欢迎访问作物学报，今天是
作物学报

作物学报 ›› 2022, Vol. 48 ›› Issue (1): 121-137.doi: 10.3724/SP.J.1006.2022.02090

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

水稻协优9308重组自交系群体白叶枯病抗性的全基因组关联分析

赵海涵1(), 练旺民1, 占小登1, 徐海明3, 张迎信1, 程式华1, 楼向阳1,*(), 曹立勇1,2,*(), 洪永波1,*()   

  1. 1中国水稻研究所水稻生物学国家重点实验室 / 浙江省超级稻重点实验室 / 国家水稻改良中心, 浙江杭州 311401
    2中国水稻研究所北方水稻研究中心, 黑龙江宝清 155600
    3浙江大学农业与生物技术学院作物科学研究所, 浙江杭州 310058
  • 收稿日期:2020-12-24 接受日期:2021-04-14 出版日期:2022-01-12 网络出版日期:2021-07-06
  • 通讯作者: 楼向阳,曹立勇,洪永波
  • 作者简介:赵海涵, E-mail: haihan3354@163.com第一联系人：**同等贡献
  • 基金资助:
    国家自然科学基金项目(31961143016);国家自然科学基金项目(31701338);国家重点研发计划项目(2018YFD0100806);农业农村部战略性科技创新合作项目资助(2020FYE0202300)

Genetic dissection of the bacterial blight disease resistance in super hybrid rice RILs using genome-wide association study

ZHAO Hai-Han1(), LIAN Wang-Min1, ZHAN Xiao-Deng1, XU Hai-Ming3, ZHANG Ying-Xin1, CHENG Shi-Hua1, LOU Xiang-Yang1,*(), CAO Li-Yong1,2,*(), HONG Yong-Bo1,*()   

  1. 1China National Center for Rice Improvement and State Key Laboratory of Rice Biology / Zhejiang Key Laboratory of Superrice / China National Rice Research Institute, Hangzhou 311401, Zhejiang, China
    2Northern Rice Center, China National Rice Research Institute, Baoqing 155600, Hebei, China
    3Institute of Crop Science and Institute of Bioinformatics, College of Agriculture and Biotechnology, Zhejiang University, Hangzhou 310058, Zhejiang, China
  • Received:2020-12-24 Accepted:2021-04-14 Published:2022-01-12 Published online:2021-07-06
  • Contact: LOU Xiang-Yang,CAO Li-Yong,HONG Yong-Bo
  • About author:First author contact:** Contributed equally to this work
  • Supported by:
    Natural Science Foundation of China(31961143016);Natural Science Foundation of China(31701338);National Key Research and Development Program(2018YFD0100806);Strategic Science and Technology Innovation Cooperation Project of the Ministry of Agriculture and Rural Affairs(2020FYE0202300)

RichHTML

27

PDF

1033

摘要：

白叶枯病是水稻生产最严重的细菌性病害, 挖掘新的白叶枯病抗性基因资源并培育抗病品种是控制该病害的重要方法。本研究利用父母本抗性差异较大的协优9308衍生的139个重组自交系群体作为遗传材料, 人工接种不同白叶枯菌后的病斑长度作为连续型表型, 结合经DNA深度测序获得的476,505个单核苷酸多态性(single-nucleotide polymorphism, SNP)标记进行全基因组关联分析(genome-wide associated study, GWAS)。结果表明在P < 1×10-4下, 4个菌株处理后共鉴定到109个与白叶枯抗性显著关联的SNPs位点, 解释表型变异率59.78%~63.29%。其中CR4接种发现了25个SNP位点其贡献率为61.00%, 在这些SNP位点附近共筛选到19个基因, 其中有2个为NBS-LRR抗病相关基因(LOC_Os11g43420和LOC_Os11g45930)。表达分析验证发现该2个基因在抗性亲本中恢9308的表达量分别为感病母本协青早B的4.42倍和8.86倍, 表明其可能在正调控白叶枯病抗性机制中发挥重要作用。进化树分析发现这2个候选基因与已克隆的抗性基因属于不同的亚组, 表明可能是新基因。本研究为进一步挖掘白叶枯抗性基因和培育抗病品种提供了理论基础和基因资源。

关键词: 水稻白叶枯病, 全基因组关联分析, NBS-LRR, 抗病育种

Abstract:

Bacterial blight caused by Xanthomonas oryzae pv. oryzae is the most destructive bacterial disease in rice production, and cultivating new disease-resistance variety by mining and utilizing the novel disease resistance genes is one of the most effective ways to control this disease. In our study, a population of 139 recombinant inbred lines (RILs) derived from super hybrid rice Xieyou 9308, between the parents of which there was genetic difference in resistance to bacterial blight, were inoculated with four Xoo strains. A genome-wide association study (GWAS) was carried out using 476,505 SNPs for the lesion length as a quantitative phenotype. The result revealed that a total of 109 significant SNPs including two genes encoding NBS-LRR containing proteins were detected to be significantly difference at P < 1×10-4 for four Xoo strains-treated lesions, accounting for phenotype variation of 59.78%-63.29%, respectively. Furthermore, 25 SNPs located in/nearby 19 candidate genes were identified by the CR4 inoculation, accounting for 61.00%. Relative expression analysis of two selected candidate genes, LOC_Os11g43420 and LOC_Os11g45930, demonstrated their higher expression levels in the resistant cultivar Zhonghui 9308 than in the susceptible variety Xieqingzao B, suggesting that these two genes might positively regulated bacterial blight resistance. Phylogenetic tree analysis indicated that these two genes were different from the previous cloned resistant genes in evolution, suggesting they were novel disease resistance genes. These results lays a theoretical basis and provides genetic resources for future resistant breeding in rice.

