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作物学报 ›› 2026, Vol. 52 ›› Issue (10): 2898-2911.doi: 10.3724/SP.J.1006.2026.63030

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

玉米抗瘤黑粉病种质评价及相关基因挖掘

杨文妍1,2(), 渠建洲1, 刘耕瑜1, 卢誉1, 刘健茁1, 张红伟2, 杜万里1,*()   

  1. 1 沈阳农业大学农学院, 辽宁沈阳 110161
    2 中国农业科学院作物科学研究所, 北京 100081
  • 收稿日期:2026-03-08 接受日期:2026-07-15 出版日期:2026-10-12 网络出版日期:2026-07-22
  • 通讯作者: 杜万里, E-mail: dwl2014@syau.edu.cn
  • 作者简介:杨文妍, E-mail: ywy02272021@163.com
  • 基金资助:
    辽宁省教育厅基本科研项目(JYTXS2024004);国家自然科学基金项目(32272152)

Evaluation of maize germplasm for resistance to common smut and identification of related genes

Yang Wen-Yan1,2(), Qu Jian-Zhou1, Liu Geng-Yu1, Lu Yu1, Liu Jian-Zhuo1, Zhang Hong-Wei2, Du Wan-Li1,*()   

  1. 1 College of Agronomy, Shenyang Agricultural University, Shenyang 110161, Liaoning, China
    2 Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081, China
  • Received:2026-03-08 Accepted:2026-07-15 Published:2026-10-12 Published online:2026-07-22
  • Contact: Du Wan-Li, E-mail: dwl2014@syau.edu.cn
  • Supported by:
    Basic Research Project of Liaoning Provincial Department of Education(JYTXS2024004);National Natural Science Foundation of China(32272152)

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摘要：

玉米瘤黑粉病是由玉蜀黍黑粉菌(Ustilago maydis)引起的世界性病害, 严重威胁玉米产量与品质, 培育抗病品种是防控该病害最经济有效的策略。为鉴定抗瘤黑粉病优异种质, 挖掘抗病关键基因, 本研究系统评价了425份玉米自交系的抗性水平, 旨在解析抗性遗传基础, 筛选抗病基因。分别于2年2点开展田间人工接种鉴定, 基于病情指数(DI)、发病率(IR)和病程曲线下面积(AUDPC)进行抗性评价。利用970,761个高质量SNP进行全基因组关联分析(GWAS), 结合时空转录组数据, 筛选抗病相关候选基因并进行功能富集分析。425份自交系中鉴定出高抗6份、抗病27份、中抗67份、感病154份、高感171份。DI、IR和AUDPC的广义遗传力均高于75%。GWAS共检测到1665个显著关联SNP位点, 筛选出376个候选基因, 其中, 79个响应病原菌侵染差异表达, 19个显著上调。GO富集分析表明, 筛选到的候选基因主要参与防御反应、细胞壁组织、激素信号传导等过程。本研究明确了玉米瘤黑粉病抗性具有遗传力高、受多基因控制的特点, 鉴定的抗病关联位点及候选基因, 为玉米抗瘤黑粉病分子育种提供重要种质资源和基因靶点。

关键词: 玉米, 瘤黑粉病, 全基因组关联分析, 转录组, 抗病候选基因

Abstract:

