小麦抗赤霉病育种回顾与展望

doi:10.3864/j.issn.0578-1752.2022.05.002

中国农业科学 ›› 2022, Vol. 55 ›› Issue (5): 837-855.doi: 10.3864/j.issn.0578-1752.2022.05.002

• 作物遗传育种·种质资源·分子遗传学 • 上一篇下一篇

小麦抗赤霉病育种回顾与展望

马鸿翔¹(),王永刚¹,高玉姣¹,何漪²,姜朋²,吴磊²,张旭²()

¹扬州大学农学院/江苏省粮食作物现代产业技术协同创新中心/江苏省作物基因组学与分子育种重点实验室,江苏扬州 225009
²江苏省农业科学院/CIMMYT-JAAS小麦病害联合研究中心/现代作物生产省部共建协同创新中心,南京 210014

收稿日期:2021-07-23 接受日期:2021-08-27 出版日期:2022-03-01 发布日期:2022-03-08
通讯作者: 张旭
作者简介:马鸿翔,E-mail： mahx@yzu.edu.cn。
基金资助:
江苏省重点研发项目(BE2021335);国家重点研发计划(2016YFD0101802);国家重点研发计划(2017YFD0100806);国家现代农业产业技术体系建设专项(CARS-03)

Review and Prospect on the Breeding for the Resistance to Fusarium Head Blight in Wheat

MA HongXiang¹(),WANG YongGang¹,GAO YuJiao¹,HE Yi²,JIANG Peng²,WU Lei²,ZHANG Xu²()

¹Agricultural College of Yangzhou University/Jiangsu Co-innovation Center of Modern Production Technology of Grain Crops/Jiangsu Key Lab of Crop Genome & Molecular Breeding, Yangzhou 225009, Jiangsu
²Jiangsu Academy of Agricultural Sciences/CIMMYT-JAAS Joint Research Center for Wheat Disease/Collaborative Innovation Center for Modern Crop Production Co-sponsored by Province and Ministry, Nanjing 210014

Received:2021-07-23 Accepted:2021-08-27 Online:2022-03-01 Published:2022-03-08
Contact: Xu ZHANG

摘要/Abstract

摘要：

由禾谷镰刀菌复合种引起的赤霉病是全世界最重要的小麦病害之一,严重影响小麦的产量和品质,受病原菌侵染的籽粒还会产生脱氧雪腐镰刀菌烯醇（deoxynivalenol,DON）为主的毒素,进一步威胁人畜健康。抗赤霉病品种选育与应用是解决小麦赤霉病及毒素危害的有效途径,中国自20世纪50年代开始抗赤霉病育种研究,70年代成立全国协作组,建立的人工接种鉴定方法被广泛用于赤霉病抗性评价,筛选出的苏麦3号、望水白等抗源被国内外广泛用于抗赤霉病研究,育成的农艺性状优良且中抗赤霉病的扬麦158号和宁麦9号等品种是抗赤霉病育种的重要突破,不仅在生产上得到大面积应用,而且作为亲本分别育成了20多个小麦品种。除常规育种外,还利用染色体工程技术将外源种质赤霉病抗性导入小麦栽培品种中,基于细胞工程技术的体细胞无性系变异和加倍单倍体创制在小麦赤霉病遗传改良中拓宽了遗传变异背景,提高了育种效率。20世纪90年代赤霉病在北美洲的流行促使欧美重视小麦赤霉病研究。国际合作促进了小麦赤霉病育种的资源、技术和信息交流,赤霉病抗性类型、鉴定技术和评价指标研究、抗赤霉病种质发掘、QTL定位、关键基因克隆、标记辅助选择及抗病品种培育等方面均取得了明显进展,利用双亲连锁作图和全基因组关联分析定位了600多个赤霉病抗性相关QTL,涉及小麦所有21条染色体,定名了Fhb1—Fhb7 7个赤霉病抗性基因/QTL。针对苏麦3号和望水白等中国抗源的主效QTL（Fhb1）,克隆了其关键基因并进行了功能验证,开发了基因的功能标记,并成功应用于标记辅助选择,培育出赤霉病抗性显著改良的小麦新品种。来自长穗偃麦草的Fhb7候选基因也被克隆,并用于提高栽培小麦品种赤霉病抗性。在精细定位的基础上,开发了紧密连锁分子标记,利用标记辅助选择将不同位点抗性QTL聚合,进一步提高了小麦赤霉病抗性。展望未来,应研究建立精准的赤霉病抗性表型评价体系、加强抗赤霉病新种质和新基因发掘、克隆主效抗病QTL的关键基因并明确其分子机制,将标记辅助选择或基因组选择与常规育种相结合,不断提升小麦抗赤霉病育种效率和水平,培育赤霉病抗性显著提高、综合性状优良的小麦品种。