Key words: rice bacterial blight, GWAS, NBS-LRR, resistant breeding

表1

139个重组自交系及双亲接种4个生理菌株后的病斑长度统计特征数"

Xoo菌株
Xoo strain		 年份/区域
Year/location		 协青早B病斑长度(P1)
Lesion length of XB (P1) (cm)		 中恢9308病斑长度(P2)
Lesion length of R9308 (P2) (cm)		 亲本差异(P1-P2)的显著性P值
Significant P-value of parental difference (P1-P2)		 重组自交系
Recombinant inbred line		 变异系数
CV (%)
均值±标准差
Mean ± SD (cm)		 变化范围
Range (cm)
PXO96 2019/Hangzhou 21.6 5.4 <0.001** 13.7±7.0 0.2-37.7 51.9
CR4 2019/Hangzhou 21.6 5.3 <0.001** 7.7±4.2 0.2-20.3 54.5
PXO61 2020/Hangzhou 19.4 0.2 0.002** 1.2±2.4 0-19.4 203.0
PXO99 2020/Hangzhou 10.4 0.8 <0.001** 5.6±3.6 0.1-21.8 64.2

图1

协B/中恢9308接种4个生理菌株的白叶枯表型和病斑长度 A~D为双亲协青早B (左)和中恢9308 (右)分别接种4种生理菌株(PXO96、CR4、PXO61和PXO99), 15 d和21 d后调查病斑长度, 标尺为10 cm。图E~H为双亲接种不同生理菌株的平均病斑长度(cm)。**表示P ≤ 0.01 (Student’s t-test)。"

图2

139个重组自交系及双亲接种4个生理菌株的病斑长度的频率分布图 A~D分别来自于4种生理菌株: A: PXO96; B: CR4; C: PXO61; D: PXO99。图中横坐标表示平均病斑长度(cm), 纵坐标表示分布频数。黑色和红色箭头分别表示中恢93008和协青早B病斑长度对应横坐标的位置, 线条表示为数据的正态分布曲线。"

图3

139个重组自交系及双亲接种4个生理菌株的白叶枯抗性等级分布图 利用139 RIL和双亲在4种Xoo生理菌株中抗性等级数据进行分组: 根据每个水稻编号的平均病斑长度(LL)划分等级: 其中LL = 0 cm, 0 cm ≤ LL < 3 cm, LL/总叶片长度< 25%, LL/总叶片长度< 50%, LL/总叶片长度< 75%和整叶发病分别代表抗病(0级)、高抗(1级)、中抗(2级)、中感(3级)、高感(4级)和易感(5级); NA代表缺失值。"

图4

全基因组范围内的连锁不平衡衰减图 LD衰减图显示了476,506 SNPs在整个基因组中的连锁不平衡(r2)随距离(kb)的衰减。"

图5

139个重组自交系及双亲接种4个生理菌株的GWAS分析 A~D分别来自于4种生理菌株: A: PXO96; B: CR4; C: PXO61; D: PXO99。横坐标对应水稻全基因组, 纵坐标表示-log10 (P值), 蓝色线条表示Bonferroni矫正的P值, 橙色线条表示-log10 (P值)大于4.0的位点, 可判断与抗性关联。E~H分别绘制了对应的Q-Q图: E: PXO96; F: CR4; G: PXO61; H: PXO99。横坐标表示为期望的-log10 (P值), 纵坐标表示为观测的-log10 (P值)。红线作为参考线。"

表2

全基因组关联分析检测的显著SNP位点和抗性相关候选基因的注释信息"