Maize common smut, caused by Ustilago maydis, is a widespread disease that seriously threatens maize yield and quality worldwide. Breeding resistant varieties is the most economical and effective strategy for controlling this disease. To identify elite germplasm resistant to common smut and uncover key resistance genes, this study systematically evaluated the resistance levels of 425 maize inbred lines, aiming to dissect the genetic basis of resistance and screen for resistance-related genes. Artificial inoculation and resistance evaluation were conducted in the field at two locations over two years, based on disease index (DI), incidence rate (IR), and area under the disease progress curve (AUDPC). A genome-wide association study (GWAS) was performed using 970,761 high-quality SNPs, and candidate resistance genes were further screened by integrating spatiotemporal transcriptomic data, followed by functional enrichment analysis. Among the 425 inbred lines, 6 were identified as highly resistant, 27 as resistant, 67 as moderately resistant, 154 as susceptible, and 171 as highly susceptible. The broad-sense heritabilities of DI, IR, and AUDPC were all above 75%. GWAS detected a total of 1665 significantly associated SNP loci, leading to the identification of 376 candidate genes, among which 79 were differentially expressed in response to pathogen infection and 19 were significantly upregulated. GO enrichment analysis indicated that the candidate genes were mainly involved in processes such as defense response, cell wall organization, and hormone signaling. This study confirms that resistance to maize common smut is highly heritable and controlled by multiple genes. The resistance-associated loci and candidate genes identified here provide important germplasm resources and genetic targets for the molecular breeding of maize varieties resistant to common smut.

Key words: maize, common smut, genome-wide association analysis, transcriptome, disease resistance candidate genes

图1

玉米瘤黑粉病抗性表型重复相关性及地点间BLUE值相关性分析 A-C: 病情指数(DI)、发病率(IR)和病程曲线下面积(AUDPC)在2年(2023、2024)2点(沈阳、锦州) 3个重复间的相关性热图。23SY、24SY、23JZ和24JZ分别表示2023年沈阳、2024年沈阳、2023年锦州和2024年锦州; 1、2、3分别代表3次重复。D: SY-DI、JZ-DI、All-DI、SY-IR、JZ-IR、All-IR、SY-AUDPC、JZ-AUDPC、All-AUDPC分别代表沈阳2年整体病情指数的BLUE值、锦州2年整体病情指数的BLUE值、2年2点整体病情指数的BLUE值、沈阳2年整体发病率的BLUE值、锦州2年整体发病率的BLUE值、2年2点整体发病率的BLUE值、沈阳2年整体病程曲线下面积的BLUE值、锦州2年整体病程曲线下面积的BLUE值、2年2点整体病程曲线下面积的BLUE值。BLUE: 最佳线性无偏估计; *、**和***分别表示P < 0.05、P < 0.01和P < 0.001。NS表示差异不显著。"

图2

玉米瘤黑粉病发病率(IR)、病情指数(DI)和病程曲线下面积(AUDPC)的频率分布图 23SY、24SY、23JZ、24JZ和BLUE分别表示2023年沈阳、2024年沈阳、2023年锦州、2024年锦州和2年2点整体BLUE值。BLUE: 最佳线性无偏估计。"

图3

玉米自交系对瘤黑粉病的抗性综合评价 HS、S、MR、R和HR分别代表高感、感病、中抗、抗病和高抗。"

图4

影响玉米瘤黑粉病的关键环境时间窗口分析 A: 病情指数(DI)关键环境窗口期确定。B: 发病率(IR)关键环境窗口确定。C: 病程曲线下面积(AUDPC)关键环境窗口确定。Tmax、Tmin、Tmean、PTT、RH和SD分别代表最高气温、最低气温、平均气温、降水量、相对湿度和日照时数。"

表1

玉米瘤黑粉病不同表型性状与气象因子的灰色关联度"

气象因子
Meteorological factor		 病情指数DI 发病率IR 病程曲线下面积AUDPC
关联度
Correlation		 排名
Rank		 关联度
Correlation		 排名
Rank		 关联度
Correlation		 排名
Rank
最高气温Maximum temperature (Tmax) 0.639 5 0.641 5 0.585 5
最低气温Minimum temperature (Tmin) 0.702 3 0.705 3 0.655 3
平均气温Mean temperature (Tmean) 0.670 4 0.673 4 0.620 4
降水量Precipitation (PTT) 0.935 1 0.933 1 0.909 1
相对湿度Relative humidity (RH) 0.405 6 0.405 6 0.347 6
日照时数Sunshine duration (SD) 0.893 2 0.898 2 0.875 2

