张晓军^1,2, 肖进³, 王海燕³, 乔麟轶¹, 李欣¹, 郭慧娟¹, 常利芳¹, 张树伟¹, 阎晓涛¹, 畅志坚^,1,2,*, 武宗信^,4,*1 山西省农业科学院作物科学研究所/作物遗传与分子改良山西省重点实验室, 山西太原 030031
2 农业部黄土高原作物基因资源与种质创制重点实验室, 山西太原 030031
3 南京农业大学作物遗传与种质创新国家重点实验室, 江苏南京 210095
4 山西省农业科学院棉花研究所, 山西运城 044000

Evaluation of resistance to Fusarium head blight in Thinopyrum-derived wheat lines

ZHANG Xiao-Jun^1,2, XIAO Jin³, WANG Hai-Yan³, QIAO Lin-Yi¹, LI Xin¹, GUO Hui-juan¹, CHANG Li-Fang¹, ZHANG Shu-Wei¹, YAN Xiao-Tao¹, CHANG Zhi-Jian^,1,2,*, WU Zong-Xin^,4,* 1 Institute of Crop Science, Shanxi Academy of Agricultural Sciences/Shanxi Key Laboratory of Crop Genetics and Molecular Improvement, Taiyuan 030006, Shanxi, China
2 Key Laboratory of Crop Gene Resources and Germplasm Enhancement on Loess Plateau of Ministry of Agriculture, Taiyuan 030006, Shanxi, China
3 State Key Laboratory of Crop Genetics and Germplasm Enhancement, Nanjing Agricultural University, Nanjing 210095, Jiangsu, China
4 Institute of Cotton, Shanxi Academy of Agricultural Sciences, Yuncheng 044000, Shanxi, China

通讯作者: *畅志坚, E-mail: wrczj@126.com。 武宗信, E-mail: mhdwzx@126.com

收稿日期:2019-02-16接受日期:2019-08-9网络出版日期:2019-09-03

基金资助:

本研究由国家重点研发计划项目.2017YFD0100600 山西省重点研发计划项目.201803D221018-5 山西省重点研发计划项目.201703D211007 山西省重点研发计划项目.201803D421020 山西省农业科学院项目.YGG17123 山西省农业科学院项目.YCX2018D2YS01 和山西省重点科技创新平台资助.201605D151002

Received:2019-02-16Accepted:2019-08-9Online:2019-09-03

Fund supported:

This study was supported by the National Key R&D Program of China.2017YFD0100600 Key R&D Program of Shanxi Province.201803D221018-5 Key R&D Program of Shanxi Province.201703D211007 Key R&D Program of Shanxi Province.201803D421020 Shanxi Academy of Agricultural Sciences.YGG17123 Shanxi Academy of Agricultural Sciences.YCX2018D2YS01 Shanxi Key Scientific and Technological Innovation Platform.201605D151002

作者简介 About authors
E-mail:zxjemail@163.com。













摘要
由镰孢属(Fusarium)真菌侵染引起的赤霉病是严重威胁小麦生产的重要病害之一, 但小麦育种中可直接利用的抗源非常有限。采用单花滴注法接种赤霉菌株F0609, 对来源于中间偃麦草或长穗偃麦草的119份小偃麦衍生品系进行3年6个环境的抗病鉴定, 发现平均病小穗率<10%的材料有13份, 抗性评价为抗病(R); 平均病小穗率介于10%~25%之间的材料有61份, 抗性评价为中抗(MR); 其余45份材料的平均病小穗率介于25%~50%或>50%, 抗性评价为中感或高感(MS和S)。在13份高抗赤霉病材料中, CH16387的抗性显著优于苏麦3号和望水白, CH16371和CH16379的抗性显著优于望水白, 其余10个品系与抗病对照苏麦3号和望水白的抗性水平相当。这13份材料分别来自小麦-中间偃麦草部分双二倍体TAI8045和小麦-长穗偃麦草部分双二倍体TAP8430与普通小麦的杂交组合, TAI8045抗性显著优于对照品种望水白, TAP8430与苏麦3号和望水白的抗性相当, 而杂交组合中的小麦亲本对赤霉病表现感病, 推测这些材料的抗性可能来自TAI8045和TAP8430。这些抗病材料为小麦抗赤霉病育种提供了新的种质资源。
关键词： 小麦;赤霉病;偃麦草;遗传改良;种质资源

Abstract
Fusarium head blight (FHB) caused by Fusarium graminearum is one of the most destructive fungal diseases in wheat production; however, only limited sources of resistance are available in wheat. In this study, we evaluated 119 lines derived from the crosses between wheat and wheat-Thinopyrum partial amphiploids for their resistance to F. graminearum isolate F0609 over six environments during 2016 to 2018 cropping seasons using single floret inoculation method. Among the wheat-Thinopyrum lines tested, 45 were moderately or highly susceptible, with 25%-50% or >50% of the average percentage of diseased spikelets (PDS), 61 were moderately resistant (MR) with 10%-25% of the average PDS, and 13 lines were identified as resistant (R), with the average PDS less than 10%. For the FHB resistance of the 13 resistant lines, CH16387 was superior to ‘Sumai 3’ and ‘Wangshuibai’, the most widely used source of resistance to FHB, CH16371 and CH16379 were superior to ‘Wangshuibai’, and the remaining ten lines were comparable to ‘Wangshuibai’ or ‘Sumai 3’, in terms of number of infected spikelets per spike and percentage of infected spikelets. Furthermore, the average PDS in these resistant lines over the six environments showed a similar distribution, suggesting a relatively stable FHB resistance. The donor parents, wheat-alien partial amphiploids, involved in development of these resistant derivatives, included wheat-Th. intermedium partial amphiploid TAI8045 and wheat-T.ponticum partial amphiploid TAP8430. As both TAI8045 and TAP8430 were resistant, but all the wheat parents were susceptible, it was likely that the resistance to FHB in these lines identified originated from TAI8045 and TAP8430. These derivatives can serve as novel sources to enhance resistance of wheat to FHB.
Keywords：wheat;Fusarium head blight;Thinopyrum;genetic improvement;germplasm resources


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本文引用格式
张晓军, 肖进, 王海燕, 乔麟轶, 李欣, 郭慧娟, 常利芳, 张树伟, 阎晓涛, 畅志坚, 武宗信. 小偃麦衍生品系的赤霉病抗性评价[J]. 作物学报, 2020, 46(1): 62-73. doi:10.3724/SP.J.1006.2020.91015
ZHANG Xiao-Jun, XIAO Jin, WANG Hai-Yan, QIAO Lin-Yi, LI Xin, GUO Hui-juan, CHANG Li-Fang, ZHANG Shu-Wei, YAN Xiao-Tao, CHANG Zhi-Jian, WU Zong-Xin. Evaluation of resistance to Fusarium head blight in Thinopyrum-derived wheat lines[J]. Acta Agronomica Sinica, 2020, 46(1): 62-73. doi:10.3724/SP.J.1006.2020.91015


由镰孢属(Fusarium)真菌侵染引起的小麦赤霉病不仅会导致小麦大幅减产, 而且, 病原菌产生的脱氧雪腐镰孢菌烯醇(deoxynivalenol, DON)等多种毒素污染还会危害人畜健康^[1]。近些年随着全球气候变暖加剧, 以及轮作倒茬、秸秆还田等耕作制度的改变, 小麦赤霉病的高发区已由长江中下游、淮河流域冬麦区和东北春麦区向西南冬麦区、黄淮冬麦区迅速扩展, 直接威胁小麦主产区的安全生产^[2]。因此, 开展抗赤霉病育种, 从根本上控制赤霉病的流行与危害对确保国家粮食安全具有重大意义。

小麦赤霉病的研究和防治一直备受关注^[3,4,5]。但由于抗源严重匮乏, 加之抗性表现形式的多样性和遗传方式的复杂性, 使得抗赤霉病育种进展缓慢。我国黄淮、西南、北部冬麦区及东北春麦区的主栽品种对赤霉病的抗性普遍较弱, 可利用的抗源也非常有限。迄今为止, 已从普通小麦及其近缘种属中鉴定出200多个抗赤霉病QTL^[6], 分布于小麦各染色体上, 但仅有7个主效抗病基因得到精确定位, 分别是来自“苏麦3号”3BS上的Fhb1^[7]和6BS上的Fhb2^[8]、“望水白”4B上的Fhb4^[9]和5A上的Fhb5^[10], 以及分别从大赖草7Lr#1S、披碱草1E^ts#1S和长穗偃麦草7E上转移的Fhb3^[11]、Fhb6^[12]和Fhb7^[13]。因此, 广泛发掘并筛选新的抗赤霉病种质资源, 特别是主效基因控制的抗病材料, 对提高栽培品种的赤霉病抗性, 拓展抗源的遗传基础, 满足小麦育种及生产需求十分必要。

2014—2015年度我们在山西省运城市播种了1100余份小偃麦衍生品系, 恰逢赤霉病严重发生, 从中筛选出119份未感病或感病较轻的品系。2016— 2018年分别在江苏南京、山西太原和四川成都通过单花滴注法对119份小偃麦衍生品系进行了赤霉病抗性鉴定与评价, 以期发掘小麦抗赤霉病优异种质资源。

1 材料与方法

1.1 供试材料

供试的119份小偃麦衍生品系(附表1)分别由TAI8045、TAI8335、TAI8505和TAP8430与普通小麦品种冀麦26、中8701、晋春5号、太原768、晋麦33和京繁309杂交1~2次后自交选育而来。TAI8045、TAI8335和TAI8505是普通小麦与中间偃麦草杂交选育的部分双二倍体小偃麦, TAP8430是普通小麦与十倍体长穗偃麦草杂交选育的部分双二倍体小偃麦。TAI8045、TAI8505、TAI8335、TAP8430以及普通小麦品种太原768、晋春5号和冀麦26均来自山西省农业科学院作物科学研究所; 普通小麦中8701来自中国农业科学院种质资源库(编号为ZM20831)。苏麦3号和望水白为抗病对照品种, 周麦27和Alondra’s为感病对照品种。其中, 苏麦3号、望水白和Alondra’s由南京农业大学作物遗传与种质创新国家重点实验室提供, 周麦27由河南省农业科学院小麦研究所提供。

Supplementary table 1
附表1
附表1119份小偃麦衍生品系不同环境下的赤霉病抗性
Supplementary table 1Responses of the 119 wheat lines derived from Thinopyrum to Fusarium head blight under different environments

