转OsLOX10基因水稻材料，进行稻瘟菌和白叶枯菌的抗病性分析，有助于揭示其调控水稻对稻瘟病和白叶枯病的抗性机制。【方法】采用CRISPR/Cas9系统构建OsLOX10的敲除载体，利用限制性内切酶XcmⅠ线性化pCXUN-HA，TA连接构建OsLOX10的过表达载体，遗传转化获得OsLOX10转基因水稻，筛选过表达株系和纯合敲除株系进行真菌和细菌的抗病性分析。在稻瘟菌（Guy11）侵染水稻后，对水杨酸（salicylic acid，SA）、茉莉酸（jasmonic acid，JA）途径的标志基因进行qRT-PCR分析;在几丁质（chitin）和flg22诱导下，观测水稻活性氧（reactive oxygen species，ROS）的暴发情况。【结果】 qRT-PCR分析表明，接种稻瘟菌和白叶枯菌24 h后，OsLOX10表达量上调;OsLOX10的纯合敲除和过表达水稻转基因株系接种稻瘟病菌Guy11孢子悬浮液，与野生型（日本晴）相比，OsLOX10敲除株系更易感病，过表达株系则无典型的病斑症状;接种 6、12、24和36 h时，3个病程相关蛋白基因OsPBZ1、OsPR1a、OsPR1b和SA通路基因OsPAL1，以及JA合成通路上的2个基因OsAOS2、OsLOX5的转录水平在敲除转基因株系中显著下调，而在过表达转基因株系中显著上调。对转OsLOX10基因水稻接种白叶枯菌（PXO99A），发现敲除OsLOX10的转基因水稻对白叶枯菌更易感病。qRT-PCR分析OsPR1b和OsPAL1以及JA合成通路上的3个基因OsAOS2、OsAOC和OsJAZ在OsLOX10过表达基因水稻中表达量明显上调，而在敲除OsLOX10的转基因水稻中却保持在较低水平，在接种7 d后表现出显著性差异。在几丁质和flg22诱导下，OsLOX10敲除株系的ROS水平显著性降低，而且在几丁质诱导下，ROS的起峰时间推迟。【结论】稻瘟病菌和白叶枯病菌能够诱导OsLOX10的表达，OsLOX10通过病原菌分子模式触发的免疫途径（PTI）参与抗病反应，其在水稻抵御稻瘟病和白叶枯病中起着正调控作用。同时，OsLOX10可能通过调节SA和JA介导的信号通路来正调控水稻对稻瘟病和白叶枯病的抗性。

关键词: OsLOX10, CRISPR/Cas9基因编辑技术, 抗病性, 稻瘟病, 白叶枯病, 信号途径

Abstract:

【Objective】Rice blast caused by Magnaporthe oryzae and bacterial blight caused by Xanthomonas oryzae pv. oryzae (Xoo) seriously affect rice yield and quality. The transgenic materials of OsLOX10 were created to evaluate the resistance to M. oryzae and Xoo. The possible mechanism of OsLOX10 in regulating the defense responses to rice blast and bacterial blight were elucidated. 【Method】The knockout vector of OsLOX10 was constructed using CRISPR/Cas9 system, and the overexpression vector of OsLOX10 was constructed by linearizing pCXUN-HA with restriction enzyme XcmⅠ and linking with TA ligation. Transgenic rice was obtained by genetic transformation, and the homozygous transgenic lines were screened for fungal and bacterial resistance analysis. The expression dynamics of SA (salicylic acid) and JA (jasmonic acid) pathway marker genes were analyzed by qRT-PCR after infection with M. oryzae. The outbreak of reactive oxygen species (ROS) induced by chitin and flg22 in rice was also observed. 【Result】qRT-PCR analysis showed that the OsLOX10 expression was up-regulated 24 h after inoculation with M. oryzae and Xoo. The OsLOX10 knockout lines were more susceptible to the rice blast than that of the wild type (Nipponbare) when inoculated with the spore suspension of M. oryzae Guy11, while the overexpressed lines had no typical disease symptoms. At 6, 12, 24 and 36 hours post inoculation, the transcriptional levels of three pathogenesis-related genes, OsPBZ1, OsPR1a and OsPR1b, and two JA pathway genes, OsAOS2 and OsLOX5, and SA pathway gene OsPAL1, were significantly down-regulated in the knockout transgenic lines. However, they were significantly up-regulated in overexpressed transgenic lines. The OsLOX10 transgenic rice was inoculated with Xoo (PXO99A), and the OsLOX10 knockout lines were more susceptible to bacterial blight. qRT-PCR analysis showed that the relative expression levels of OsPR1b, OsPAL1 and the three genes (OsAOS2, OsAOC and OsJAZ) involved in the JA synthesis pathway were up-regulated in OsLOX10 overexpression lines, significantly down-regulated in OsLOX10 knockout lines seven days after Xoo inoculation. OsLOX10 knockout also significantly reduced the ROS level induced by chitin and flg22 in rice, delayed the peaking time of ROS induced by chitin, and reduced the sensitivity of rice to chitin and flg22.【Conclusion】OsLOX10 could be induced by infection with M. oryzae and Xoo. OsLOX10 may be involved in pathogen-associated molecular patterns triggered immunity, and play a positive regulatory role in rice resistance to rice blast and bacterial blight. Moreover, the OsLOX10 may regulate rice resistance to M. oryzae and Xoo through SA and JA mediated signaling pathways.

Key words: OsLOX10, CRISPR/Cas9, disease resistance, rice blast, bacterial blight, signal pathway

周永林, 申小磊, 周立帅, 林巧霞, 王朝露, 陈静, 冯慧捷, 张振文, 陈晓婷, 鲁国东. OsLOX10正调控水稻对稻瘟病和白叶枯病的抗性[J]. 中国水稻科学, 2022, 36(4): 348-356.

ZHOU Yonglin, SHEN Xiaolei, ZHOU Lishuai, LIN Qiaoxia, WANG Zhaolu, CHEN Jing, FENG Huijie, ZHANG Zhenwen, CHEN Xiaoting, LU Guodong. OsLOX10 Positively Regulates Defense Responses of Rice to Rice Blast and Bacterial Blight[J]. Chinese Journal OF Rice Science, 2022, 36(4): 348-356.

图/表 9

表1 本研究所使用引物

Table 1. Primers used in this study.