菌株
Xoo		 SNP标记
SNP		 染色体
Chr.		 物理位置
Position		 P值
P-value		 基因名称(MSU)
Gene ID (MSU)		 基因注释
Gene annotation
CR4 11_26502942_C_A 11 26502942 1.71260×10-5 LOC_Os11g43890 WD domain, G-beta repeat domain containing protein, expressed
CR4 9_1843902_G_A 9 1843902 1.83954×10-5 LOC_Os09g03660 Retrotransposon protein, putative, unclassified, expressed
CR4 7_23436266_C_T 7 23436266 1.88459×10-5 LOC_Os07g39114 Expressed protein
CR4 7_23436267_C_G 7 23436267 1.88459×10-5
CR4 7_23436291_C_T 7 23436291 1.88459×10-5
CR4 5_1383235_C_A 5 1383235 1.92989×10-5 LOC_Os05g03360 Retrotransposon protein, putative, unclassified, expressed
CR4 7_23436858_C_T 7 23436858 2.13921×10-5 LOC_Os07g39114 Expressed protein
CR4 7_23436871_A_T 7 23436871 2.13921×10-5
CR4 8_26662359_G_A 8 26662359 2.44334×10-5 LOC_Os08g42210 Expressed protein
CR4 11_27792921_A_G 11 27792921 2.70257×10-5 LOC_Os11g45930 NBS-LRR type disease resistance protein, putative, expressed
CR4 11_26056269_A_G 11 26056269 2.90443×10-5 LOC_Os11g43200 Tropinone reductase 2, putative, expressed
CR4 2_11969901_C_T 2 11969901 3.20777×10-5 LOC_Os02g20330 Expressed protein
CR4 11_23030967_G_A 11 23030967 3.55194×10-5 LOC_Os11g38760 Retrotransposon protein, putative, LINE subclass, expressed
CR4 12_10252946_G_T 12 10252946 4.65692×10-5 LOC_Os12g17880 Armadillo/beta-catenin repeat protein-related, putative, expressed
CR4 11_26220673_C_T 11 26220673 5.00378×10-5 LOC_Os11g43420 LZ-NBS-LRR class RGA, putative, expressed
CR4 6_23077582_G_A 6 23077582 6.20380×10-5 LOC_Os06g38890 Transposon protein, putative, unclassified, expressed
CR4 11_26410429_C_A 11 26410429 6.81889×10-5 LOC_Os11g43740 OsMADS68 - MADS-box family gene with MIKC* type-box
CR4 11_25840166_C_A 11 25840166 7.97330×10-5 LOC_Os11g42900 Expressed protein
CR4 11_25849255_T_C 11 25849255 8.02272×10-5
CR4 6_23441121_G_C 6 23441121 8.04677×10-5 LOC_Os06g39470 Transferase family protein, putative, expressed
CR4 2_1831384_A_G 2 1831384 8.31200×10-5 LOC_Os02g04180 Transposon protein putative CACTA En/Spm sub-class expressed
CR4 4_3331408_G_A 4 3331408 8.31200×10-5 LOC_Os04g06374 Retrotransposon protein putative Ty3-gypsy subclass expressed
CR4 7_9599243_T_C 7 9599243 8.31200×10-5 LOC_Os07g16400 Retrotransposon protein putative Ty3-gypsy subclass expressed
CR4 8_20838490_G_A 8 20838490 9.02380×10-5 LOC_Os08g33410 Hypothetical protein
CR4 8_20838500_T_C 8 20838500 9.02380×10-5 LOC_Os02g21320 Retrotransposon protein putative unclassified expressed
PXO96 6_25611297_T_C 6 25611297 8.66084×10-7 LOC_Os10g05510
PXO96 5_14854774_G_A 5 14854774 3.83500×10-6 LOC_Os06g42010
PXO96 2_12653706_C_A 2 12653706 4.27352×10-6 LOC_Os03g35500 Expressed protein
PXO96 10_2723114_A_G 10 2723114 6.44760×10-6 LOC_Os10g20020 Expressed protein
PXO96 6_25220112_T_G 6 25220112 1.45559×10-5 LOC_Os06g37790 Transposon protein putative unclassified expressed
PXO96 3_19680871_T_C 3 19680871 2.49545×10-5 LOC_Os10g01510 Retrotransposon protein putative Ty3-gypsy subclass
PXO96 10_10043595_C_T 10 10043595 2.53663×10-5 LOC_Os03g28010 Retrotransposon protein putative Ty3-gypsy subclass expressed
PXO96 6_22366894_G_A 6 22366894 2.89696×10-5 LOC_Os06g42010 Expressed protein
PXO96 10_330955_C_T 10 330955 2.98270×10-5 LOC_Os06g42250 Transposon protein putative CACTA En/Spm sub-class expressed
PXO96 3_16101988_G_T 3 16101988 3.07716×10-5 LOC_Os06g42370 Retrotransposon protein putative unclassified expressed
PXO96 6_25219787_G_A 6 25219787 3.14674×10-5 LOC_Os06g33630 Transposon protein putative unclassified expressed
PXO96 6_25373580_A_G 6 25373580 3.85706×10-5 LOC_Os02g32680 Retrotransposon protein putative Ty3-gypsy subclass expressed
PXO96 6_25462404_C_T 6 25462404 4.00401×10-5 LOC_Os06g49185 Retrotransposon protein putative unclassified expressed
PXO96 6_19579269_C_T 6 19579269 4.66700×10-5 LOC_Os06g45410 Retrotransposon protein putative Ty3-gypsy subclass
PXO96 2_19389629_A_G 2 19389629 4.76090×10-5 LOC_Os06g42310 Lectin receptor-type protein kinase putative expressed
PXO96 6_29801791_T_C 6 29801791 4.82490×10-5 LOC_Os06g42010 Expressed protein
PXO96 6_27455797_A_T 6 27455797 5.46665×10-5 LOC_Os06g42370 MYB family transcription factor putative expressedMYB
PXO96 6_25409464_C_T 6 25409464 6.23488×10-5 LOC_Os07g33300 Beta-galactosidase precursor putative expressed
PXO96 6_25226687_G_C 6 25226687 6.81127×10-5 LOC_Os05g40940 Transposon protein putative unclassified expressed
PXO96 6_25465479_C_T 6 25465479 6.91484×10-5 LOC_Os06g41120 Retrotransposon protein putative unclassified expressed
PXO96 7_19906050_T_A 7 19906050 7.23635×10-5 LOC_Os02g32520 Expressed protein
PXO96 5_24009603_G_A 5 24009603 7.38552×10-5 LOC_Os06g42350 Retrotransposon protein putative unclassified expressed
PXO96 6_24594013_A_C 6 24594013 7.54406×10-5 LOC_Os06g42560 Expressed protein
PXO96 2_19249914_C_T 2 19249914 7.84868×10-5 LOC_Os06g42020 ERD1 protein chloroplast precursor putative expressed
PXO96 6_25451288_C_G 6 25451288 9.02863×10-5 LOC_Os06g42350 Transposon protein putative CACTA En/Spm sub-class expressed
PXO96 6_25592476_C_T 6 25592476 9.10497×10-5 LOC_Os06g42560 Tryptophan synthase beta chain 2 putative expressed
PXO96 6_25233994_G_A 6 25233994 9.17863×10-5 LOC_Os06g42020 CSLA9 - cellulose synthase-like family A expressed
PXO96 6_25446478_G_A 6 25446478 9.81605×10-5 LOC_Os06g42350 Transposon protein putative CACTA En/Spm sub-class expressed
PXO96 6_25226517_G_A 6 25226517 9.81690×10-5 LOC_Os06g42010 Transposon protein putative unclassified expressed
PXO96 6_25226850_C_T 6 25226850 9.81690×10-5