编号 No.	材料 名称 Lines	6个环境病小穗率 PDS in six environments (%)							6个环境病小穗率分布 PDS distribution in six environments				严重度分级 Severity degree	抗性评价 FHB resistance
		2018 太原 Taiyuan	2018 成都 Chengdu	2018 南京 Nanjing	2017 太原 Taiyuan	2017 南京 Nanjing	2016 南京 Nanjing		最小 Min	最大 Max	极差 Range	平均 Average
1	苏麦3号 Sumai 3	16.8	3.6	7.9	5.1	4.0	5.1		3.6	16.8	13.2	7.1	1	R
2	望水白 Wangshuibai	13.9	6.1	12.3	9.6	7.5	6.8		6.1	13.9	7.8	9.4	1	R
3	CH16387	8.7	4.9	4.4	4.5	1.3	1.3		1.3	8.7	7.4	4.2**a	1	R
4	CH16371	15.1	7.3	4.6	4.6	2.0	3.0		2.0	15.1	13.1	5.0**b	1	R
5	CH16379	7.0	10.2	7.2	5.1	2.0	4.2		2.0	10.2	8.2	6.0*b	1	R
6	CH16373	10.5	4.2	5.1	4.8	10.0	4.0		4.0	10.5	6.5	6.7	1	R
7	CH16352	7.3	11.6	5.5	4.2	-	6.0		4.2	11.6	7.3	6.7	1	R
8	CH16388	6.5	7.8	14.5	4.0	4.6	2.0		2.0	14.5	12.5	6.7	1	R
9	CH16419	6.5	11.4	6.6	4.8	-	4.0		4.0	11.4	7.4	6.7	1	R
10	CH16367	12.5	9.2	9.3	4.6	2.8	3.0		2.8	12.5	9.7	7.0	1	R
11	CH16378	11.0	5.1	-	4.9	-	-		4.9	11.0	6.1	7.1	1	R
12	CH16374	9.4	9.8	8.3	5.4	9.2	3.0		3.0	9.8	6.8	7.5	1	R
13	CH16375	12.4	10.4	7.1	5.3	8.6	2.0		2.0	12.4	10.4	7.6	1	R
14	CH16427	8.4	10.5	-	5.9	-	6.7		5.9	10.5	4.6	7.7	1	R
15	CH16432	5.8	12.1	-	12.7	-	4.6		4.6	12.7	8.1	8.4	1	R
16	CH16390	6.7	17.2	20.0	3.7	9.0	4.0		3.7	20.0	16.3	10.1	2	MR
17	CH16424	-	17.6	-	6.8	-	6.1		6.1	17.6	11.5	10.2	2	MR
18	CH16399	7.1	10.6	-	7.3	-	17.0		7.1	17.0	9.8	10.5	2	MR
19	CH16431	11.9	9.5	-	6.5	-	15.0		6.5	15.0	8.6	10.7	2	MR
20	CH16369	11.3	11.5	14.0	16.0	10.0	3.0		3.0	16.0	13.0	11.0	2	MR
21	CH16389	11.4	14.1	17.6	15.7	5.0	3.0		3.0	17.6	14.6	11.1	2	MR
22	CH1357	11.2	13.3	-	10.0	-	11.0		10.0	13.3	3.3	11.4	2	MR
编号 No.	材料 名称 Lines	6个环境病小穗率 PDS in six environments (%)							6个环境病小穗率分布 PDS distribution in six environments				严重度分级 Severity degree	抗性评价 FHB resistance
		2018 太原 Taiyuan	2018 成都 Chengdu	2018 南京 Nanjing	2017 太原 Taiyuan	2017 南京 Nanjing	2016 南京 Nanjing		最小 Min	最大 Max	极差 Range	平均 Average
23	CH16423	5.0	12.3	12.0	11.1	23.0	6.0		5.0	23.0	18.0	11.6	2	MR
24	CH16392	16.9	14.9	-	10.0	-	4.6		4.6	16.9	12.2	11.6	2	MR
25	CH16346	24.3	8.1	-	4.1	-	10.0		4.1	24.3	20.2	11.6	2	MR
26	CH16381	15.8	10.2	24.0	8.6	8.0	5.0		5.0	24.0	19.0	11.9	2	MR
27	CH16118	8.5	20.4	-	8.1	-	12.1		8.1	20.4	12.3	12.3	2	MR
28	CH16372	23.7	19.3	14.0	9.1	5.0	3.0		3.0	23.7	20.7	12.3	2	MR
29	CH16368	27.4	11.0	20.0	4.8	12.0	2.0		2.0	27.4	25.4	12.9	2	MR
30	CH16426	24.0	12.0	-	7.1	-	8.8		7.1	24.0	16.9	12.9	2	MR
31	CH16360	13.4	7.1	-	4.7	-	27.0		4.7	27.0	22.3	13.0	2	MR
32	CH16380	10.8	15.8	29.0	18.4	1.0	4.0		1.0	29.0	28.0	13.2	2	MR
33	CH16409	17.6	10.2	-	19.5	-	6.0		6.0	19.5	13.5	13.3	2	MR
34	CH16359	22.6	18.5	8.0	19.1	12.0	2.0		2.0	22.6	20.6	13.7	2	MR
35	CH16417	10.8	12.3	-	12.0	-	20.4		10.8	20.4	9.6	13.9	2	MR
36	CH16408	5.0	28.5	-	13.3	-	8.8		5.0	28.5	23.5	13.9	2	MR
37	CH16376	42.7	6.2	11.0	5.6	15.0	3.0		3.0	42.7	39.7	13.9	2	MR
38	CH1349	24.5	7.5	-	6.8	-	17.0		6.8	24.5	17.7	14.0	2	MR
39	CH1556	24.5	13.3	-	6.3	-	12.1		6.3	24.5	18.2	14.0	2	MR
40	CH16366	11.4	8.0	8.0	23.9	28.0	6.0		6.0	28.0	22.0	14.2	2	MR
41	CH16382	8.7	18.2	26.0	22.8	11.0	5.0		5.0	26.0	21.0	15.3	2	MR
42	CH16391	11.3	13.1	29.0	6.8	29.0	3.0		3.0	29.0	26.0	15.4	2	MR
43	CH16403	23.1	20.9	-	7.1	-	11.0		7.1	23.1	16.1	15.5	2	MR
44	CH16377	14.0	16.9	-	6.2	-	26.0		6.2	26.0	19.8	15.8	2	MR
45	CH16440	23.6	12.6	-	11.0	-	17.6		11.0	23.6	12.6	16.2*	2	MR
46	CH16439	27.8	15.4	-	13.1	-	9.5		9.5	27.8	18.3	16.5	2	MR
47	CH16404	20.2	13.7	14.9	25.4	21.0	4.0		4.0	25.4	21.4	16.5*	2	MR
48	CH16342	29.7	23.7	11.0	9.6	17.0	10.0		9.6	29.7	20.1	16.8*c	2	MR
49	CH16393	17.5	6.7	-	30.8	-	13.3		6.7	30.8	24.1	17.1	2	MR
50	CH16425	21.8	20.5	-	6.5	-	20.2		6.5	21.8	15.3	17.2*	2	MR
51	CH16386	13.3	19.5	21.9	26.4	15.0	8.0		8.0	26.4	18.4	17.3*	2	MR
52	CH16397	8.8	17.4	50.0	8.7	18.0	3.0		3.0	50.0	47.0	17.7	2	MR
53	CH16406	20.4	33.8	-	4.6	-	12.0		4.6	33.8	29.2	17.7	2	MR
54	CH16384	23.9	10.4	13.0	19.6	30.0	10.0		10.0	30.0	20.0	17.8*	2	MR
55	CH16418	23.0	31.0	-	6.1	-	12.0		6.1	31.0	24.9	18.0	2	MR
56	CH16394	31.9	14.7	-	13.1	-	12.5		12.5	31.9	19.5	18.1	2	MR
57	CH16104	29.7	18.7	-	18.4	-	10.0		10.0	29.7	19.7	19.2*	2	MR
58	CH16365	-	26.2	14.0	4.0	46.0	7.0		4.0	46.0	42.0	19.4	2	MR
59	CH16422	18.8	31.9	26.0	6.0	30.0	5.0		5.0	31.9	26.9	19.6*c	2	MR
60	CH16412	14.4	37.3	-	14.9	-	14.0		14.0	37.3	23.3	20.1*c	2	MR
61	CH16430	27.1	17.6	-	28.4	-	11.0		11.0	28.4	17.4	21.0*	2	MR
62	CH16442	30.9	14.5	-	21.4	-	19.5		14.5	30.9	16.4	21.6**	2	MR
63	CH16370	37.5	35.3	7.0	36.9	11.0	3.0		3.0	37.5	34.5	21.8*	2	MR
编号 No.	材料 名称 Lines	6个环境病小穗率 PDS in six environments (%)							6个环境病小穗率分布 PDS distribution in six environments				严重度分级 Severity degree	抗性评价 FHB resistance
		2018 太原 Taiyuan	2018 成都 Chengdu	2018 南京 Nanjing	2017 太原 Taiyuan	2017 南京 Nanjing	2016 南京 Nanjing		最小 Min	最大 Max	极差 Range	平均 Average
64	CH16416	49.9	8.3	-	7.9	-	-		7.9	49.9	42.0	22.0*	2	MR
65	CH16445	26.0	26.2	-	18.8	-	17.6		17.6	26.2	8.6	22.2**	2	MR
66	CH16421	13.4	51.1	-	12.1	-	12.0		12.0	51.1	39.1	22.2*	2	MR
67	CH16420	11.1	40.6	-	26.1	-	11.0		11.0	40.6	29.6	22.2*c	2	MR
68	CH16351	32.0	12.1	-	35.9	-	9.4		9.4	35.9	26.6	22.4*	2	MR
69	CH16341	23.9	16.0	-	18.4	-	32.0		16.0	32.0	16.0	22.6**	2	MR
70	CH16443	27.9	16.3	-	23.9	-	-		16.3	27.9	11.6	22.7**	2	MR
71	CH16449	-	25.3	-	20.2	-	23.6		20.2	25.3	5.1	23.0**	2	MR
72	CH16112	30.1	20.3	-	15.0	-	27.0		15.0	30.1	15.1	23.1**	2	MR
73	CH16383	30.6	14.0	-	15.7	-	33.0		14.0	33.0	19.0	23.3**a	2	MR
74	CH16414	40.2	20.3	-	6.7	-	27.1		6.7	40.2	33.4	23.6*	2	MR
75	CH16401	17.1	8.3	85.0	11.4	17.0	3.0		3.0	85.0	82.0	23.6*	2	MR
76	CH16415	21.5	6.7	-	21.4	-	49.0		6.7	49.0	42.3	24.6*	2	MR
77	CH16410	15.0	38.6	-	21.8	-	-		15.0	38.6	23.6	25.1*	3	MS
78	CH16435	25.3	-	-	19.5	-	31.0		19.5	31.0	11.5	25.2**	3	MS
79	CH16362	38.1	33.9	-	13.0	-	18.0		13.0	38.1	25.0	25.8*	3	MS
80	CH16438	16.9	31.9	-	15.4	-	39.7		15.4	39.7	24.3	26.0**	3	MS
81	CH16348	32.1	41.4	-	18.8	-	13.0		13.0	41.4	28.4	26.3*	3	MS
82	CH16364	18.8	53.6	13.0	8.1	59.0	6.0		6.0	59.0	53.0	26.4*	3	MS
83	CH1364	36.5	21.5	-	31.1	-	17.0		17.0	36.5	19.5	26.6**	3	MS
84	CH16402	33.3	21.5	-	27.4	-	25.0		21.5	33.3	11.9	26.8**	3	MS
85	CH16396	12.2	34.1	35.3	40.0	-	13.0		12.2	40.0	27.9	26.9**	3	MS
86	CH16385	30.0	34.2	-	23.7	-	20.2		20.2	34.2	14.0	27.0**	3	MS
87	CH16343	31.1	60.6	9.0	28.2	-	8.0		8.0	60.6	52.6	27.4*	3	MS
88	CH16433	18.6	47.4	-	18.8	-	26.2		18.6	47.4	28.9	27.8*	3	MS
89	CH16340	28.8	32.3	-	31.1	-	19.1		19.1	32.3	13.2	27.8**	3	MS
90	CH16437	39.7	20.5	-	21.8	-	31.9		20.5	39.7	19.1	28.5**	3	MS
91	CH16407	38.2	43.3	-	18.9	-	14.1		14.1	43.3	29.2	28.6*	3	MS
92	CH16405	26.4	11.4	85.0	9.5	31.0	9.0		9.0	85.0	76.0	28.7*	3	MS
93	CH16354	52.7	12.9	-	30.5	-	19.3		12.9	52.7	39.8	28.8*	3	MS
94	CH16444	38.1	23.1	-	31.0	-	29.0		23.1	38.1	15.0	30.3**	3	MS
95	CH16395	16.8	52.3	91.0	6.0	14.0	2.0		2.0	91.0	89.0	30.4*	3	MS
96	CH16428	33.4	40.2	-	11.3	-	38.4		11.3	40.2	28.9	30.8**	3	MS
97	CH16429	31.8	31.8	-	27.8	-	31.9		27.8	31.9	4.1	30.8**	3	MS
98	CH16413	23.2	44.4	-	24.4	-	31.8		23.2	44.4	21.2	31.0**	3	MS
99	CH16344	38.6	23.1	-	36.0	-	30.0		23.1	38.6	15.5	31.9**	3	MS
100	CH16358	35.6	46.8	-	14.0	-	34.2		14.0	46.8	32.9	32.6**	3	MS
101	CH16441	52.2	19.2	-	31.0	-	30.0		19.2	52.2	33.0	33.1**	3	MS
102	CH16398	30.1	49.8	-	16.5	-	37.5		16.5	49.8	33.3	33.5**	3	MS
103	CH16448	6.1	57.5	-	38.5	-	31.8		6.1	57.5	51.5	33.5*	3	MS
104	CH16447	31.7	37.7	-	40.2	-	27.1		27.1	40.2	13.0	34.2**	3	MS
编号 No.	材料 名称 Lines	6个环境病小穗率 PDS in six environments (%)							6个环境病小穗率分布 PDS distribution in six environments				严重度分级 Severity degree	抗性评价 FHB resistance
		2018 太原 Taiyuan	2018 成都 Chengdu	2018 南京 Nanjing	2017 太原 Taiyuan	2017 南京 Nanjing	2016 南京 Nanjing		最小 Min	最大 Max	极差 Range	平均 Average
105	CH16363	51.7	79.7	29.0	24.6	20.0	2.0		2.0	79.7	77.7	34.5*	3	MS
106	CH16347	53.9	37.2	-	30.1	-	24.4		24.4	53.9	29.5	36.4**	3	MS
107	CH16446	46.0	41.0	-	28.2	-	34.1		28.2	46.0	17.8	37.3**	3	MS
108	CH16361	62.7	42.8	-	9.0	-	-		9.0	62.7	53.7	38.2*	3	MS
109	CH1673	50.8	-	57.0	32.1	-	16.0		16.0	57.0	41.0	39.0**	3	MS
110	CH16436	40.8	46.2	-	23.1	-	51.1		23.1	51.1	28.0	40.3**	3	MS
111	CH16411	17.0	75.4	51.1	24.8	-	39.7		17.0	75.4	58.4	41.6**	3	MS
112	CH16355	47.9	64.0	-	23.2	-	34.1		23.2	64.0	40.8	42.3**	3	MS
113	CH16345	33.4	73.8	-	21.5	-	46.0		21.5	73.8	52.2	43.7**	3	MS
114	CH16400	47.4	80.2	74.9	20.6	-	15.0		15.0	80.2	65.2	47.6**	3	MS
115	CH16357	40.2	54.5	62.7	27.8	-	55.0		27.8	62.7	34.9	48.0**	3	MS
116	CH16356	36.3	58.6	-	37.5	-	60.7		36.3	60.7	24.5	48.3**	3	MS
117	CH16353	53.2	82.1	-	43.1	-	21.1		21.1	82.1	61.0	49.9**	3	MS
118	CH16434	38.4	96.3	-	31.0	-	35.3		31.0	96.3	65.2	50.2**	4	S
119	CH16106	78.4	61.4	-	40.2	-	41.4		40.2	78.4	38.3	55.3**	4	S
120	CH16350	63.8	93.9	-	53.6	-	55.6		53.6	93.9	40.4	66.7**	4	S
121	CH16349	95.0	100.0	-	33.6	-	46.0		33.6	100.0	66.4	68.6**	4	S
122	周麦27 Zhoumai 27	64.3	57.0	63.0	58.6	38.6	53.9		38.6	64.3	25.8	55.9**	4	S
123	Alondra’s	53.1	65.5	63.8	51.7	48.1	61.4		48.1	65.5	17.4	57.3**	4	S