关键词: 小麦, 遗传, 育种, 赤霉病, 毒素

Abstract:

Fusarium head blight (FHB) caused by Fusarium graminearum complex, is one of the most disastrous diseases seriously affecting yield and quality in wheat. Wheat kernels infected by Fusarium pathogen produce Fusarium mycotoxin, especially deoxynivalenol (DON), which may threaten the health of human beings and livestock. Breeding and application FHB resistant varieties is effective to control the disease and mycotoxin. Breeding for the resistance to FHB in wheat was commenced in 1950s in China. A national network on FHB research in wheat was established in 1970s. Inoculation methods were established and widely used for evaluating FHB resistance. Sumai 3, Wangshuibai and other resources with high resistance to FHB were selected and applied worldwide. Release of Yangmai 158 and Ningmai 9 with desirable agronomic traits and moderate resistance to FHB was a breakthrough in wheat breeding for FHB resistance. Such varieties have not only been widely applied in wheat production, but also produced more than 20 wheat varieties as parents of each variety. In addition to conventional breeding, chromosome engineering was used for transfer alien germplasm into wheat cultivated varieties, somaclonal variation and double haplotype produced by cell engineering techniques broadened genetic background and improved breeding efficiency in wheat genetic improvement for FHB resistance. The outbreak of FHB in North America in 1990s initiated the attention to FHB research in the United States and Europe. International cooperation prompted the exchange on material, technology and information in wheat breeding for FHB resistance. As results, significant progress in the research of FHB resistance type, inoculation techniques and evaluation index, germplasm development, QTL mapping, gene cloning, marker assisted selection and breeding for FHB resistance has been made. More than 600 QTL associated with FHB resistance were identified and located on all 21 chromosomes of wheat by using bi-parents linkage mapping and whole genome association analysis. Seven major genes/QTL were named as Fhb1-Fhb7. For the major QTL, Fhb1, associated with FHB resistance derived from Sumai 3 and Wangshuibai, the key candidate gene was cloned and validated, functional markers were developed and effectively used for marker assisted selection and new varieties possessing Fhb1 were released. The Fhb7 gene from Thinopyrum ponticum has also been isolated and used to improve FHB resistance in wheat. Pyramiding multiple QTL by marker assisted selection enhanced the resistance to FHB in wheat base on QTL fine mapping and close linked marker development. In the future, we should establish accurate phenotypic evaluation systems for evaluating the resistance to Fusarium head blight, strengthen the discovery of novel resistance germplasms and genes, isolate key genes related to FHB resistance and uncover their molecular mechanism, combine marker-assisted selection or genome selection with conventional breeding to continuously improve FHB resistance, breed wheat varieties with FHB resistance significantly improved and excellent agronomic traits.

Key words: wheat, genetics, breeding, Fusarium head blight, mycotoxin

马鸿翔, 王永刚, 高玉姣, 何漪, 姜朋, 吴磊, 张旭. 小麦抗赤霉病育种回顾与展望[J]. 中国农业科学, 2022, 55(5): 837-855.

MA HongXiang, WANG YongGang, GAO YuJiao, HE Yi, JIANG Peng, WU Lei, ZHANG Xu. Review and Prospect on the Breeding for the Resistance to Fusarium Head Blight in Wheat[J]. Scientia Agricultura Sinica, 2022, 55(5): 837-855.