PXO96 6_28762618_T_C 6 28762618 9.93356×10-5 LOC_Os06g47500 Retrotransposon protein putative Ty3-gypsy subclass
PXO96 6_27492113_G_A 6 27492113 9.99334×10-5 LOC_Os06g45470 Expressed protein
PXO99 1_19343496_G_A 1 19343496 2.88795×10-7 LOC_Os01g34960 Retrotransposon putative centromere-specific
PXO99 1_29753419_G_A 1 29753419 2.88795×10-7 LOC_Os01g51740 Expressed protein
PXO99 1_33782923_T_C 1 33782923 2.88795×10-7 LOC_Os01g58444 Expressed protein
PXO99 2_5398259_G_A 2 5398259 2.88795×10-7 LOC_Os02g10280 Transposon protein putative unclassified expressed
PXO99 3_13482702_G_A 3 13482702 2.88795×10-7 LOC_Os03g23790 Retrotransposon protein putative Ty3-gypsy subclass expressed
PXO99 4_1315077_T_C 4 1315077 2.88795×10-7 LOC_Os04g03150 Retrotransposon protein putative unclassified expressed
PXO99 4_25911404_C_T 4 25911404 2.88795×10-7 LOC_Os04g43770 Expressed protein
PXO99 4_35500148_C_A 4 35500148 2.88795×10-7 LOC_Os04g59630 Prenylcysteine oxidase 1 precursor putative expressed
PXO99 7_26936907_G_A 7 26936907 2.88795×10-7 LOC_Os07g45120 Expressed protein
PXO99 8_10707525_C_A 8 10707525 2.88795×10-7 LOC_Os08g17480 Retrotransposon protein putative Ty3-gypsy subclass expressed
PXO99 9_13062864_A_G 9 13062864 2.88795×10-7 LOC_Os09g21580 OsFBX324 - F-box domain containing protein expressed
PXO99 10_770633_A_C 10 770633 2.88795×10-7 LOC_Os10g02220 Peptide transporter PTR2 putative expressed
PXO99 10_4768566_T_C 10 4768566 1.88802×10-6 LOC_Os10g08820 Retrotransposon protein putative Ty3-gypsy subclass expressed
PXO99 5_25082791_T_C 5 25082791 3.02499×10-6 LOC_Os05g43190 Expressed protein
PXO99 9_682155_C_T 9 682155 3.65396×10-6 LOC_Os09g01980 Retrotransposon protein putative unclassified expressed
PXO99 1_35729850_C_T 1 35729850 3.96584×10-6 LOC_Os01g61780 Vacuolar ATP synthase 98 kDa subunit putative expressed
PXO99 3_20089598_G_A 3 20089598 4.86058×10-6 LOC_Os03g36220 Retrotransposon protein putative unclassified expressed
PXO99 4_23009161_T_C 4 23009161 5.65336×10-6 LOC_Os04g38730 Hypothetical protein
PXO99 10_10280800_G_A 10 10280800 8.21681×10-6 LOC_Os10g20430 Transposon protein putative CACTA En/Spm sub-class expressed
PXO99 4_22488159_C_T 4 22488159 8.54439×10-6 LOC_Os04g37830 Retrotransposon protein putative unclassified expressed
PXO99 1_6763731_G_A 1 6763731 1.45237×10-5 LOC_Os01g12381 GDSL-like lipase/acylhydrolase putative expressed
PXO99 10_10622194_C_T 10 10622194 2.05682×10-5 LOC_Os10g20980 Transposon protein putative CACTA En/Spm sub-class expressed
PXO99 11_19490121_G_C 11 19490121 2.45252×10-5 LOC_Os11g32980 Transposon protein putative unclassified expressed
PXO99 6_15683763_C_T 6 15683763 2.50426×10-5 LOC_Os06g27700 Retrotransposon protein putative unclassified expressed
PXO99 2_11530742_G_A 2 115307424 2.50426×10-5 LOC_Os02g19710 Retrotransposon protein putative unclassified expressed
PXO99 8_14551707_C_T 8 14551707 2.58147×10-5 LOC_Os08g24080 Retrotransposon protein putative unclassified
PXO99 1_37300757_C_T 1 37300757 2.64067×10-5 LOC_Os01g64230 Long cell-linked locus protein putative expressed
PXO99 4_29550982_A_G 4 29550982 2.67698×10-5 LOC_Os04g49550 RING-H2 finger protein ATL2A putative expressed
PXO99 4_29550988_A_T 4 29550988 2.67698×10-5
PXO99 4_29550990_T_C 4 29550990 2.67698×10-5
PXO99 8_20326896_C_T 8 20326896 2.96673×10-5 LOC_Os08g32790 Retrotransposon protein putative Ty3-gypsy subclass expressed
PXO99 7_9714575_A_T 7 9714575 3.05773×10-5 LOC_Os07g16560 Retrotransposon protein putative Ty3-gypsy subclass expressed
PXO99 12_23369262_A_G 12 23369262 3.32972×10-5 LOC_Os12g38030 Retrotransposon protein putative Ty3-gypsy subclass expressed
PXO99 6_9880157_A_G 6 9880157 3.68628×10-5 LOC_Os06g17040 Retrotransposon protein putative unclassified expressed
PXO99 6_27238355_G_T 6 27238355 3.79721×10-5 LOC_Os06g45030 Transposon protein putative unclassified expressed
PXO99 10_2015605_G_A 10 2015605 3.98789×10-5 LOC_Os10g04290 Transposon protein putative CACTA En/Spm sub-class expressed
PXO99 10_2015606_T_C 10 2015606 3.98789×10-5
PXO99 10_2015616_T_C 10 2015616 3.98789×10-5
PXO99 7_9797928_A_T 7 9797928 4.00610×10-5 LOC_Os07g16710 Retrotransposon protein putative Ty3-gypsy subclass
PXO99 12_23610190_C_T 12 23610190 4.37810×10-5 LOC_Os12g38460 RNA recognition motif family protein expressed
PXO99 11_3704859_G_A 11 3704859 4.37973×10-5 LOC_Os11g07350 Expressed protein
PXO99 6_27238423_T_C 6 27238423 4.48254×10-5 LOC_Os06g45030 Transposon protein putative unclassified expressed
PXO99 6_27238426_A_G 6 27238426 4.48254×10-5 LOC_Os06g45030 Transposon protein putative unclassified expressed
PXO99 11_16720749_G_A 11 16720749 5.24152×10-5 LOC_Os11g28890 Retrotransposon protein putative Ty3-gypsy subclass expressed
PXO99 3_28507186_C_T 3 28507186 5.35983×10-5 LOC_Os03g49980 Retrotransposon protein putative unclassified expressed
PXO99 9_8088480_G_C 9 8088480 5.59333×10-5 LOC_Os09g13800 Retrotransposon protein putative Ty1-copia subclass expressed
PXO99 8_15034247_T_G 8 15034247 6.82446×10-5 LOC_Os08g24850 Retrotransposon putative centromere-specific expressed
PXO99 1_40956241_G_A 1 40956241 7.49332×10-5 LOC_Os01g70760 Retrotransposon protein putative Ty3-gypsy subclass expressed
PXO99 1_40956243_C_T 1 40956243 7.49332×10-5
PXO99 1_40956268_A_G 1 40956268 7.49332×10-5
PXO99 8_10707195_C_A 8 10707195 9.39077×10-5 LOC_Os08g17480 Retrotransposon protein putative Ty3-gypsy subclass expressed
PXO99 12_23610183_G_A 12 23610183 9.70588×10-5 LOC_Os12g38460 RNA recognition motif family protein expressed