R: 抗病; MR: 中抗; MS: 中感; S: 感病; “-”: 未调查或数据缺失。苏麦3号, 望水白: 抗病对照; 周麦27, Alondra’s: 感病对照。^a: 与抗病对照苏麦3号有显著差异(P < 0.05), 与望水白极显著差异(P < 0.01); ^b: 仅与抗病对照望水白差异显著(P < 0.05), 与苏麦3号无显著差异; ^c: 仅与苏麦3号有显著差异(P < 0.05), 与望水白无显著差异。^*、^**分别表示在P < 0.05、P < 0.01水平上与抗病对照有显著差异。
R: resistant; MR: moderately resistant; MS: moderately susceptible; S: susceptible;“-”: no results or missing data. PDS: percentage of diseased spikelet. Sumai 3, Wangshuibai: resistant control; Zhoumai 27, Alondra’s: susceptible control. ^a: significantly different from Sumai 3 at P < 0.05, and from Wangshuibai at P < 0.01. ^b: significantly different from Wangshuibai at P < 0.05, but not from Sumai 3. ^c: significantly different from Sumai 3 at P < 0.05, but not from Wangshuibai. ^*, ^**: significantly different from the resistant control at P < 0.05, P < 0.01, respectively.

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1.2 赤霉病鉴定地点及接种方法

2016、2017和2018年在江苏省南京市南京农业大学江浦基地日光温室, 2017年和2018年在山西省太原市山西省农业科学院东阳基地日光温室, 2018年在四川省成都市四川省农业科学院新都基地田间分别进行接种鉴定。接种赤霉菌菌株为F0609, 由南京农业大学作物遗传与种质创新国家重点实验室王秀娥博士惠赠。

播种鉴定材料于温室或田间, 行长1.5~2.0 m, 每行播种30粒, 行距0.25 m。在小麦完全抽穗后开始扬花时选取中部小穗采用单花滴注法^[14]接种赤霉病, 接种剂量为10 μL (约50~80个孢子 μL^-1), 每行材料接种10个穗子, 接种后用保鲜膜包裹保湿3~5 d以使发病, 颖壳变褐色时揭开保鲜膜, 每日喷雾保湿4次以上, 以保证发病环境的湿度。

1.3 赤霉病抗性调查与记载

接种后27 d开始调查发病小穗数, 调查每个材料10个单穗, 抗病调查和记载标准按照《中华人民共和国农业行业标准NY/T 1443.4-2007: 小麦抗赤霉病评价技术规范》^[15]。小麦赤霉病严重度按病小穗率(percentage of diseased spikelet, PDS)分5级: 免疫(I), 平均严重度为0, 接种小穗无可见发病症状; 高抗(R), 0<平均严重度<2.0, 仅接种小穗发病, 或相邻的个别小穗发病, 但病斑不扩展到穗轴; 中抗(MR), 2.0≤平均严重度<3.0, 穗轴发病, 发病小穗占总小穗数的1/4以下; 中感(MS), 3.0≤平均严重度<3.5, 穗轴发病, 发病小穗占总小穗数的1/4~1/2; 高感(S), 平均严重度≥3.5, 穗轴发病, 发病小穗占总小穗数的1/2以上。

1.4 染色体数目鉴定

采用Han等^[16]描述的方法鉴定染色体数目, 滴片后在Olympus BX53相差显微镜下观察, 每个材料5~10个根尖, 选择分裂相完整的细胞进行染色体计数。

1.5 统计分析

利用SAS (Statistical Analysis System) v9.2对不同环境小麦赤霉病病小穗率进行基本数据统计和相关分析。利用Microsoft Excel 2013进行t测验分析。

2 结果与分析

2.1 亲本材料抗赤霉病鉴定与评价

来源于中间偃麦草的TAI8045表现抗病, 病小穗率显著低于抗病对照望水白(P<0.05); TAI8505表现中抗, 病小穗率与望水白无显著差异, 但与苏麦3号有显著差异; TAI8335表现中感, 病小穗率与2个抗病对照存在极显著差异(P<0.01); 来源于长穗偃麦草的TAP8430表现抗病, 病小穗率与苏麦3号和望水白均无显著差异; 其他普通小麦亲本冀麦26、中8701、晋春5号和太原768均表现感病, 严重度3~4级(表1)。未鉴定普通小麦亲本晋麦33和京繁309。

Table 1
表1
表1小偃麦衍生品系亲本的病小穗率及抗性评价
Table 1Fusarium head blight (FHB) resistance and percentage of diseased spikelet (PDS) in the parents of Thinopyrum-derived wheat lines