引物名称 Primer name	引物序列 Primer sequence（5＇-3＇）	用途 Purpose	片段长度Length/bp
OsLOX10-F	ATGCAGCAGCCGCAGGCGAG	过表达载体构建 Construction of OE vector	2607
OsLOX10-R	TCAGATGGAAGCGCTGTTGG	过表达载体构建 Construction of OE vector	2607
UbiP-seq	TTTTAGCCCTGCCTTCATACGC	过表达载体测序 Sequencing of OE vector	2795
NosR-seq	AGACCGGCAACAGGATTCAATC	过表达载体测序 Sequencing of OE vector	2795
OsLOX10-U3-F	ggcaGCTACACCCAACTCCCTACG	CRISPR/Cas9第1靶点验证 Verification of first-target site of CRISPR/Cas9	259
OsLOX10-U3-R	aaacCGTAGGGAGTTGGGTGTAGC		259
OsLOX10-U6a-F	gccgCGCCAGAGTCTGATCAACGC	CRISPR/Cas9第2靶点验证 Verification of second-target site of CRISPR/Cas9	329
OsLOX10-U6a-R	aaacGCGTTGATCAGACTCTGGCG		329
U-F	CTCCGTTTTACCTGTGGAATCG	敲除靶点第1轮扩增 First round amplification of knockout targets	534
gRNA-R	CGGAGGAAAATTCCATCCAC	敲除靶点第1轮扩增 First round amplification of knockout targets	534
Uctcg-B1＇	TTCAGAggtctcTctcgCACTGGAATCGGCAGCAAAGG	CRISPR/Cas9第1靶点扩增 Amplification of first-target site	564
gRctga-B2	AGCGTGggtctcGtcagGGTCCATCCACTCCAAGCTC	CRISPR/Cas9第1靶点扩增 Amplification of first-target site	564
Uctga-B2＇	TTCAGAggtctcTctgaCACTGGAATCGGCAGCAAAGG	CRISPR/Cas9第2靶点扩增 Amplification of second-target site target site	629
gRcggt-BL	AGCGTGggtctcGaccgGGTCCATCCACTCCAAGCTC		629
Actin-QF	GAGTATGATGAGTCGGGTCCAG	Actin定量PCR qRT-PCR of Actin	143
Actin-QR	ACACCAACAATCCCAAACAGAG	Actin定量PCR qRT-PCR of Actin	143
OsPR1a-QF	CGTCTTCATCACCTGCAACTACTC	OsPR1a定量PCR qRT-PCR of OsPR1a	132
OsPR1a-QR	CATGCATAAACACGTAGCATAGCA	OsPR1a定量PCR qRT-PCR of OsPR1a	132
OsPR1b-QF	ACGGGCGTACGTACTGGCTA	OsPR1b定量PCR qRT-PCR of OsPR1b	105
OsPR1b-QR	CTCGGTATGGACCGTGAAG	OsPR1b定量PCR qRT-PCR of OsPR1b	105
OsAOS2-QF	CAATACGTGTACTGGTCGAATGG	OsAOS2定量PCR qRT-PCR of OsAOS2	134
OsAOS2-QR	AAGGTGTCGTACCGGAGGAA	OsAOS2定量PCR qRT-PCR of OsAOS2	134
OsPAL1-QF	AGGAGCTCGGCTGCGTATT	OsPAL1定量PCR qRT-PCR of OsPAL1	79
OsPAL1-QR	ATGCCGAGGAACACCTTGTT	OsPAL1定量PCR qRT-PCR of OsPAL1	79
OsLOX5-QF	CTGATGAGGAGTTTGCACGA	OsLOX5定量PCR qRT-PCR of OsLOX5	542
OsLOX5-QR	TCGTCCTTCAGGAGCAGAAT	OsLOX5定量PCR qRT-PCR of OsLOX5	542
OsAOC-QF	CCAAGGTGCAGGAGATGTT	OsAOC定量PCR qRT-PCR of OsAOC	150
OsAOC-QR	TACAGCTTGTTGGTGAAGGG	OsAOC定量PCR qRT-PCR of OsAOC	150
OsJAZ-QF	GAAGGCTCAACAGCTGACCAT	OsJAZ定量PCR qRT-PCR of OsJAZ	69
OsJAZ-QR	TTGGTGGACGGGAAGTTCTC	OsJAZ定量PCR qRT-PCR of OsJAZ	69
OsUG-F	TTCTGGTCCTTCCACTTTCAG	泛素融合蛋白定量PCR qRT-PCR of ubiquitin fusion protein OsUG	92
OsUG-R	ACGATTGATTTAACCAGTCCATGA	泛素融合蛋白定量PCR qRT-PCR of ubiquitin fusion protein OsUG	92
MoPot2-F	ACGACCCGTCTTTACTTATTTGG	MoPot2定量PCR qRT-PCR of MoPot2	99
MoPot2-R	AAGTAGCGTTGGTTTTGTTGGAT	MoPot2定量PCR qRT-PCR of MoPot2	99

表1 本研究所使用引物

Table 1. Primers used in this study.