图5

玉米关联群体的遗传结构分析 A: 基于全基因组SNP的主成分分析(PCA)散点图。B: 个体间亲缘关系矩阵热图; 对角线元素接近1, 表明个体基因型完整且自交纯合度高; 非对角线区域以浅蓝色为主, 亲缘系数普遍较低, 表明该群体个体间遗传背景差异较大, 亲缘关系相对较远; 热图未呈现明显的块状高亲缘聚类, 与PCA结果一致, 说明该群体不存在强烈的亚群分化导致的近亲聚集, 遗传结构较为分散。C: 群体结构分析结果, 不同颜色代表材料归属于不同祖先亚群的比例(K = 6)。横坐标代表421份玉米自交系材料, 纵坐标表示各自交系分布到不同亚群的概率。"

图6

玉米瘤黑粉病抗性性状的全基因组关联分析曼哈顿图 A: 病情指数(DI); B: 发病率(IR); C: 病程曲线下面积(AUDPC)。红色水平虚线表示显著性阈值(P = 2.87×10-6)。"

图7

玉米瘤黑粉病病情指数(DI)、发病率(IR)、病程曲线下面积(AUDPC)三者共同关联的SNP位点韦恩图"

图8

GWAS候选基因与转录组差异表达基因的整合分析 79个候选基因在接种玉蜀黍黑粉菌标准菌株SG200后不同时间点的表达模式热图。蓝色到红色表示表达量由低到高。Inf1 d、Inf2 d、Inf4 d、Inf6 d、Inf8 d、Inf10 d和Mock1 d、Mock2 d、Mock4 d、Mock6 d、Mock8 d、Mock10 d分别代表接菌第1、2、4、6、8、10天感病材料和接菌第1、2、4、6、8、10天对照材料。FPKM: 每百万个映射读数的转录本片段数。"

附表1

玉米抗瘤黑粉病候选基因及其功能注释"

基因编号
Gene ID		 染色体
Chromosome		 起始位置
Start position		 终止位置
End position		 描述
Description
Zm00001eb363720 8 159,957,159 159,961,745 Aminopeptidase
Zm00001eb425600 10 127,370,239 127,371,829 Endochitinase B
Zm00001eb298310 7 1,229,176 1,233,458 Monosaccharide transporter 1
Zm00001eb325320 7 160,143,290 160,148,271 Pheophytinase, chloroplastic
Zm00001eb400350 9 147,292,024 147,294,766 Putative MYB DNA-binding domain superfamily protein
Zm00001eb220250 5 22,100,009 22,104,260 Trihelix transcription factor GT-2
Zm00001eb342970 8 65,781,256 65,785,083 Unknown
Zm00001eb325790 7 161,346,807 161,349,645 Unknown
Zm00001eb152660 3 195,940,555 195,943,584 Unknown
Zm00001eb136360 3 125,744,774 125,746,538 Unknown
Zm00001eb296740 6 165,988,425 165,989,350 Calmodulin
Zm00001eb154300 3 201,227,822 201,232,263 Subtilisin-like protease SBT5.6
Zm00001eb029990 1 161,025,196 161,028,265 Unknown
Zm00001eb169930 4 20,398,975 20,399,475 Unknown
Zm00001eb116370 2 232,357,624 232,362,179 Unknown
Zm00001eb325340 7 160,215,143 160,216,461 Stress-induced protein 1
Zm00001eb404840 10 1,146,198 1,148,897 Unknown
Zm00001eb233720 5 89,152,269 89,156,855 4-coumarate-CoA ligase 1
Zm00001eb325300 7 160,098,845 160,101,078 Putative lectin-like receptor protein kinase family protein

图9

玉米抗瘤黑粉病候选基因的GO功能富集分析 FDR (错误发现率)是统计学中用于多重假设检验的错误控制指标, 定义为被错误拒绝的原假设数量占所有被拒绝原假设数量比例的期望值。"

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