亲本来源 Source of parents	亲本名称 Name of parents	病小穗率 PDS (%)				严重度分级 Severity degree	抗性评价 FHB resistance	染色体数目 Number of chromosomes
		最小 Min.	最大 Max.	极差 Range	平均 Average
长穗偃麦草Th. ponticum	TAP8430	4.4	5.9	1.5	5.4	1	R	56
中间偃麦草Th. intermedium	TAI8045	1.1	4.5	3.4	3.3*a	1	R	56
	TAI8505	7.0	20.8	13.8	14.6**b	2	MR	56
	TAI8335	20.4	65.4	45.0	34.5**	3	MS	56
普通小麦Wheat	太原768 Taiyuan 768	36.0	50.0	14.0	42.1**	3	MS	42
	冀麦26 Jimai 26	40.0	52.2	12.2	47.3**	3	MS	42
	晋春5号Jinchun 5	48.5	70.0	21.5	58.7**	4	S	42
	中8701 Zhong 8701	29.2	84.0	54.8	63.6**	4	S	42
	苏麦3号Sumai 3	1.4	24.0	22.6	7.1	1	R	42
	望水白Wangshuibai	1.4	28.0	26.6	9.4	1	R	42
	周麦27 Zhoumai 27	38.6	64.3	25.7	55.9**	4	S	42
	Alondra’s	48.1	65.5	17.4	57.3**	4	S	42

R: resistant; MR: moderately resistant; MS: moderately susceptible; S: susceptible. ^*, ^**: significantly different from the resistant control at P < 0.05, P < 0.01, respectively. ^a: significantly different from Wangshuibai at P < 0.05, but not from Sumai 3; ^b: significantly different from Sumai 3 at P < 0.01, but not from Wangshuibai.
R: 抗病; MR: 中抗; MS: 中感; S: 感病。^*、^**分别表示在P < 0.05, P < 0.01水平上与抗病对照有显著差异; ^a: 仅与望水白在P < 0.05水平上显著, 与苏麦3号不显著; ^b: 仅与苏麦3号在P < 0.01水平上差异显著, 与望水白差异不显著。

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图1

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图1小偃麦衍生品系亲本对赤霉病的抗性反应(山西太原, 2018)

Fig. 1Responses to Fusarium head blight in the parents (Taiyuan, Shanxi, 2018)

2.2 119份小偃麦衍生品系赤霉病抗性鉴定

2016—2018年3年6点的抗病鉴定中, 抗病对照苏麦3号和望水白的病害严重度均为1级, 表现高抗或中抗, 二者病小穗率无显著差异。感病对照Alondra’s和周麦27的病害严重度均为4级, 都表现高感。119份小偃麦衍生系接种后均有侵染反应, 未发现病小穗率为0、赤霉病抗性免疫(I)的品系(附表1)。平均病小穗率<10%的材料有13份, 抗性分级为抗病(R), 占鉴定材料的10.9%; 平均病小穗率介于10%~25%之间的材料有61份, 抗性评价为中抗(MR), 占51.3%; 平均病小穗率介于25%~50%之间的材料有41份, 表现中感(MS), 占34.5%; 平均病小穗率大于50%的材料有4份, 表现感病(S)(表2和表3)。鉴定材料分别来自9个杂交组合(表2), 其中以小麦-长穗偃麦草部分双二倍体TAP8430为亲本的3个组合共有73份材料, 4份表现抗病(R), 36份表现中抗(MR), 占组合材料总数的54.8%; 以小麦-中间偃麦草部分双二倍体TAI8045为亲本的3个组合共有37份材料, 9份表现抗病(R), 21份表现中抗(MR), 占组合材料总数的81.1%; 以小麦-中间偃麦草部分双二倍体TAI8335为亲本的2个组合共6份材料, 1份表现中抗(MR), 其余5份为中感(MS); 以小麦-中间偃麦草部分双二倍体TAI8505为亲本的3份材料均表现中抗(MR)。

Table 2
表2
表2119份小偃麦衍生品系的赤霉病抗性评价
Table 2Responses to Fusarium head blight in 119 wheat lines derived from Thinopyrum

供体亲本Donor parent	系谱来源 Pedigree	品系个数 Number of lines	抗病 R	中抗 MR	中感 MS	感病 S
TAP8430	太原768//冀麦26/TAP8430 Taiyuan 768//Jimai 26/TAP8430	73	1	20	19	4
	中8701//冀麦26/TAP8430 Zhong 8701//Jimai 26/TAP8430		2	14	4
	中8701//京繁309/TAP8430 Zhong 8701//Jingfan 309/TAP8430		1	2	6
TAI8045	京繁309//晋春5号/TAI8045 Jingfan 309//Jinchun 5/TAI8045	37	8	11	1
	太原768//冀麦26/TAI8045 Taiyuan 768//Jimai 26/TAI8045		1	7	4
	京繁309//冀麦26/TAI8045 Jingfan 309//Jimai 26/TAI8045			3	2
TAI8335	中8701//晋麦33/TAI8335 Zhong 8701//Jinmai 33/TAI8335	6		1	2
	冀麦26//晋麦33/TAI8335 Jimai 26//Jinmai 33/TAI8335				3
TAI8505	太原768//冀麦26/TAI8505 Taiyuan 768//Jimai 26/ATI8505	3		3
合计 Total		119	13	61	41	4

R: resistant; MR: moderately resistant; MS: moderately susceptible; S: susceptible.
R: 抗病; MR: 中抗; MS: 中感; S: 感病。

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Table 3
表3
表313份小偃麦衍生品系不同环境下的抗病性
Table 3Disease resistance of the 13 wheat lines derived from Thinopyrum across different environments

材料名称 Line	6个环境病小穗率 PDS in six environments (%)							6个环境病小穗率分布 PDS distribution in six environments				染色体数目 Number of chromosomes
	2018 太原 Taiyuan	2018 成都 Chengdu	2018 南京 Nanjing	2017 太原 Taiyuan	2017 南京 Nanjing	2016 南京 Nanjing		最小 Min.	最大 Max.	极差 Range	平均 Average
CH16387	8.7	4.9	4.4	4.5	1.3	1.3		1.3	8.7	7.4	4.2**a	42
CH16371	15.1	7.3	4.6	4.6	2.0	3.0		2.0	15.1	13.1	5.0** b	42
CH16379	7.0	10.2	7.2	5.1	2.0	4.2		2.0	10.2	8.2	6.0* b	42
CH16373	10.5	4.2	5.1	4.8	10.0	4.0		4.0	10.5	6.5	6.7	42
CH16352	7.3	11.6	5.5	4.2	-	6.0		4.2	11.6	7.3	6.7	42
CH16388	6.5	7.8	14.5	4.0	4.6	2.0		2.0	14.5	12.5	6.7	42
CH16419	6.5	11.4	6.6	4.8	-	4.0		4.0	11.4	7.4	6.7	42
CH16367	12.5	9.2	9.3	4.6	2.8	3.0		2.8	12.5	9.7	7.0	42
CH16378	11.0	5.1	-	4.9	-	-		4.9	11.0	6.1	7.1	42
CH16374	9.4	9.8	8.3	5.4	9.2	3.0		3.0	9.8	6.8	7.5	42
CH16375	12.4	10.4	7.1	5.3	8.6	2.0		2.0	12.4	10.4	7.6	42
CH16427	8.4	10.5	-	5.9	-	6.7		5.9	10.5	4.6	7.7	42
CH16432	5.8	12.1	-	12.7	-	4.6		4.6	12.7	8.2	8.4	42
苏麦3号Sumai 3	16.8	3.6	7.9	5.1	4.0	5.1		3.6	16.8	13.2	7.1	42
望水白Wangshuibai	13.9	6.1	12.3	9.6	7.5	6.8		6.1	13.9	7.8	9.4	42
周麦27 Zhoumai 27	64.3	57.0	63.0	58.6	38.6	53.9		38.6	64.3	25.7	55.9	42
Alondra’s	53.1	65.5	63.8	51.7	48.1	61.4		48.1	65.5	17.4	57.3	42

^*, ^**: significantly different from the resistant control at P < 0.05, P < 0.01, respectively. ^a: significantly different from Sumai 3 at P < 0.05, and from Wangshuibai at P < 0.01; ^b: significantly different from Wangshuibai at P < 0.01, but not from Sumai 3. R: resistant; MR: moderately resistant; MS: moderately susceptible; S: susceptible; -: no results or missing data. PDS: percentage of diseased spikelet.
^*和^**分别表示在P < 0.05, P < 0.01水平上与抗病对照有显著差异。^a: 与抗病对照苏麦3号有显著差异(P < 0.05),与望水白极显著差异(P < 0.01); ^b: 仅与抗病对照望水白在P < 0.01水平上差异显著, 与苏麦3号无显著差异。R: 抗病; MR: 中抗; MS: 中感; S: 感病; -: 未调查或数据缺失。

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2.3 13份优良品系的抗赤霉病分析

来源于部分双二倍体TAP8430和TAI8045的5个组合中, 分别有4份和9份材料在6个环境下均表现出优良的赤霉病抗性, 平均病小穗率均小于10.0%, 病害严重度均为1级, 均表现抗病(R)(表2、表3和图2)。t测验表明, CH16387的病小穗率显著低于苏麦3号和望水白(P<0.05), CH16371和CH16379的病小穗率分别显著低于望水白(P<0.05、P<0.01), 其余10个品系的病小穗率与苏麦3号和望水白无显著差异(表3)。

图2

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图2抗病材料和感病材料对赤霉病的反应类型(山西太原, 2018)

a: 13个抗病反应为R的材料; b: 抗病反应为S的部分材料。
Fig. 2Responses to Fusarium head blight in 13 wheat lines (Taiyuan, Shanxi, 2018)

a: resistant wheat lines; b: susceptible wheat lines.


