植物学报 ›› 2021, Vol. 56 ›› Issue (6): 687-698.DOI: 10.11983/CBB21025
王田幸子, 朱峥, 陈悦, 刘玉晴, 燕高伟, 徐珊, 张彤, 马金姣, 窦世娟, 李莉云(
), 刘国振()
首页收稿日期:2021-01-31
接受日期:2021-08-09
出版日期:2021-11-01
发布日期:2021-11-12
通讯作者:
李莉云,刘国振
作者简介:gzhliu@hebau.edu.cn基金资助:
Tianxingzi Wang, Zheng Zhu, Yue Chen, Yuqing Liu, Gaowei Yan, Shan Xu, Tong Zhang, Jinjiao Ma, Shijuan Dou, Liyun Li(), Guozhen Liu()
Received:2021-01-31
Accepted:2021-08-09
Online:2021-11-01
Published:2021-11-12
Contact:
Liyun Li,Guozhen Liu
摘要： Xa21对白叶枯病菌(Xanthomonas oryzae pv. oryzae, Xoo)具有广谱抗性, 是最早被克隆的水稻(Oryza sativa)抗白叶枯病基因。前期研究表明, OsWRKY42可能在Xa21介导的抗病反应中发挥作用。在Xa21基因遗传背景下制备了OsWRKY42的RNA干扰株系, 将经免疫印迹确认的转基因株系接种白叶枯病菌, 结果表明, 与抗病对照4021相比, 转基因株系的病斑长度增加, 说明OsWRKY42的丰度下调抑制了Xa21对白叶枯病的抗性反应。免疫印迹分析表明, 在OsWRKY42-RNAi转基因水稻中, OsPR6、OsPR15和OsPR16的蛋白质丰度降低, OsPR1A、OsPR1B、OsPR2和OsPR10A的蛋白质丰度升高, 表明这些病程相关蛋白质可能位于OsWRKY42基因的下游, 受OsWRKY42调控并参与Xa21介导的抗病性。研究结果表明, OsWRKY42是Xa21介导的抗白叶枯病途径新元件, 增进了对Xa21介导的水稻抗病机理的认识。
王田幸子, 朱峥, 陈悦, 刘玉晴, 燕高伟, 徐珊, 张彤, 马金姣, 窦世娟, 李莉云, 刘国振. 水稻OsWRKY42是Xa21介导的抗白叶枯病途径新元件. 植物学报, 2021, 56(6): 687-698.
Tianxingzi Wang, Zheng Zhu, Yue Chen, Yuqing Liu, Gaowei Yan, Shan Xu, Tong Zhang, Jinjiao Ma, Shijuan Dou, Liyun Li, Guozhen Liu. Rice OsWRKY42 is a Novel Element in Xa21-mediated Resistance Pathway Against Bacterial Leaf Blight. Chinese Bulletin of Botany, 2021, 56(6): 687-698.
图1 OsWRKY42-RNAi转基因水稻的鉴定 将等量的4021和OsWRKY42-RNAi转基因水稻总蛋白质用Tricine-SDS-PAGE分离, 用Mini Chemiluminescent Imager and Sage Capture软件检测蛋白信号。PCR: PCR鉴定结果; WB: Western blot鉴定结果; CK: 转基因受体对照4021; R01-R03: 独立的OsWRKY42-RNAi转基因株系; 1-8: 同一转化株系中的不同转化植株; W42: 抗OsWRKY42多克隆抗体识别条带; HSP: 抗OsHSP82单克隆抗体识别条带。
Figure 1 Verification of rice OsWRKY42-RNAi transgenic lines Equal amounts of total protein from 4021 and OsWRKY42-RNAi transgenic lines were separated by Tricine-SDS-PAGE. The protein signal was detected by Mini Chemiluminescent Imager and Sage Capture software. PCR: PCR verification; WB: Western blot verification; CK: 4021 control of transgenic donor; R01-R03: Independent OsWRKY42-RNAi transgenic lines; 1-8: Individual plants within same transgenic lines; W42: Anti-OsWRKY42 polyclonal antibody detected band; HSP: Anti-OsHSP82 monoclonal antibody detected band.
图2 OsWRKY42-RNAi转基因水稻的叶绿素含量 (A)-(D) 依次为OsWRKY42-RNAi转基因水稻中叶绿素a (Chl a)、叶绿素b (Chl b)、总叶绿素(total Chl)及叶绿素a/叶绿素b的比值(Chl a/b)。TP309: 野生型; 4021: 表达Xa21的转基因TP309纯合植株; R01-R03: OsWRKY42-RNAi转基因水稻。误差线表示标准差(SD) (n=3); *表示差异显著(P<0.05); **表示差异极显著(P<0.01) (Student's t-test)。
Figure 2 Chlorophyll contents in rice OsWRKY42-RNAi transgenic lines (A)-(D) Chlorophyll a (Chl a) content, chlorophyll b (Chl b) content, total chlorophyll (total Chl) content and the ratio of Chl a/b in OsWRKY42-RNAi transgenic lines. TP309: Wild type; 4021: Homozygous transgenic TP309 line expressing Xa21; R01-R03: OsWRKY42-RNAi transgenic lines. Error bars are standard deviation (SD) (n=3); * indicate significant differences at P<0.05; ** indicate significant differences at P<0.01 (Student's t-test).