图6

候选白叶枯抗性基因的相对表达量分析 A~D分别包括关联位点SNP_11_26220673_C_T (P = 5.00378× 10-5)、SNP_11_27792921_A_G (P = 2.70257×10-5)和SNP_6_ 25611297_T_C (P = 8.66×10-7)和Xa21基因的qRT-PCR分析。横坐标表示经过接菌时间: “0”表示未接生理菌株; “1”表示接种生理菌株当天; “4”表示接种生理菌株后4 d, 在相同的时间点对同一批次水稻植株的叶片进行取样。纵坐标表示以actin作为内参基因计算相对表达水平(folds)。E表示3个候选基因(LOC_Os11g43420、LOC_Os11g45930和LOC_Os06g42590) SNP突变分别在编码区和启动子区域。*表示P ≤ 0.05和**表示P ≤ 0.01 (Student’s t-test)。"

表3

已克隆的水稻白叶枯病抗性基因"

R基因
R gene		 位点
Loci (MSU format)		 染色体
Chr.		 物理位置
Position		 参考文献
Reference
Xa1 LOC_Os04g53120 4 31638099-31644795 Yoshimura et al. [9]
Xa3/Xa26 LOC_Os11g47210 11 28399360-28402773 Sun et al. [13]
Xa4 LOC_Os11g46870 11 28142192-28148337 Sun et al. [14]
xa5 LOC_Os05g01710 5 437013-443270 Blair et al. [11];
Xa10 LOC_Os11g37570 11 22181556-22181152 Tian et al. [16]
xa13 LOC_Os08g42350 8 26728795-26725898 Chu et al. [10]
Xa21 LOC_Os11g35500 11 20802924-20806518 Song et al. [17 ];
Xa23 LOC_Os11g37620 11 22204676-22203734 Wang et al. [15 ]
Xa25 LOC_Os12g29220 12 17305326-17302073 Triplettl et al [55].
Xa27 LOC_Os06g39810 6 23654303-23653851 Gu et al. [12]
xa41(t) LOC_Os11g31190 11 18171678-18174478 Hutin et al. [6]

图7

候选白叶枯抗性基因的进化分析 利用多重比对软件Clastal X将LOC_Os11g43420、LOC_Os11g45930、LOC_Os11g32980、LOC_Os12g17880、Xa1、Xa3/Xa26、Xa4、xa5、Xa10、xa13、Xa21、Xa23、Xa25、Xa27和xa41(t)比对后, 使用MEGA软件最大邻间法构建进化树。"