从不同环境病小穗率的变化幅度来看, 除CH16371和CH16388的极差分别为13.1%和12.5%外, 其他11份材料的极差均低于苏麦3号的13.2%, 在试验的各个年度及地点间表现较为稳定的抗病性。尤其是CH16387, 在6个点的试验中, 病小穗率最高仅为8.7%, 最低为1.3%, 极差为7.4%, 低于望水白的7.8%。此外, 不同环境间相关分析表明2018年成都与2018年太原、2017年太原与2017年和2016年南京、2016年与2017年南京之间分别存在一定的相关性, 但相关性均较弱(r<0.5), 其余环境间并无显著相关性(表4)。其原因可能是赤霉病发病易受温、湿度等气候因素影响, 病小穗率在不同环境间变异较大, 但在相同环境下与抗、感病对照相比, 仍具有显著抗病性, 品种抗病性鉴定结果相对稳定, 筛选的抗源材料可用于抗赤霉病的遗传和育种研究。

Table 4
表4
表413个抗病品系在不同环境条件下赤霉病病小穗率的相关系数
Table 4Correction coefficients of percentage of diseased spikes in 13 lines resistant to Furarium head blight under different environments

环境 Environment	2018太原 2018 Taiyuan	2018成都 2018 Chengdu	2018南京 2018 Nanjing	2017太原 2017 Taiyuan	2017南京 2017 Nanjing
2018成都 2018 Chengdu	-0.2764*
2018南京 2018 Nanjing	0.0562	-0.0724
2017太原 2017 Taiyuan	0.0398	0.0247	-0.0686
2017南京 2017 Nanjing	0.0818	-0.0851	-0.0264	0.4086**
2016南京 2016 Nanjing	0.2281	0.0080	-0.0756	0.3199*	0.3141*

^*,^**: significant difference at P < 0.05 and P < 0.01, respectively.
^*和^**分别表示在P < 0.05和P < 0.01水平上差异显著。

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3 讨论

近年来, 小麦赤霉病呈爆发速度快、流行范围广、发生面积大和危害损失重的严重态势, 直接威胁着小麦的安全生产^[2]。抗病种质资源筛选是抗病育种的基础, 许多小麦近缘种对赤霉病具有较高抗病性, 是小麦抗赤霉病基因的重要来源。从这些近缘种中发掘抗病基因, 对于提高小麦抗赤霉病能力具有重要意义^[17]。20世纪80年代以来, 一些研究利用近缘种属创造小麦抗赤霉病新材料, 将纤毛鹅观草^[18]、大赖草^[19]、鹅观草^[20]、华山新麦草^[21]、黑麦草、长穗偃麦草^[22]、簇毛麦等物种的抗赤霉病基因通过外源染色体附加、代换、易位和双二倍体的方式导入小麦背景中, 育成了许多含有外源染色体或染色体片段的抗赤霉病种质^[23]。

偃麦草属植物对赤霉病具有很强的抗性^[24], 通过远缘杂交将其抗病基因导入普通小麦, 是改良小麦赤霉病抗性的重要途径。经过研究已从中间偃麦草和长穗偃麦草中鉴定出多个抗赤霉病双二倍体^[25,26,27]、附加系^[28]、代换系^[27]以及易位系^{[22, 29-30]}。其中易位系在导入抗赤霉病基因的同时, 尽量地减少了不利基因的影响, 而且能够保证小麦染色体构成的稳定性, 因而是利用外源基因的最佳材料。本研究通过多个环境鉴定筛选出的13份高抗赤霉病材料, 有9份来源于小麦-中间偃麦草部分双二倍体TAI8045与普通小麦的杂交组合, 4份来源于小麦-十倍体长穗偃麦草部分双二倍体TAP8430与普通小麦的杂交组合(表2), 其普通小麦亲本对赤霉病都表现中感或感病(表1和图1), 而八倍体亲本TAI8045和TAP8430则高抗赤霉病。因此推测这些材料的赤霉病抗性可能来源于部分双二倍体TAI8045和TAP8430。利用细胞学和分子标记技术可以进一步确定这些材料的赤霉病抗性是否来自偃麦草。根据多年的田间表现, 这些材料的穗型(图2)、株高、株型、籽粒、结实性等农艺性状以及染色体数目都与普通小麦完全一致, 根据系谱及抗病性分析, 推测它们不含或仅含很小的外源片段, 可直接用于小麦抗病育种和QTL/基因定位。

小麦赤霉病是由镰孢属真菌侵染引起的一种土传真菌病害, 引起小麦赤霉病的镰孢菌至少有20种, 发病机制复杂, 不同菌株间的致病力存在显著差异^[31-32], 因此致病菌株的选择是否恰当直接影响着抗病鉴定结果的准确性。本研究使用赤霉菌菌株F0609进行接种鉴定, 经过抗感对照的发病情况对比, 发现苏麦3号和望水白对F0609菌株均表现抗病, 周麦27和Alondra’s则表现感病(表2和图2)。在3年6个环境的接种鉴定中, F0609对周麦27和Alondra’s的致病性表现一致。在鉴定的119份材料中, 表现抗或中抗的品系占总数的62.2%, 感病品系相对较少, 其原因可能是这些材料是经过2015年赤霉病高发期的田间抗病鉴定筛选出的抗病品系, 但由于田间自然发病不均匀, 因而仍有45份材料人工接种鉴定时表现中感或感病(MS或S)。

本研究筛选出的13份抗病品系在测试的6个环境中, 病小穗率变化幅度不大, 在不同环境条件下表现出较好的抗病性。但部分材料在不同年度和地点间表现出较大的差异, 如CH16376的最高病小穗率为42.7%, 最低仅为3.0%, CH16379最高为50.0%, 最低3.0% (附表1)。究其原因, 既有可能是人为接种造成的差异, 也有可能是赤霉病发病易受环境影响所致^[33,34]。因此, 要获得具有良好、稳定抗性的抗赤霉病材料, 需要连续多年在不同环境下进行抗病性鉴定^[35], 并且设置适宜的抗感病对照来评价材料的抗性, 以保证获得的抗病材料具有可靠抗性。13份抗病品系中CH16387、CH16371和CH16379的抗性显著优于苏麦3号或望水白, 其余10份与抗病对照苏麦3号和望水白的抗性水平相当; CH16352、CH16388、CH16427和CH16432则表现出矮秆、早熟、结实性好、籽粒饱满、株型紧凑等优良农艺性状, 可为小麦抗病遗传育种提供有价值的亲本材料。

4 结论

采用单花滴注法对来源于中间偃麦草和长穗偃麦草的119份小偃麦衍生品系进行多年多点表型鉴定, 发现有13份材料在多个环境下与高抗对照品种苏麦3号和望水白抗性水平相当, 在不同环境条件下均表现出较好的抗病性; 61份材料表现中抗, 41份材料表现中感, 4份材料表现高感。这些抗赤霉病材料为小麦抗病遗传育种提供有效抗源。

致谢:

对南京农业大学作物遗传与种质创新国家重点实验室王秀娥博士、刘玉博士, 四川农业大学罗培高博士、黄强兰博士, 四川省农业科学院作物所杨恩年博士在种质抗病鉴定方面给予的帮助; 电子科技大学杨足君博士、李光蓉博士, 江苏里下河地区农业科学研究所臧淑江老师在接种与鉴定方法给予的宝贵指导与帮助, 谨致谢忱。

参考文献 View Option 原文顺序
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Fusarium head blight (FHB) is a devastating disease of wheat and barley worldwide. Resistant cultivars could reduce damage from FHB. Chinese wheat cultivar Sumai 3 and its derivatives represent the greatest degree of resistance to FHB known. A major quantitative trait locus (QTL) on chromosome 3BS and other minor QTL for FHB resistance have been identified in these cultivars and used in wheat-breeding programs worldwide. Many breeding lines with the 3BS resistance QTL and improved agronomic traits have been developed. In barley, only limited sources of FHB resistance are available, especially in six-rowed barley, and none of them contains a DON level low enough to meet the safety requirement of the brewing industry. Several QTL have been identified for lower FHB severity, DON content, and kernel discoloration and used to enhance FHB resistance in barley. Marker-assisted selection for FHB resistance QTL on 3BS of wheat and on 2H of barley is in progress.

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[5] Gilbert J, Haber S . Overview of some recent research developments inFusarium head blight of wheat
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This article reviews the recent progress of research on fusarium head blight (FHB) of wheat. It addresses the broad areas of strategies for disease management, biological control, the pathogen (Fusarium graminearum (teleomorph Gibberella zeae)), mycotoxins, the effects of dwarfing genes on FHB severity, quantitative trait loci (QTLs) and new perspectives. Where there are recent reviews on this subject, we have deliberately examined the subsequent literature to provide an update on research. With few resistant cultivars available even now, the main tools to manage the disease remain rotation, varietal selection, disease forecasting and fungicides. A few biocontrol organisms are being considered for commercial application. The pathogen's sexual life cycle has been investigated in depth, and with its complete genome sequence known, the pathways and genes controlling the sexual development and ascospore release of F. graminearum are being explored. The 3-acetyldeoxynivalenol chemotype of F. graminearum has increased in prevalence in Canada with attendant risks of higher DON levels in cereal grain. Stringent limits on allowable levels of Fusarium mycotoxins in the food/feed chain have been enacted in Europe and the USA, but regulations for Canada are only at the discussion stage with the Canadian Food Inspection Agency. Efforts to develop FHB-resistant lines proceed apace, as these can be selected in most wheat populations despite the adverse effects of dwarfing genes on FHB severity. While more quantitative trait loci (QTLs) for disease resistance continue to be identified and mapped, new resistant cultivars remain disappointingly few. We present some encouraging early results from an alternative approach based on epigenetics.

[6] 张爱民, 阳文龙, 李欣, 孙家柱 . 小麦抗赤霉病研究现状与展望
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A major fusarium head blight (FHB) resistance gene Fhb1 (syn. Qfhs.ndsu-3BS) was fine mapped on the distal segment of chromosome 3BS of spring wheat (Triticum aestivum L.) as a Mendelian factor. FHB resistant parents, Sumai 3 and Nyubai, were used as sources of this gene. Two mapping populations were developed to facilitate segregation of Qfhs.ndsu-3BS in either a fixed resistant (Sumai 3*5/Thatcher) (S/T) or fixed susceptible (HC374/3*98B69-L47) (HC/98) genetic background (HC374=Wuhan1/Nyubai) for Type II resistance. Type II resistance (disease spread within the spike) was phenotyped in the greenhouse using single floret injections with a mixture of macro-conidia of three virulent strains of Fusarium graminearum. Due to the limited heterogeneity in the genetic background of the crosses and based on the spread of infection, fixed recombinants in the interval between molecular markers XGWM533 and XGWM493 on 3BS could be assigned to discrete “resistant” and “susceptible” classes. The phenotypic distribution was bimodal with progeny clearly resembling either the resistant or susceptible parent. Marker order for the two maps was identical with the exception of marker STS-3BS 142, which was not polymorphic in the HC/98 population. The major gene Fhb1 was successfully fine mapped on chromosome 3BS in the same location in the two populations within a 1.27-cM interval (S/T) and a 6.05-cM interval (HC/98). Fine mapping of Fhb1 in wheat provides tightly linked markers that can reduce linkage drag associated with marker-assisted selection of Fhb1 and assist in the isolation, sequencing and functional identification of the underlying resistance gene.