引物名称 Primer name	引物序列 Primer sequence（5＇-3＇）	用途 Purpose	片段长度Length/bp
OsLOX10-F	ATGCAGCAGCCGCAGGCGAG	过表达载体构建 Construction of OE vector	2607
OsLOX10-R	TCAGATGGAAGCGCTGTTGG	过表达载体构建 Construction of OE vector	2607
UbiP-seq	TTTTAGCCCTGCCTTCATACGC	过表达载体测序 Sequencing of OE vector	2795
NosR-seq	AGACCGGCAACAGGATTCAATC	过表达载体测序 Sequencing of OE vector	2795
OsLOX10-U3-F	ggcaGCTACACCCAACTCCCTACG	CRISPR/Cas9第1靶点验证 Verification of first-target site of CRISPR/Cas9	259
OsLOX10-U3-R	aaacCGTAGGGAGTTGGGTGTAGC		259
OsLOX10-U6a-F	gccgCGCCAGAGTCTGATCAACGC	CRISPR/Cas9第2靶点验证 Verification of second-target site of CRISPR/Cas9	329
OsLOX10-U6a-R	aaacGCGTTGATCAGACTCTGGCG		329
U-F	CTCCGTTTTACCTGTGGAATCG	敲除靶点第1轮扩增 First round amplification of knockout targets	534
gRNA-R	CGGAGGAAAATTCCATCCAC	敲除靶点第1轮扩增 First round amplification of knockout targets	534
Uctcg-B1＇	TTCAGAggtctcTctcgCACTGGAATCGGCAGCAAAGG	CRISPR/Cas9第1靶点扩增 Amplification of first-target site	564
gRctga-B2	AGCGTGggtctcGtcagGGTCCATCCACTCCAAGCTC	CRISPR/Cas9第1靶点扩增 Amplification of first-target site	564
Uctga-B2＇	TTCAGAggtctcTctgaCACTGGAATCGGCAGCAAAGG	CRISPR/Cas9第2靶点扩增 Amplification of second-target site target site	629
gRcggt-BL	AGCGTGggtctcGaccgGGTCCATCCACTCCAAGCTC		629
Actin-QF	GAGTATGATGAGTCGGGTCCAG	Actin定量PCR qRT-PCR of Actin	143
Actin-QR	ACACCAACAATCCCAAACAGAG	Actin定量PCR qRT-PCR of Actin	143
OsPR1a-QF	CGTCTTCATCACCTGCAACTACTC	OsPR1a定量PCR qRT-PCR of OsPR1a	132
OsPR1a-QR	CATGCATAAACACGTAGCATAGCA	OsPR1a定量PCR qRT-PCR of OsPR1a	132
OsPR1b-QF	ACGGGCGTACGTACTGGCTA	OsPR1b定量PCR qRT-PCR of OsPR1b	105
OsPR1b-QR	CTCGGTATGGACCGTGAAG	OsPR1b定量PCR qRT-PCR of OsPR1b	105
OsAOS2-QF	CAATACGTGTACTGGTCGAATGG	OsAOS2定量PCR qRT-PCR of OsAOS2	134
OsAOS2-QR	AAGGTGTCGTACCGGAGGAA	OsAOS2定量PCR qRT-PCR of OsAOS2	134
OsPAL1-QF	AGGAGCTCGGCTGCGTATT	OsPAL1定量PCR qRT-PCR of OsPAL1	79
OsPAL1-QR	ATGCCGAGGAACACCTTGTT	OsPAL1定量PCR qRT-PCR of OsPAL1	79
OsLOX5-QF	CTGATGAGGAGTTTGCACGA	OsLOX5定量PCR qRT-PCR of OsLOX5	542
OsLOX5-QR	TCGTCCTTCAGGAGCAGAAT	OsLOX5定量PCR qRT-PCR of OsLOX5	542
OsAOC-QF	CCAAGGTGCAGGAGATGTT	OsAOC定量PCR qRT-PCR of OsAOC	150
OsAOC-QR	TACAGCTTGTTGGTGAAGGG	OsAOC定量PCR qRT-PCR of OsAOC	150
OsJAZ-QF	GAAGGCTCAACAGCTGACCAT	OsJAZ定量PCR qRT-PCR of OsJAZ	69
OsJAZ-QR	TTGGTGGACGGGAAGTTCTC	OsJAZ定量PCR qRT-PCR of OsJAZ	69
OsUG-F	TTCTGGTCCTTCCACTTTCAG	泛素融合蛋白定量PCR qRT-PCR of ubiquitin fusion protein OsUG	92
OsUG-R	ACGATTGATTTAACCAGTCCATGA	泛素融合蛋白定量PCR qRT-PCR of ubiquitin fusion protein OsUG	92
MoPot2-F	ACGACCCGTCTTTACTTATTTGG	MoPot2定量PCR qRT-PCR of MoPot2	99
MoPot2-R	AAGTAGCGTTGGTTTTGTTGGAT	MoPot2定量PCR qRT-PCR of MoPot2	99