[1] Savary S, Willocquet L, Pethybridge S J, Esker P, McRoberts N, Nelson A. The global burden of pathogens and pests on major food crops. Nat Ecol Evol, 2019, 3:430-439.
[2] Mew T W. Focus on bacterial blight of rice. Plant Dis, 1993, 77:5-12.
[3] Niño-Liu D O, Ronald P C, Bogdanove A J. Xanthomonas oryzae pathovars: model pathogens of a model crop. Mol Plant Pathol, 2006, 7:303-324.
[4] Mew W T. Current status and future prospects of research on bacterial blight of rice. Annu Rev Phytopathol, 1987, 25:359-382.
[5] Kim S M. Identification of novel recessive gene Xa44(t) conferring resistance to bacterial blight races in rice by QTL linkage analysis using an SNP chip. Theor Appl Genet, 2018, 131:2733-2743.
[6] Hutin M, Sabot F, Ghesquiere A, Koebnik R, Szurek B. A knowledge-based molecular screen uncovers a broad-spectrum OsSWEET14 resistance allele to bacterial blight from wild rice. Plant J, 2015, 84:694-703.
[7] Chen X, Liu P, Mei L, He X, Chen L, Liu H, Shen S, Ji Z, Zheng X, Zhang Y, Gao Z, Zeng D, Qian Q, Ma B. Xa7, a new executor R gene that confers durable and broad-spectrum resistance to bacterial disease in rice. Plant Commun, 2021, 2:100143.
[8] Luo D, Huguet-Tapia J, Raborn R T, White F F, Brendel V P, Yang B. The Xa7 resistance gene guards the susceptibility gene SWEET14 of rice against exploitation by bacterial blight pathogen. Plant Commun, 2021, 2:100164.
[9] Yoshimura S, Yamanouchi U, Katayose Y, Toki S, Wang Z X, Kono I. Expression of Xa1, a bacterial blight resistance gene in rice, is induced by bacterial inoculation. Proc Natl Acad Sci USA, 1998, 95:1663-1668.
[10] Chu Z H, Fu B Y, Yang H, Xu C G, Li Z K, Sanchez A, Park Y J, Bennetzen J L, Zhang Q F, Wang S P. Targeting xa13, a recessive gene for bacterial blight resistance in rice. Theor Appl Genet, 2006, 112:455-461.
[11] Blair M W, Garris A J, Iyer A S, Chapman B, Kresovich S, McCouch S R. High resolution genetic mapping and candidate gene identification at the Xa5 locus for bacterial blight resistance in rice(Oryza sativa L.). Theor Appl Genet, 2003, 107:62-73.
[12] Gu K, Tian D, Yang F, Wu L, Skeekala C, Wang D. High-resolution genetic mapping of Xa27(t), a new bacterial blight resistance gene in rice, Oryza sativaL. Theor Appl Genet, 2004, 5:800-807.
[13] Sun X, Cao Y, Yang Z, Xu C, Li X, Wang S. Xa26, a gene conferring resistance to Xanthomonas oryzae pv. oryzae in rice, encodes an LRR receptor kinase-like protein. Plant J, 2004, 37:517-527
[14] Sun X, Yang Z, Wang S, Zhang Q. Identification of a 47-kb DNA fragment containing Xa4, a locus for bacterial blight resistance in rice. Theor Appl Genet, 2003, 4:683-687.
[15] 王春连. 水稻抗白叶枯病基因Xa23的图位克隆, 中国农业科学院博士学位论文,北京, 2006.
Wang C L. Mapping Cloning of Rice Bacterial Blight Resistance Gene Xa23 in China. PhD Dissertation of Chinese Academy of Agricultural Sciences, Beijing,China, 2006 (in Chinese with English abstract).
[16] Tian D, Wang J, Zeng X, Gu K, Qiu C, Yang X. The rice TAL effector-dependent resistance protein Xa10 triggers cell death and calcium depletion in the endoplasmic reticulum. Plant Cell, 2014, 26:497-515.
[17] Song W Y, Wang G L, Chen L L, Kim H S, Pi L Y, Holsten T, Gardner J, Wang B, Zhai W X, Zhu L H. A receptor kinase-like protein encoded by the rice disease resistance gene,Xa21. Science, 1995, 270:1804-1806.
[18] Wang C L, Zhang X P, Fan Y L, Gao Y, Zhu Q L, Zheng C K, Qin T F, Li Y Q, Che J Y. Xa23 is an executor R protein and confers broad-spectrum disease resistance in rice. Mol Plant, 2015. 8:290-302.
[19] Heath M C. Hypersensitive response-related death. Plant Mol Biol, 2000, 44:321-334.
[20] Shirasu K, Schulze-Lefert P. Regulators of cell death in disease resistance. Plant Mol Biol, 2000, 44:371-385.
[21] Boller T, Felix G. A renaissance of elicitors: perception of microbe-associated molecular patterns and danger signals by pattern-recognition receptors. Annu Rev Plant Biol, 2009, 60:379-406.
[22] Monteiro F, Nishimura M T. Structure fuction and genomic diversity of plant NLR proteins: an evolved resource for rational engineering of plant immunity. Annu Rev Phytopathol, 2018, 56:243-267.
[23] Andersen E J, Nepal M P, Purintun J M, Nelson D, Mermigka G, Sarris P F. Wheat disease resistance genes and their diversification through integrated domain fusions. Front Genet, 2020, 11.