[8] Cuthbert P A, Somers D J, Brule-Babel A . Mapping of Fhb2 on chromosome 6BS: a gene controlling Fusarium head blight field resistance in bread wheat(Triticum aestivum L.)
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Fusarium head blight (FHB) is one of the most important fungal wheat diseases worldwide. Understanding the genetics of FHB resistance is key to facilitate the introgression of different FHB resistance genes into adapted wheat. The objective of this project was to study the FHB resistance QTL on chromosome 6B, quantify the phenotypic variation, and qualitatively map the resistance gene as a Mendelian factor. The FHB resistant parent BW278 (AC Domain*2/Sumai 3) was used as the source of the resistance allele. A large recombinant inbred line (RIL) mapping population was developed from the cross BW278/AC Foremost. The population segregated for three known FHB resistance QTL located on chromosomes 3BSc, 5A, and 6B. Molecular markers on chromosome 6B (WMC104, WMC397, GWM219), 5A (GWM154, GWM304, WMC415), and 3BS (WMC78, GWM566, WMC527) were amplified on approximately 1,440 F_2:7 RILs. The marker information was used to select 89 RILs that were fixed homozygous susceptible for the 3BSc and 5A FHB QTLs and were recombinant in the 6B interval. Disease response was evaluated on 89 RILs and parental checks in the greenhouse and field nurseries. Dual floret injection (DFI) was used in greenhouse trials to evaluate disease severity (DS). Macroconidial spray inoculations were used in field nurseries conducted at two locations in southern Manitoba (Carman and Glenlea) over two years 2003 and 2004, to evaluate disease incidence, disease severity, visual rating index, and Fusarium-damaged kernels. The phenotypic distribution for all five-disease infection measurements was bimodal, with lines resembling either the resistant or susceptible checks and parents. All of the four field traits for FHB resistance mapped qualitatively to a coincident position on chromosome 6BS, flanked by GWM133 and GWM644, and is named Fhb2. The greenhouse-DS trait mapped 2cM distal to Fhb2. Qualitative mapping of Fhb2 in wheat provides tightly linked markers that can reduce linkage drag associated with marker assisted selection of Fhb2 and aid the pyramiding of different resistance loci for wheat improvement.

[9] Xue S L, Li G Q, Jia H Y, Xu F, Lin F, Tang M Z, Wang Y, An X, Xu H B, Zhang L X, Kong Z X, Ma Z Q . Fine mapping Fhb4, a major QTLs conditioning resistance to Fusarium infection in bread wheat(Triticum aestivum L.)
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DOI:10.1007/s00122-010-1298-5URL [本文引用: 1]
Qfhi.nau-4B is a major quantitative trait locus (QTL) against Fusarium graminearum infection identified in the Fusarium head blight-resistant germplasm Wangshuibai. To fine map this QTL, a recombinant inbred line (RIL) population of 530 lines derived from Nanda2419×Wangshuibai and the BC₃F₂ population derived from the cross of a Qfhi.nau-4B near isogenic line (NIL) with susceptible cultivar Mianyang 99-323 as the recurrent parent were screened for recombinants occurred between microsatellite markers Xbarc20 and Xwmc349 that flank Qfhi.nau-4B. A total of 95 recombinants were obtained, including 45 RIL recombinants obtained through reverse-selection of Qfhi.nau-5A and 50 NIL recombinants from the BC₃F₂ population. Genotyping these recombinant lines with 22 markers mapping to the Xbarc20 and Xwmc349 interval revealed fourteen genotypes of the RIL recombinants as well as of the NIL recombinants. Two-year field evaluation of their resistance to Fusarium infection showed that these lines could be clearly classified into two groups according to percentage of infected spikes. The more resistant class had over 60% less infection than the susceptible class and were common to have Wangshuibai chromatin in the 1.7-cM interval flanked by Xhbg226 and Xgwm149. None of the susceptible recombinants had this Wangshuibai chromatin. Qfhi.nau-4B was thus confined between Xhbg226 and Xgwm149 and named Fhb4. The interval harboring Fhb4 was mapped to 4BL5-0.86–1.00 bin using Chinese Spring deletion lines, a region with about 5.7 times higher recombination rate than the genome average. This study established the basis for map-based cloning of Fhb4.

[10] Xue S L, Xu F, Tang M Z, Zhou Y, Li G Q, An X, Lin F, Xu H B, Jia H Y, Zhang L X, Kong Z X, Ma Z Q . Precise mappingFhb5, a major QTLs conditioning resistance to Fusarium infection in bread wheat(Triticum aestivum L.)
Theor Appl Genet, 123:1055-1063.
DOI:10.1007/s00122-011-1647-zURLPMID:21739138 [本文引用: 1]
Qfhi.nau-5A is a major quantitative trait locus (QTL) against Fusarium graminearum infection in the resistant wheat germplasm Wangshuibai. Genetic analysis using BC(3)F(2) and BC(4)F(2) populations, derived from selfing two near-isogenic lines (NIL) heterozygous at Qfhi.nau-5A that were developed, respectively, with Mianyang 99-323 and PH691 as the recurrent parent, showed that Qfhi.nau-5A inherited like a single dominant gene. This QTL was thus designated as Fhb5. To fine map it, these two backcross populations and a recombinant inbred line (RIL) population derived from Nanda2419?×?Wangshuibai were screened for recombinants occurring between its two flanking markers Xbarc56 and Xbarc100. Nineteen NIL recombinants were identified from the two backcross populations and nine from the RIL population. In the RIL recombinant selection process, selection against Fhb4 present in the RIL population was incorporated. Genotyping these recombinant lines with ten markers mapping to the Xbarc56-Xbarc100 interval revealed four types of Mianyang 99-323-derived NIL recombinants, three types of PH691-derived NIL recombinants, and four types of RIL recombinants. In different field trials, the percentage of infected spikes of these lines displayed a distinct two-peak distribution. The more resistant class had over 55% less infection than the susceptible class. Common to these resistant genotypes, the 0.3-cM interval flanked by Xgwm304 and Xgwm415 or one of these two loci was derived from Wangshuibai, while none of the susceptible recombinants had Wangshuibai chromatin in this interval. This interval harboring Fhb5 was mapped to the pericentromeric C-5AS3-0.75 bin through deletion bin mapping. The precise localization of Fhb5 will facilitate its utilization in marker-assisted wheat breeding programs.

[11] Qi L L, Pumphrey M O, Friebe B, Chen P D, Gill B S . Molecular cytogenetic characterization of alien introgressions with geneFhb3 for resistance to Fusarium head blight disease of wheat
Theor Appl Genet, 2008,117:1155-1166.
DOI:10.1007/s00122-008-0853-9URL [本文引用: 1]
Fusarium head blight (FHB) resistance was identified in the alien species Leymus racemosus, and wheat-Leymus introgression lines with FHB resistance were reported previously. Detailed molecular cytogenetic analysis of alien introgressions T01, T09, and T14 and the mapping of Fhb3, a new gene for FHB resistance, are reported here. The introgression line T09 had an unknown wheat-Leymus translocation chromosome. A total of 36 RFLP markers selected from the seven homoeologous groups of wheat were used to characterize T09 and determine the homoeologous relationship of the introgressed Leymus chromosome with wheat. Only short arm markers for group 7 detected Leymus-specific fragments in T09, whereas 7AS-specific RFLP fragments were missing. C-banding and genomic in situ hybridization results indicated that T09 has a compensating Robertsonian translocation T7AL·7Lr#1S involving the long arm of wheat chromosome 7A and the short arm of Leymus chromosome 7Lr#1 substituting for chromosome arm 7AS of wheat. Introgression lines T01 (2n=44) and T14 (2n=44) each had two pairs of independent translocation chromosomes. T01 had T4BS·4BL-7Lr#1S+T4BL-7Lr#1S·5Lr#1S. T14 had T6BS·6BL-7Lr#1S+T6BL·5Lr#1S. These translocations were recovered in the progeny of the irradiated line Lr#1 (T5Lr#1S·7Lr#1S). The three translocation lines, T01, T09, and T14, and the disomic addition 7Lr#1 were consistently resistant to FHB in greenhouse point-inoculation experiments, whereas the disomic addition 5Lr#1 was susceptible. The data indicated that at least one novel FHB resistance gene from Leymus, designated Fhb3, resides in the distal region of the short arm of chromosome 7Lr#1, because the resistant translocation lines share a common distal segment of 7Lr#1S. Three PCR-based markers, BE586744-STS, BE404728-STS, and BE586111-STS, specific for 7Lr#1S were developed to expedite marker-assisted selection in breeding programs.

[12] Cainong J C, Bockus W W, Feng Y G, Chen P D, Qi L L, Sehgal S K, Danilova T V, Koo D H, Friebe B, Gill B S . Chromosome engineering, mapping, and transferring of resistance toFusarium head blight disease from Elymus tsukushiensis into wheat
Theor Appl Genet, 2015,128:1019-1027.
DOI:10.1007/s00122-015-2485-1URLPMID:25726000 [本文引用: 1]
This manuscript describes the transfer and molecular cytogenetic characterization of a novel source of Fusarium head blight resistance in wheat. Fusarium head blight (FHB) caused by the fungus Fusarium graminearum Schwabe [telomorph = Gibberella zeae (Schwein. Fr.) Petch] is an important disease of bread wheat, Triticum aestivum L. (2n = 6x = 42, AABBDD) worldwide. Wheat has limited resistance to FHB controlled by many loci and new sources of resistance are urgently needed. The perennial grass Elymus tsukushiensis thrives in the warm and humid regions of China and Japan and is immune to FHB. Here, we report the transfer and mapping of a major gene Fhb6 from E. tsukushiensis to wheat. Fhb6 was mapped to the subterminal region in the short arm of chromosome 1E(ts)#1S of E. tsukushiensis. Chromosome engineering was used to replace corresponding homoeologous region of chromosome 1AS of wheat with the Fhb6 associated chromatin derived from 1E(ts)#1S of E. tsukushiensis. Fhb6 appears to be new locus for wheat as previous studies have not detected any FHB resistance QTL in this chromosome region. Plant progenies homozygous for Fhb6 had a disease severity rating of 7 % compared to 35 % for the null progenies. Fhb6 has been tagged with molecular markers for marker-assisted breeding and pyramiding of resistance loci for effective control of FHB.