图1 OsLOX10在水稻接种稻瘟菌（A）和白叶枯菌（B）后表达量的动态变化图中数据为平均值±标准误，*、**分别代表处理间在P<0.05和P<0.01水平差异达显著水平(t-检验）。

Fig. 1. Expression profiles of OsLOX10 after inoculation with M. oryzae (A) and Xoo (B) in rice. Data in the figure are Mean±SE, and *, ** represent significant difference at the 0.05 and 0.01 levels (t-test), respectively.

图2 OsLOX3编辑纯合突变体的鉴定 A－OsLOX10结构和编辑位点;B－OsLOX10 T0代18号植株的突变位点检测;C－OsLOX10 T0代24号植株的突变位点检测。

Fig. 2. Identification of homozygous mutant of OsLOX10 by gene editing. A, Schematic presentation of the OsLOX10 structure and gene editing site; B, Mutation site in OsLOX10 gene editing line-18; C, Mutation sites in OsLOX10 gene editing line-24.

图3 qRT-PCR分析OsLOX10在其过表达植株中的表达量

Fig. 3. Expression analysis of OsLOX10 by qRT-PCR in overexpressed lines.

图4 OsLOX10转基因水稻对稻瘟病的抗性分析

Fig. 4. Response of OsLOX10 transgenic rice to M. oryzae.

图5 OsLOX10转基因稻接种白叶枯菌的症状

Fig. 5. Symptom of OsLOX10 transgenic lines inoculated with Xoo by leaf clipping method.

图6 OsLOX10转基因水稻相关抗病基因在稻瘟病菌侵染早期的qRT-PCR分析

Fig. 6. qRT-PCR analysis of disease resistance-related genes in OsLOX10 transgenic lines during the early infection of rice blast fungus.

图7 OsLOX10转基因水稻相关抗病基因在白叶枯病菌侵染早期的qRT-PCR分析

Fig. 7. qRT-PCR analysis of disease resistance-related genes in OsLOX10 transgenic lines during the early infection of bacterial blight.

图8 转基因水稻OsLOX10-KO对几丁质和flg22诱导的响应 A－几丁质诱导的转基因水稻OsLOX10-KO和日本晴中活性氧的暴发情况，水稻叶片用400 nmol/L的几丁质和水处理;B－flg22诱导的转基因水稻OsLOX10-KO和日本晴中活性氧的暴发情况，水稻叶片用500 nmol/L的flg22和水处理。

Fig. 8. Response of OsLOX10-KO transgenic rice and wild-type Nipponbare to flg22 and chitin. A, Chitin-induced ROS burst in the transgenic rice OsLOX10-KO and Nipponbare (rice leaf disks were exposed to 500 nmol/L chitin and water); B, flg22-induced ROS burst in the transgenic rice OsLOX10-KO and NPB (rice leaf disks were exposed to 400 nmol/L flg22 and water).

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OsLOX10正调控水稻对稻瘟病和白叶枯病的抗性

OsLOX10 Positively Regulates Defense Responses of Rice to Rice Blast and Bacterial Blight

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