图3 OsWRKY42-RNAi转基因水稻接种Xoo的表型 (A) OsWRKY42-RNAi转基因水稻接种14天的叶片表型(bars=1 cm); (B) 病斑长度扩展曲线; (C) 接种14天病斑长度统计。误差线表示标准差SD (n>9); **差异极显著(P<0.01)。R01-R03: OsWRKY42-RNAi转基因水稻; TP309: 野生型; 4021: 表达Xa21的转基因TP309植株
Figure 3 Phenotype of rice OsWRKY42-RNAi transgenic lines inoculated with Xoo (A) The phenotype of OsWRKY42-RNAi transgenic lines at 14-day post inoculation (bars=1 cm); (B) Lesion development of inoculated plant; (C) Lesion length of inoculated plants at 14 day post inoculation. Error bars are standard deviation (SD) (n>9); ** significant differences at P<0.01 level. R01-R03: OsWRKY42-RNAi transgenic lines; TP309: Wild type; 4021: Transgenic TP309 expressing Xa21
图4 水稻接种叶片中Xoo总蛋白质丰度 将等量的多个样品的总蛋白质用SDS-PAGE分离蛋白后转膜, 用抗XOO抗体检测(附图1), 抗体为anti-XOO多克隆抗体和anti-OsHSP82单克隆抗体。用Mini Chemiluminescent Imager and Sage Capture软件检测免疫印迹(WB)信号, 用Lane 1D软件对免疫印迹检测结果进行信号采集。计算3次实验的平均值和标准差, 误差线表示标准差(SD)。TP309: 野生型; 4021: 表达Xa21的转基因TP309植株; R01和R02为OsWRKY42-RNAi转基因纯合株系。
Figure 4 Expression abundance of total Xoo protein in inoculated rice leaves Equal amounts of total protein from multiple samples were resolved by SDS-PAGE (appendix figure 1), antibodies are anti-XOO polyclonal antibody and anti-OsHSP82 monoclonal antibody. Western blot (WB) signals were detected by Mini Chemiluminescent Imager and Sage Capture software. Lane 1D software was used to extract signals of Western blot. Average and standard deviation was calculated for three repeats. Error bars are standard deviation (SD). * P<0.05; ** P<0.01; TP309: Wild type; 4021: Transgenic TP309 expressing Xa21; R01 and R02 were homozygous OsWRKY42- RNAi transgenic lines.
图5 OsWRKY42蛋白质在水稻-Xoo互作中的表达特征 (A) 接种后不同时间点OsWRKY42蛋白质的表达。分别在接种后0、1、2、4和6天5个时间点取材进行免疫印迹(WB)检测; (B) 接种叶片样品采集示意图。以病斑线为0点, 在接种后6天取材, 叶片取材部位分别为±1、1-3、3-5和5-7 cm; (C) 接种后叶片不同部位中OsWRKY42蛋白质的表达。以接种6天的水稻叶片为材料, 接种后的病斑线为0点, 采集接种后不同部位的叶片, 将等量的多个样品的总蛋白质用Tricine-SDS-PAGE分离, 一抗为抗OsWRKY42多克隆抗体和抗OsHSP82单克隆抗体, 用Sage Capture软件检测OsWRKY42蛋白质丰度。TP309: 野生型; 4021: 表达Xa21的转基因TP309植株; Xoo: Xoo总蛋白质; W42和HSP见图1。
Figure 5 Expression profiling of the OsWRKY42 protein in rice-Xoo interaction (A) Expression profiling of OsWRKY42 protein at different time points after inoculation. Samples were collected at 0, 1, 2, 4, 6-day post inoculation, respectively; (B) Schematic diagram for sample collection of inoculated leaves. Based on the lesion line (0), samples were taken at ±1, 1-3, 3-5, and 5-7 cm of the leaves at 6-day post inoculation, respectively; (C) Expression profiling of OsWRKY42 protein at different position of leaves at 6-day post inoculation. Based on lesion line, different portion of leaf tissues were collected after inoculation. Equal amounts of total protein from multiple samples were resolved by Tricine-SDS-PAGE. Antibodies are anti-OsWRKY42 polyclonal antibody and anti-OsHSP82 monoclonal antibody. OsWRKY42 protein signal were detected by Sage Capture software. TP309: Wild type; 4021: Transgenic TP309 expressing Xa21; Xoo: Total protein isolated from Xoo cells; W42 and HSP are shown in Figure 1.
图6 OsWRKY42-RNAi水稻植株中病程相关蛋白质的表达 以接种3天和6天的叶片(病斑线±1 cm)为材料, 等量的总蛋白质样品通过SDS-PAGE或者Tricine-SDS-PAGE分离, 一抗为抗病程相关蛋白质抗体, 分别为抗OsPR1A、抗OsPR1B及抗OsPR10A多克隆抗体(Wu et al., 2011); 抗OsPR2、抗OsPR6、抗OsPR15及抗OsPR16多克隆抗体(Hou et al., 2012)和抗OsHSP82单克隆抗体(Li et al., 2011)。TP309: 野生型; 4021: 表达Xa21的转基因TP309植株; R01-R03: OsWRKY42-RNAi转基因株系; HSP见图1。
Figure 6 Expression profiling of pathogenesis-related proteins in rice OsWRKY42-RNAi transgenic lines Equal amounts of total protein, isolated from inoculated leaves (lesion line ±1 cm) of 3-day and 6-day, were resolved by SDS-PAGE or Tricine-SDS-PAGE. The antibodies were anti-OsPR1A, -OsPR1B and -OsPR10A polyclonal antibodies (Wu et al., 2011); anti-OsPR2, -OsPR6, -OsPR15 and -OsPR16 polyclonal antibodies (Hou et al., 2012) and anti-OsHSP82 monoclonal antibody (Li et al., 2011). TP309: Wild type; 4021: Transgenic TP309 individual expressing Xa21; R01-R03: OsWRKY42-RNAi transgenic lines; HSP is shown in Figure 1.
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