[24] Monosi B, Wisser R J, Pennill L, Hulbert S H. Full-genome analysis of resistance gene homologues in rice. Theor Appl Genet, 2004, 109:1434-1447.
[25] Ronald P C, Albano B, Tabien R, Abenes L, Tanksley S D. Genetic and physical analysis of the rice bacterial blight disease resistance locus,Xa21. Mol Gen Genet, 1992, 236:113-120.
[26] Kiyosawa S, Yamaguchi H, Yamada M. The influence of resistance gene frequencies in rice plants on virulence gene frequencies in blast fungus population in Japan. Jpn J Phytopathol, 1982, 48:199-209.
[27] Khan M A, Naeem M, Iqbal M. Breeding approaches for bacterial leaf blight resistance in rice (Oryza sativa L.), current status and future directions. Eur J Plant Pathol, 2014, 139:27-37.
[28] Li Z K, Arif M, Zhong D B, Fu B Y, Xu J L, Domingo-Rey J. Complex genetic networks underlying the defensive system of rice (Oryza sativa L.) to Xanthomonas oryzae pv. oryzae. Proc Natl Acad Sci USA, 2006, 103:7994-7999
[29] Zhang F, Xie X, Xu M, Wang W, Xu J, Zhou Y. Detecting major QTL associated with resistance to bacterial blight using a set of rice reciprocal introgression lines with high density SNP markers. Plant Breed, 2015, 134:286-292.
[30] 杨长登, 曾大力, 马良勇. 水稻籼粳交DH群体白叶枯病抗性的QTL定位. 中国水稻科学, 2006, 20:102-104.
Yang C D, Zeng D L, Ma L Y. Mapping QTLs for bacterial blight resistance in a DH population from japonica/indica cross of rice(Oryza sativa). Chin J Rice Sci, 2006, 20:102-104 (in Chinese with English abstrat).
[31] 陈天晓, 朱亚军, 密雪飞. 利用水稻MAGIC群体关联定位白叶枯病抗性QTL和创制抗病新种质. 作物学报, 2016, 42:1437-1447.
Chen T X, Zhu Y J, Mi X F. Mapping of QTLs for bacterial blight resistance and screening of resistant materials using MAGIC populations of rice. Acta Agron Sin, 2016, 42:1437-1447 (in Chinese with English abstrat).
[32] Antoni R J. Association genetics in crop improvement. Curr Opin Plant Biol, 2010, 13:174-180.
[33] Huang X, Wei X, Sang T, Zhao Q, Feng Q, Zhao Y, Li C Y, Zhu C R, Lu T T, Zhang Z W, Li M, Fan D L, Guo Y L, Wang A H, Wang L, Deng L W. Genome-wide association studies of 14 agronomic traits in rice landraces. Nat Genet, 2010, 42:961.
[34] Tseng H Y, Lin D G, Hsieh H Y, Tseng Y J, Tseng W B, Chen C W, Wang C S. Genetic analysis and molecular mapping of QTLs associated with resistance to bacterial blight in a rice mutant,SA0423. Euphytica, 2015, 205:231-241.
[35] Niño-liu D O, Ronald P C, Bogdanove A J. Xanthomonas oryzae pathovars: model pathogens of a model crop. Mol Plant Pathol, 2010, 7:303-324.
[36] Ogawa T, Khush G S. Major genes for resistance to bacterial blight in rice. Bacterial Blight Rice. 1989.
[37] Basavaraj S H, Singh V K, Singh A, Singh A, Singh A, Anand D. Marker-assisted improvement of bacterial blight resistance in parental lines of Pusa RH10, a superfine grain aromatic rice hybrid. Mol Breed, 2010, 26:293-305.
[38] Zhou L Y, Liu S Y, Wu W X, Chen D B, Zhan X D, Zhu A K, Zhang Y X, Cheng S H, Cao L Y, Lou X Y, Xu H M. Dissection of genetic architecture of rice plant height and heading date by multiple-strategy-based association studies. Sci Rep, 2016, 6:29718.
[39] Mather K A, Caicedo A L, Polato N R, Olsen K M, McCouch S, Purugganan M D, The extent of linkage disequilibrium in rice (Oryza sativa L.). Genetics, 2007, 177:2223-2232.
[40] Chen X, Shang J, Chen D. A B-lectin receptor kinase gene conferring rice blast resistance. Plant J, 2006, 46:794-804.
[41] Cheng Q, Mao W, Xie W, Liu Q, Cao J, Yuan M, Zhang Q, Li X H, Wang S P. Characterization of a disease susceptibility locus for exploring an efficient way to improve rice resistance against bacterial blight. Sci China-Life Sci, 2017, 60:298-306.
[42] Li W T, Zhu Z, Chern M, Yin J, Yang C, Ran L, Cheng M, He M, Zhu L H, Li S G, Chen X W. A natural allele of a transcription factor in rice confers broad-spectrum blast resistance. Cell, 2017, 170:114-126.
[43] 章琦, 林汉明. 章琦稻病抗性研究选集. 北京: 中国农业出版社, 2010. p 555.
Zhang Q, Lam H M. Seleted Works of Zhang Qi on Rice Disease Resistance. Beijing: China Agriculture Press, 2010. p 555 (in Chinese).
[44] Kou Y J, Li X H, Xiao J H, Wang S P. Identification of genes contributing to quantitative disease resistance in rice. Sci China: Life Sci, 2010, 53:1263-1273.