[13] Guo J, Zhang X L, Hou Y L, Cai J J, Shen X R, Zhou T T, Xu H H, Ohm H W, Wang H W, Li A F, Han F P, Wang H G, Kong L R . High-density mapping of the major FHB resistance geneFhb7 derived from Thinopyrum ponticum and its pyramiding with Fhb1 by marker-assisted selection
Theor Appl Genet, 2015,128:2301-2316.
DOI:10.1007/s00122-015-2586-xURLPMID:26220223 [本文引用: 1]
Wheat lines with shortened Th. ponticum chromatin carrying Fhb7 and molecular markers linked to Fhb7 will accelerate the transfer of Fhb7 to breeding lines and provide an important resource for future map-based cloning of this gene. Fusarium head blight is a major wheat disease globally. A major FHB resistance gene, designated as Fhb7, derived from Thinopyrum ponticum, was earlier transferred to common wheat, but was not used in wheat breeding due to linkage drag. The aims of this study were to (1) saturate this FHB resistance gene region; (2) develop and characterize secondary translocation lines with shortened Thinopyrum segments carrying Fhb7 using ph1b; (3) pyramid Fhb7 and Fhb1 by marker-assisted selection. Fhb7 was mapped in a 1.7 cM interval that was flanked by molecular markers XsdauK66 and Xcfa2240 with SSR, diversity arrays technology, EST-derived and conserved markers. KS24-2 carrying Fhb7 was analyzed with molecular markers and genomic in situ hybridization, confirming it was a 7DS.7el2L Robertsonian translocation. To reduce the Thinopyrum chromatin segments carrying Fhb7, a BC1F2 population (Chinese Spring ph1bph1b*2/KS24-2) was developed and genotyped with the markers linked to Fhb7. Two new translocation lines (SDAU1881 and SDAU1886) carrying Fhb7 on shortened alien segments (approximately 16.1 and 17.3% of the translocation chromosome, respectively) were developed. Furthermore, four wheat lines (SDAU1902, SDAU1903, SDAU1904, and SDAU1906) with the pyramided markers flanking Fhb1 and Fhb7 were developed and the FHB responses indicated lines with mean NDS ranging from 1.3 to 1.6 had successfully combined Fhb7 and Fhb1. Three new molecular markers associated with Fhb7 were identified and validated in 35 common wheat varieties. The translocation lines with shortened alien segments carrying Fhb7 (and Fhb1) and the markers closely linked to Fhb7 will be useful for improving wheat scab resistance.

[14] Bai G H, Kolb F L, Shaner G, Domier L L . Amplified fragment length polymorphism markers linked to a major quantitative trait locus controlling scab resistance in wheat
Phytopathology, 1999,89:343-348.
DOI:10.1094/PHYTO.1999.89.4.343URLPMID:18944781 [本文引用: 1]
Scab is a destructive disease of wheat. To accelerate development of scab-resistant wheat cultivars, molecular markers linked to scab resistance genes have been identified by using recombinant inbred lines (RILs) derived by single-seed descent from a cross between the resistant wheat cultivar Ning 7840 (resistant to spread of scab within the spike) and the susceptible cultivar Clark. In the greenhouse, F(5), F(6), F(7), and F(10) families were evaluated for resistance to spread of scab within a spike by injecting about 1,000 conidiospores of Fusarium graminearum into a central spikelet. Inoculated plants were kept in moist chambers for 3 days to promote initial infection and then transferred to greenhouse benches. Scab symptoms were evaluated four times (3, 9, 15, and 21 days after inoculation). The frequency distribution of scab severity indicated that resistance to spread of scab within a spike was controlled by a few major genes. DNA was isolated from both parents and F(9) plants of the 133 RILs. A total of 300 combinations of amplified fragment length polymorphism (AFLP) primers were screened for polymorphisms using bulked segregant analysis. Twenty pairs of primers revealed at least one polymorphic band between the two contrasting bulks. The segregation of each of these bands was evaluated in the 133 RILs. Eleven AFLP markers showed significant association with scab resistance, and an individual marker explained up to 53% of the total variation (R(2)). The markers with high R(2) values mapped to a single linkage group. By interval analysis, one major quantitative trait locus for scab resistance explaining up to 60% of the genetic variation for scab resistance was identified. Some of the AFLP markers may be useful in marker-assisted breeding to improve resistance to scab in wheat.

[15] 中华人民共和国农业行业标准NY/T 1443.4-2007, 小麦病虫性评价技术规范, 第4部分: 小麦抗赤霉病评价技术规范

[本文引用: 1]

Agricultural Standard of the People’s Republic of China, NY/T 1443.4-2007. Rules for Resistance Evaluation of Wheat to Diseases and Insect Pests, Part 4. Rule for Resistance Evaluation to Wheat Scab
(in Chinese).
[本文引用: 1]

[16] Han F P, Fedak G, Benabdelmouna A, Armstrong K C, Ouellet T . Characterization of six wheat ×Thinopyrum intermedium derivativesby GISH, RFLP and multicolor GISH
Genome, 2003,46:490-495.
DOI:10.1139/g03-032URLPMID:12834067 [本文引用: 1]
Restriction fragment length polymorphism (RFLP) analysis and multicolor genomic in situ hybridization (GISH) are useful tools to precisely characterize genetic stocks derived from crosses of wheat (Triticum aestivum) with Thinopyrum intermedium and Thinopyrum elongatum. The wheat x Th. intermedium derived stocks designated Z1, Z2, Z3, Z4, Z5, and Z6 were initially screened by multicolor GISH using Aegilops speltoides genomic DNA for blocking and various combinations of genomic DNA from Th. intermedium, Triticum urartu, and Aegilops tauschii for probes. The probing (GISH) results indicated that lines Z1 and Z3 were alien disomic addition lines with chromosome numbers of 2n = 44. Z2 was a substitution line in which chromosome 2D was substituted by a pair of Th. intermedium chromosomes; this was confirmed by RFLP and muticolour GISH. Z4 (2n = 44) contained two pairs of wheat--Th. intermedium translocated chromosomes; one pair involved A-genome chromosomes, the other involved D- and A- genome chromosomes. Z5 (2n = 44) contained one pair of wheat--Th. intermedium translocated chromosomes involving the D- and A-genome chromosomes of wheat. Z6 (2n = 44) contained one pair of chromosomes derived from Th. intermedium plus another pair of translocated chromosomes involving B-genome chromosomes of wheat Line Z2 was of special interest because it has some resistance to infection by Fusarium graminearum.

[17] Zhang L, Luo P G, Ren Z L, Zhang H Y . Controlling fusarium head blight of wheat (Triticum aestivum L.) with genetics
Adv Biosci Biotech, 2011,2:263-270.
DOI:10.3835/plantgenome2015.04.0028URLPMID:27898754 [本文引用: 1]
Fusarium head blight (FHB) is one of the most important wheat ( L.) diseases worldwide, and host resistance displays complex genetic control. A genome-wide association study (GWAS) was performed on 273 winter wheat breeding lines from the midwestern and eastern regions of the United States to identify chromosomal regions associated with FHB resistance. Genotyping-by-sequencing (GBS) was used to identify 19,992 single-nucleotide polymorphisms (SNPs) covering all 21 wheat chromosomes. Marker-trait associations were performed with different statistical models, the most appropriate being a compressed mixed linear model (cMLM) controlling for relatedness and population structure. Ten significant SNP-trait associations were detected on chromosomes 4A, 6A, 7A, 1D, 4D, and 7D, and multiple SNPs were associated with on chromosome 3B. Although combination of favorable alleles of these SNPs resulted in lower levels of severity (SEV), incidence (INC), and deoxynivalenol concentration (DON), lines carrying multiple beneficial alleles were in very low frequency for most traits. These SNPs can now be used for creating new breeding lines with different combinations of favorable alleles. This is one of the first GWAS using genomic resources from the International Wheat Genome Sequencing Consortium (IWGSC).

[18] Wang X E, Chen P D, Liu D J, Zhang P, Zhou B, Friebe B, Gill B S . Molecular cytogenetic characterization ofRoegneria ciliaris chromosome additions in common wheat
Theor Appl Genet, 2001,102:651-657.
DOI:10.1007/s001220051693URL [本文引用: 1]
The development of alien addition lines is important both for transferring useful genes from related species into common wheat and for studying the relationship between alien chromosomes and those of wheat. Roegneria ciliaris (2n=4x=28, S^cS^cY^cY^c) is reported to be a potential source of resistance to wheat scab, which may be useful in wheat improvement. The amphiploid common wheat-R. ciliaris and BC₁F₇ or BC₂F₆ derivatives were screened by C-banding, genomic in situ hybridization (GISH), fluorescent in situ hybridization (FISH) and restriction fragment length polymorphism (RFLP) for the presence of R. ciliaris chromatin introgressed into wheat. Six lines were identified as disomic chromosome additions (DA), one as a ditelosomic addition (Dt), two as double disomic additions (dDA) and one as a monosomic chromosome addition (MA). RFLP analysis using wheat homoeologous group-specific clones indicated that the R. ciliaris chromosomes involved in these lines belong to groups 1, 2, 3, 5 and 7. The genomic affinities of the added R. ciliaris chromosomes were determined by FISH analysis using the repetitive sequence pCbTaq4.14 as a probe. These data suggest that the R. ciliaris chromosomes in five lines belong to the S^c genome. Based on the molecular cytogenetic data, the lines are designated as DA2S^c#1, Dt2S^c#1L, DA3S^c#1, dDA1S^c#2+5Y^c#1, DA5Y^c#1, DA7S^c#1, DA7Y^c#1 and MA?Y^c#1. Based on the present and previous work, 8 of the 14 chromosomes of R. ciliaris have been transferred into wheat.

[19] 王林生, 张雅莉, 南广慧 . 普通小麦-大赖草易位系T5AS-7Lr.7LrS分子细胞遗传学鉴定
作物学报, 2018,44:1442-1447.
DOI:10.3724/SP.J.1006.2018.01442URL [本文引用: 1]
大赖草对赤霉病具有较好的抗性, 将大赖草赤霉病抗性基因转入普通小麦, 对拓宽小麦赤霉病抗性基础有重要意义。本研究在获得抗赤霉病普通小麦-大赖草异附加系基础上, 采用 ⁶⁰Co-γ射线(1200 Rad, 剂量率100 Rad min ^-1)处理小麦-大赖草二体附加系DA7Lr, 并用处理后的花粉授给去雄的普通小麦中国春, 对其M₁代种子根尖细胞有丝分裂中期染色体进行GISH分析, 获得1株具有一条普通小麦-大赖草易位染色体的植株, 对其自交后代中具有2条易位染色体植株的花粉母细胞减数分裂中期I观察发现, 2条易位染色体形成了稳定的环状二价体, 表明该植株为纯合体。利用顺序GISH-双色FISH分析, 结合C-分带、小麦D组专化探针Oligo-pAs1-2和B组专化探针Oligo-pSc119.2-2, 进一步鉴定出该易位系为T5AS-7LrL·7LrS, 同时筛选出可追踪该易位系的3个EST-STS分子标记, 即BE591127、BQ168298和BE591737。该易位系的育成也为小麦赤霉病遗改良提供了新种质。
Wang L S, Zhang Y L, Nan G H . Molecular and cytogenetic identification of Triticum aestivum-Leymus racemosus translocation line T5AS-7Lr.7LrS
Acta Agron Sin, 2018,44:1442-1447 (in Chinese with English abstract).
DOI:10.3724/SP.J.1006.2018.01442URL [本文引用: 1]
大赖草对赤霉病具有较好的抗性, 将大赖草赤霉病抗性基因转入普通小麦, 对拓宽小麦赤霉病抗性基础有重要意义。本研究在获得抗赤霉病普通小麦-大赖草异附加系基础上, 采用 ⁶⁰Co-γ射线(1200 Rad, 剂量率100 Rad min ^-1)处理小麦-大赖草二体附加系DA7Lr, 并用处理后的花粉授给去雄的普通小麦中国春, 对其M₁代种子根尖细胞有丝分裂中期染色体进行GISH分析, 获得1株具有一条普通小麦-大赖草易位染色体的植株, 对其自交后代中具有2条易位染色体植株的花粉母细胞减数分裂中期I观察发现, 2条易位染色体形成了稳定的环状二价体, 表明该植株为纯合体。利用顺序GISH-双色FISH分析, 结合C-分带、小麦D组专化探针Oligo-pAs1-2和B组专化探针Oligo-pSc119.2-2, 进一步鉴定出该易位系为T5AS-7LrL·7LrS, 同时筛选出可追踪该易位系的3个EST-STS分子标记, 即BE591127、BQ168298和BE591737。该易位系的育成也为小麦赤霉病遗改良提供了新种质。