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图/表 4

图1

表1

宁麦9号为亲本育成的小麦品种及其赤霉病抗性"

品种 Variety	审定号 Released code	亲本 Parents	抗性等级 Resistance level
镇麦5号Zhenmai 5	苏审麦200406 Jiangsu200406	扬麦158/宁麦9号 Yangmai 158/Ningmai 9	MR
宁麦14 Ningmai 14	苏审麦200601 Jiangsu200601	宁麦9号系统选择Ningmai 9 system selection	MR
宁麦13 Ningmai 13	国审麦2006004 National2006004	宁麦9号系统选择Ningmai 9 system selection	MR
镇麦8号Zhenmai 8	国审麦2006008 National2006008	扬麦158/宁麦9号 Yangmai 158/Ningmai 9	MS
生选4号Shengxuan 4	苏审麦200606 Jiangsu200606	宁麦8号/宁麦9号 DH Ningmai 8/Ningmai 9 DH	MR
扬辐麦4号Yangfumai 4	苏审麦200801 Jiangsu200801	宁麦8号/宁麦9号辐照Ningmai 8/Ningmai 9 irradiation	MR
生选6号Shengxuan 6	国审麦2009004 National2009004	宁麦8号/宁麦9号 DH Ningmai 8/Ningmai 9 DH	R
宁麦16 Ningmai 16	国审麦2009003 National2009003	宁麦8号/宁麦9号 Ningmai 8/Ningmai 9	MR
扬麦18 Yangmai 18	苏审麦200901 Jiangsu200901	4×宁麦9号/3/6×扬麦158//88-128/NNP045 4×Niangmai 9/3/6×Yangmai1 58//88-128/NNP045	MR
南农0686 Nannong 0686	国审麦2010003 National2010003	MV964091/宁麦9号MV964091/Ningmai 9	MR
扬麦21 Yangmai 21	苏审麦201102 Jiangsu201102	宁麦9号/红卷芒Ningmai 9/HJM	MR
宁麦20 Ningmai 20	苏审麦201202 Jiangsu201202	Y18//宁麦8号/宁麦9号DH Y18//Ningmai 8/Ningmai 9 DH	R
宁麦18 Ningmai 18	国审麦2012003 National2012003	宁麦9号3/杨93-111 Ningmai 93/Yang 93-111	MR
苏麦8号 Sumai 8	苏审麦201302 Jiangsu201302	宁麦9号/扬麦11 Ningmai 9/Yangmai 11	MS
宁麦21 Ningmai 21	苏审麦201303 Jiangsu201303	宁麦9号/扬麦158//宁麦9号Ningmai 9/Yangmai 158//Ningmai 9	MR
苏麦9号 Sumai 9	徽审麦201303 Anhui201303	宁麦9号/扬麦11 Ningmai 9/Yangmai 11	R
宁麦24 Ningmai 24	徽审麦201509 Anhui201509	宁麦9号系统选择Ningmai 9 system selection	MR
宁麦26 Ningmai 26	苏审麦2016004 Jiangsu2016004	宁9351/宁麦9号 Ning 9351/Ningmai 9	MR
苏麦11 Sumai 11	国审麦2016002	宁麦9号/扬麦11 Ningmai 9/Yangmai 11	MS
苏麦10号Sumai 10	徽审麦2016014 Anhui2016014	宁麦9号/扬麦11 Ningmai 9/Yangmai 11	MR
徽麦202 Huimai 202	徽审麦2016024 Anhui2016024	宁麦9号/扬麦158 Ningmai 9/Yangmai 158	MR
苏隆128 Sunong 128	徽审麦2016007 Anhui2016007	5E007/宁麦9号5E007/Ningmai 9	MR
光明麦1311 Guangmingmai 1311	国审麦20180005 National20180005	3E158/宁麦9 3E158/Ningmai 9	MR
农麦126 Nongmai 126	国审麦20180008 National20180008	扬麦16/宁麦9号Yangmai 16/Ningmai 9	MS
扬辐麦8号 Yangfumai 8	国审麦2018011 National2018011	宁麦8号/宁麦9号//扬麦11 Ningmai 8/Ningmai 9//Yangmai 11	MR
中垦麦212 Zhongkeng 212	国审麦20200005 National2020005	郑麦9023/宁麦9号Zhengmai 9023/Ningmai 9	MS
凯麦1778 Kaimai 1778	皖审麦20210026 Anhui20210026	宁麦9号/扬麦158 Ningmai 9/Yangmai 158	MR
扬麦36 Yangmai 36	国审麦20210012 National20210012	宁麦9号/扬麦152//镇麦9号Ningmai 9/Yangmai 152//Zhengmai 9	MR