[45] 王永军, 吴晓蕾, 贺超英, 张劲松, 陈受宜, 盖钧镒. 大豆作图群体检验与调整后构建的遗传图谱. 中国农业科学, 2003, 36:1254-1260.
Wang Y J, Wu X L, He C Y, Zhang J S, Chen S Y, Gai J Y. A soybean genetic linkage map constructed after the mapping population being tested and adjusted. Sci Agric Sin, 2003, 36:1254-1260 (in Chinese with English abstrat).
[46] Liu M H, Kang H X, Xu Y C, Peng Y, Wang D, Gao L J, Wang X L, Ning Y S, Wu J, Liu W D, Li C Y, Liu B, Wang G L. Genome-wide association study identifies an NLR gene that confers partial resistance to Magnaporthe oryzae in rice. Plant Biotech J, 2020, 18:1376-1383.
[47] Wang X Q, Pang Y L, Zhang J, Wu Z C, Chen K, Ali J, Ye G Y, Xu J L, Li Z K. Genome-wide and gene-based association mapping for rice eating and cooking characteristics and protein content. Sci Rep, 2017, 7:17203.
[48] Zhao K Y, Tung C W, Eizenga G C, Wright M H, Ali M L, Price A H, Norton G J, Islam M R, Reynolds A, Mezey J, McClung A M, Bustamante C D, McCouch S R. Genome-wide association mapping reveals a rich genetic architecture of complex traits in Oryza sativa. Nat Commun, 2011, 1:467.
[49] Zhang F, Wu Z C, Wang M M, Zhang F, Dingkuhn M, Xu J L, Zhou Y L, Li Z K. Genome-wide association analysis identifies resistance loci for bacterial blight in a diverse collection of indica rice germplasm. PLoS One, 2017, 12:e0174598.
[50] Song W Y, Pi L Y, Bureau T E, Ronald P C. Identification and characterization of 14 transposon-like elements in the noncoding regions of members of the Xa21 family of disease resistance genes in rice. Mol Gen Genet, 1998, 258:449-456.
[51] Sesma A, Osbourn A E. The rice leaf blast pathogen undergoes developmental processes typical of root-infecting fungi. Nature, 2004, 431:582-586.
[52] Poland J A, Bradbury P J, Buckler E S. Genome-wide nested association mapping of quantitative resistance to northern leaf blight in maize. Proc Natl Acad Sci USA, 2011, 108:6893-6898.
[53] Monosi B, Wisser R J, Pennill L. Full-genome analysis of resistance gene homologues in rice. Theor Appl Genet, 2004, 109:1434-1447.
[54] Basavaraj S H, Singh V K, Singh A. Marker-assisted improvement of bacterial blight resistance in parental lines of Pusa RH10, a superfine grain aromatic rice hybrid. Mol Breed, 2010, 26:293-305.
[55] Triplett L R, Cohen S P, Heffelfinger C. A resistance locus in the American heirloom rice variety carolina gold select is triggered by TAL effectors with diverse predicted targets and is effective against African strains of Xanthomonas oryzae pv. oryzicola. Plant J, 2016, 87:472-483.
[1] 习千辉, 徐梓瑗, 刘梦梦, 王宏艺, 郎凯琳, 井震海, 陈锋, 赵磊. 小麦籽粒铜含量的全基因组关联分析及候选基因预测[J]. 作物学报, 2026, 52(6): 1604-1617.
[2] 毛嘉琦, 黄朋雨, 赵佳佳, 郑兴卫, 武棒棒, 郝宇琼, 屈非, 刘成, 马朋涛, 郑军. 山西小麦品种白粉病抗性评价及抗病基因分子检测[J]. 作物学报, 2026, 52(6): 1669-1681.
[3] 唐宽强, 李公允, 宋美毅, 赵雪, 常春玲. 大豆株高性状全基因组关联分析及预测模型构建[J]. 作物学报, 2026, 52(6): 1743-1756.
[4] 杨欣雨, 崔文涛, 迪力尼格尔·阿力木, 汪凯翔, 吴鹏昊, 任姣姣. 玉米穗上叶片数全基因组关联分析和全基因组选择[J]. 作物学报, 2026, 52(5): 1573-1590.
[5] 闫安, 蒋昆炜, 王蓉圆, 田林, 张璐, 王韵, 徐建龙. 水稻剑叶小维管束数基因SVN7的鉴定与克隆[J]. 作物学报, 2026, 52(5): 1364-1372.
[6] 鲁雅妮, 丁超杰, 张煜, 杜习军, 齐学礼, 胡琳, 许为钢. 河南省200份小麦品种苗期茎基腐病抗性鉴定与全基因组关联分析[J]. 作物学报, 2026, 52(2): 363-375.
[7] 李诗晴, 王茜, 王素华, 张耀文, 王丽侠. 绿豆种质资源苗期耐盐性鉴定及相关基因发掘[J]. 作物学报, 2026, 52(2): 376-388.
[8] 苏爱国, 肖森林, 易红梅, 段赛茹, 王帅帅, 张如养, 邢锦丰, 李春辉, 孙轩, 徐瑞斌, 徐田军, 李志勇, 张勇, 王荣焕, 宋伟, 赵久然. 玉米穗腐病的抗性遗传研究进展与育种应用[J]. 作物学报, 2026, 52(1): 1-13.
[9] 李云香, 郭千纤, 侯万伟, 张小娟. 引进ICARDA小麦苗期根系抗旱性状的全基因组关联分析[J]. 作物学报, 2025, 51(9): 2387-2398.
[10] 李璐琪, 程宇坤, 白斌, 雷斌, 耿洪伟. 小麦叶片气孔相关性状全基因组关联分析[J]. 作物学报, 2025, 51(9): 2266-2284.
[11] 蔡金珊, 李超男, 王景一, 李宁, 柳玉平, 景蕊莲, 李龙, 孙黛珍. 小麦幼苗根系性状全基因组关联分析及TaSRL-3B优异等位基因发掘[J]. 作物学报, 2025, 51(8): 2020-2032.
[12] 李宜谦, 徐守振, 刘萍, 马麒, 谢斌, 陈红. 基于40K SNP芯片的陆地棉产量构成因素全基因组关联分析及单铃重位点挖掘[J]. 作物学报, 2025, 51(8): 2128-2138.
[13] 高梦娟, 赵贺莹, 陈家辉, 陈晓倩, 牛萌康, 钱琪润, 崔陆飞, 邢江敏, 银庆淼, 郭雯, 张宁, 孙丛苇, 阳霞, 裴丹, 贾奥琳, 陈锋, 余晓东, 任妍. 小麦抗纹枯病新位点Qse.hnau-5AS的定位及其候选基因鉴定[J]. 作物学报, 2025, 51(8): 2240-2250.
[14] 陈惠莹, 何嘉欣, 朱斌, 黄士轩, 周星佑, 伍君权, 杨美艳. 水稻黄单胞菌噬菌体vB_XaS_HDB2的全基因组分析和生物学特性研究[J]. 作物学报, 2025, 51(8): 2087-2099.
[15] 赵超男, 王金凤, 张玉, 张丽, 李瑞琦, 王鹏飞, 李鸽子, 张宏军, 虞波, 康国章. 全基因组关联分析定位与挖掘小麦氮高效基因[J]. 作物学报, 2025, 51(7): 1801-1813.
Viewed
Full text


Abstract

Cited
  Shared   
  Discussed   
No Suggested Reading articles found!
京ICP备15008789号-2