表1

表2

表3

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年份 Year	审定品种数 No. of released varieties	长江中下游中抗品种 Varieties with moderate resistance to FHB in the middle to lower reaches of Yangtze River	黄淮麦区中感品种 Varieties with moderate susceptible to FHB in the reaches of Yellow River to Huai River	长江上游中感品种 Varieties with moderate susceptible to FHB in the upper reaches of Yangtze River	东北春麦区中感品种 Varieties with moderate susceptible to FHB in the Northeast China
2017	26	宁麦26 Ningmai 26	恒进麦8号、天益科麦5号、瑞华麦523 Hengjinmai 8, Tianyikemai 5, Ruihuamai 523
2018	77	光明麦1311、华麦1028、扬麦28、扬辐麦8号 Guangmingmai 1311, Huamai 1028, Yangmai 28, Yangfumai 8	瑞华麦516 Ruihuamai 516	川麦601、川农32 Chuanmai 601, Chuannong 32	北麦16、垦红24、龙辐麦23 Beimai 16, Kenhong 24, Longfumai 23
2019	59	宁麦27、苏研麦017、扬辐麦9号、扬麦30、中麦5051 Ningmai 27, Suyanmai 017, Yangfumai 9, Yangmai 30, Zhongmai 5051	安科1401、郑麦0943、中麦4072 Anke 1401, Zhengmai 0943, Zhongmai 4072		龙垦401、龙垦402 Longken 401, Longken 402
2020	106	扬辐麦10号、镇麦13、扬麦24、丰麦216、宁麦28、华麦8号 Yangfumai 10, Zhenmai 13, Yangmai 24, Fengmai 216, Ningmai 28, Huamai 8	中麦578、天益科麦6号、西农501、淮麦43、濮麦087、涡麦606、荷麦23、中育9302、大平原1号、西农369、平麦189、农麦168 Zhongmai 578, Tianyikemai 6, Xinong 501, Huaimai 43, Pumai 087, Guomai 606, Hemai 23, Zhongyu 9302, Dapingyuan 1, Xinong 369, Pingmai 189, Nongmai 168

小麦抗赤霉病育种回顾与展望

Review and Prospect on the Breeding for the Resistance to Fusarium Head Blight in Wheat

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年份 Year	审定品种数 No. of released varieties	抗或中抗品种数 No. of varieties with resistance or moderate resistance to FHB	品种 Varieties	抗性等级 FHB resistance	亲本 Parents	Fhb1位点 Fhb1 allele
2001	13	0
2002	0	0
2003	45	0
2004	25	0
2005	22	1	扬麦17 Yangmai 17	MR	92F101/川育21526 92F101/Chuanyu 21526	-
2006	32	1	宁麦13 Ningmai13	MR	宁麦9号系选 System selection from Ningmai 9	+
2007	30	1	镇麦168 Zhengmai 168	MR	苏麦6号/97G59 Sumai 6/97G59	-
2008	20	2	宁麦15 Ningmai 15	MR	宁9914系选 System selection from Ning 9914	+
2008	20	2	淮麦21 Huaimai 21	MR	淮麦17/豫麦54 Huaimai 17/Yumai 54	-
2009	33	2	宁麦16 Ningmai 16	MR	宁麦8号/宁麦9号 Ningmai 8/ Ningmai 9	+
2009	33	2	生选6号Shnegxuan 6	R	宁麦8号/宁麦9号 Ningmai 8/ Ningmai 9	+
2010	22	1	南农0686 Nannong 0686	MR	MV964091/宁麦9号MV964091/Ningmai 9	+
2011	18	0	-
2012	16	2	宁麦18 Ningmai 18	MR	宁9312/扬93-111 Ning 9312/Yang 93-111	+
2012	16	2	苏麦188 Sumai 188	MR	扬辐麦2号系选 System selection from Yangmai 2	-
2013	25	1	宁麦23 Ningmai 23	MR	宁9543/扬麦158//宁9543 Ning 9543/Yangmai 158//Ning 9543	+
2014	21	0	-
2015	34	1	华麦6号 Huamai 6	MR	扬麦13/苏麦6号 Yangmai 13/Sumai 6	-

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