[20] 丁春邦, 周永红 . 小麦族拟鹅观草属研究进展
四川农业大学学报, 2004,22:269-273.
[本文引用: 1]

Ding C B, Zhou Y H . Advances in research on Pseudoroegneria in the tribe Triticeae
J Sichuan Agric Univ, 2004,22:269-273 (in Chinese with English abstract).
[本文引用: 1]

[21] 王益, 康厚扬, 原红军, 蒋云, 张海琴, 周永红 . 普通小麦与华山新麦草衍生后代的农艺性状和细胞遗传学研究
四川农业大学学报, 2008,26:405-410.
[本文引用: 1]

Wang Y, Kang H Y, Yuan H J, Jiang Y, Zhang H Q, Zhou Y H . Cytogenetic and morphological studies on the derivative progenies of Triticum aestivum × Psathyrostachys huashanica
J Sichuan Agric Univ, 2008,26:405-410 (in Chinese with English abstract).
[本文引用: 1]

[22] 张璐璐, 陈士强, 李海凤, 刘慧萍, 戴毅, 高勇, 陈建民 . 小麦-长穗偃麦草7E抗赤霉病易位系培育
中国农业科学, 2016,49:3477-3488.
DOI:10.3864/j.issn.0578-1752.2016.18.002URL [本文引用: 2]
【目的】将二倍体长穗偃麦草7E染色体导入主栽小麦背景,培育小麦-长穗偃麦草7E染色体抗赤霉病易位系,为小麦抗赤霉病遗传改良利用外源优良基因提供新种质。【方法】利用中国春-长穗偃麦草7E代换系DS7E（7B）与扬麦16杂交的F₂种子进行⁶⁰Co辐射（30 000 rad）和种植,表现型选择收获存活M₁植株的种子,从M₂连续通过表型农艺性状选择、单花滴注法进行赤霉病抗性鉴定和长穗偃麦草7E染色体或染色体臂特异分子标记PCR扩增筛选,最后在M₄代对中选材料以长穗偃麦草基因组DNA为探针进行基因组原位杂交（genomic in situ hybridization,GISH）证实。【结果】M₁选择了赤霉病发病率不同的13个单株进行繁殖。利用前期开发的长穗偃麦草7E染色体和7EL、7ES特异标记检测13株的后代M₂单株,获得含有长穗偃麦草7EL片段7株和7ES片段14株；对21株M₂衍生的222个M₃植株进行特异标记检测,共选择含有长穗偃麦草7EL片段13株和7ES片段3株；利用来自12株M₃的后代（M₄）进行GISH,共9株M₃的后代具有小麦-长穗偃麦草易位染色体,体细胞染色体2n=42。2株M₃的后代显示附加2条长穗偃麦草染色体短臂,体细胞染色体2n=44。连续多年多途径的筛选,获得4份材料,3份材料均为长穗偃麦草7E染色体长臂易位系,命名为TW-7EL1、TW-7EL2和TW-7EL3。1份为7E染色体短臂附加系,命名为W-DA7ES,最后所获得的材料是源自M₁代2个单株。连续3年赤霉病抗性鉴定结果表明长穗偃麦草7EL易位系抗性高,发病率明显低于中国春和扬麦16,与苏麦3号相当,而7ES附加系的赤霉病抗性明显较低,发病率明显高于7EL易位系。【结论】通过赤霉病抗性鉴定、染色体特异分子标记筛选和GISH证实相结合培育了小麦-长穗偃麦草7EL抗赤霉病易位系,长穗偃麦草易位片段鉴定快速和准确。二倍体长穗偃麦草7E染色体长臂中含有抗小麦赤霉病的基因。
Zhang L L, Chen S Q, Li H F, Liu H P, Dai Y, Gao Y, Chen J M . Development of wheat- Thinopyrum elongatum translocation lines resistant to Fusarium head blight
Sci Agric Sin, 2016,49:3477-3488 (in Chinese with English abstract).
DOI:10.3864/j.issn.0578-1752.2016.18.002URL [本文引用: 2]
【目的】将二倍体长穗偃麦草7E染色体导入主栽小麦背景,培育小麦-长穗偃麦草7E染色体抗赤霉病易位系,为小麦抗赤霉病遗传改良利用外源优良基因提供新种质。【方法】利用中国春-长穗偃麦草7E代换系DS7E（7B）与扬麦16杂交的F₂种子进行⁶⁰Co辐射（30 000 rad）和种植,表现型选择收获存活M₁植株的种子,从M₂连续通过表型农艺性状选择、单花滴注法进行赤霉病抗性鉴定和长穗偃麦草7E染色体或染色体臂特异分子标记PCR扩增筛选,最后在M₄代对中选材料以长穗偃麦草基因组DNA为探针进行基因组原位杂交（genomic in situ hybridization,GISH）证实。【结果】M₁选择了赤霉病发病率不同的13个单株进行繁殖。利用前期开发的长穗偃麦草7E染色体和7EL、7ES特异标记检测13株的后代M₂单株,获得含有长穗偃麦草7EL片段7株和7ES片段14株；对21株M₂衍生的222个M₃植株进行特异标记检测,共选择含有长穗偃麦草7EL片段13株和7ES片段3株；利用来自12株M₃的后代（M₄）进行GISH,共9株M₃的后代具有小麦-长穗偃麦草易位染色体,体细胞染色体2n=42。2株M₃的后代显示附加2条长穗偃麦草染色体短臂,体细胞染色体2n=44。连续多年多途径的筛选,获得4份材料,3份材料均为长穗偃麦草7E染色体长臂易位系,命名为TW-7EL1、TW-7EL2和TW-7EL3。1份为7E染色体短臂附加系,命名为W-DA7ES,最后所获得的材料是源自M₁代2个单株。连续3年赤霉病抗性鉴定结果表明长穗偃麦草7EL易位系抗性高,发病率明显低于中国春和扬麦16,与苏麦3号相当,而7ES附加系的赤霉病抗性明显较低,发病率明显高于7EL易位系。【结论】通过赤霉病抗性鉴定、染色体特异分子标记筛选和GISH证实相结合培育了小麦-长穗偃麦草7EL抗赤霉病易位系,长穗偃麦草易位片段鉴定快速和准确。二倍体长穗偃麦草7E染色体长臂中含有抗小麦赤霉病的基因。

[23] 丛雯雯, 郭长虹 . 小麦近缘野生植物的赤霉病抗源筛选及其利用
分子植物育种, 2010,8:1043-1049.
[本文引用: 1]

Cong W W, Guo C H . Selection and utilization of resistance sources to Fusarium head blight in wheat wild relatives
Mol Plant Breed, 2010,8:1043-1049 (in Chinese with English abstract).
[本文引用: 1]

[24] Li H J, Wang X M . Thinopyrum ponticum and Th. intermedium: the promising source of resistance to fungal and viral diseases of wheat
J Genet Genomics, 2009,36:557-565.
DOI:10.1016/S1673-8527(08)60147-2URL [本文引用: 1]

Abstract

Thinopyrum ponticum and Th. intermedium provide superior resistance against various diseases in wheat (Ttricum aestivum). Because of their readily crossing with wheat, many genes for disease resistance have been introduced from the wheatgrasses into wheat. Genes for resistance to leaf rust, stem rust, powdery mildew, Barley yellow dwarf virus, Wheat streak mosaic virus, and its vector, the wheat curl mite, have been transferred into wheat by producing chromosome translocations. These genes offer an opportunity to improve resistance of wheat to the diseases; some of them have been extensively used in protecting wheat from damage of the diseases. Moreover, new resistance to diseases is continuously detected in the progenies of wheat-Thinopyrum derivatives. The present article summaries characterization and application of the genes for fungal and viral disease-resistance derived from Th. ponticum and Th. intermedium.

[25] Oliver R E, Cai X, Xu S S, Chen X, Stack R W . Wheat-alien species derivatives: a novel source of resistance toFusarium head blight in wheat
Crop Sci, 2005,45:1353-1360.
DOI:10.2135/cropsci2004.0503URL [本文引用: 1]

[26] Zeng J, Cao W, Fedak G, Sun S, Mccallum B, Fetch T, Xue A, Zhou Y . Molecular cytological characterization of two novel durum-Thinopyrum intermedium partial amphiploids with resistance to leaf rust, stem rust and Fusarium head blight
Hereditas, 2013,150:10-16.
DOI:10.1111/j.1601-5223.2012.02262.xURL [本文引用: 1]
Thinopyrum intermedium, a wild relative of wheat, is an excellent source of disease resistance. Two novel partial amphiploids, 08-47-50 and 08-53-55 (2n = 6x = 42), were developed from wide crosses between durum wheat and Th. intermedium. Meiotic analysis showed that pollen mother cells of the two partial amphiploids formed an average 20.49 bivalents for 08-47-50 and 20.67 bivalents for 08-53-55, indicating that they are basically cytologically stable. GISH analysis revealed that the two partial amphiploids carried different chromosome compositions. 08-47-50 had fourteen chromosomes from Th. intermedium and its alien chromosomes included six St-, four Ee- and four Ee-St translocated chromosomes, whereas 08-53-55 had four St- and ten Ee-St translocated chromosomes. Fungal disease evaluation indicated that both partial amphiploids had a high level of resistance to FHB, leaf rust and stem rust race Ug99. These two novel partial amphiploids with multiple disease resistance could be used as a new source of multiple disease resistance in bread wheat and durum wheat breeding programs.

[27] Turner M K, De Haan L R, Jin Y, Anderson J . Wheatgrass-wheat partial amphiploids as a novel source of stem rust andFusarium head blight resistance
Crop Sci, 2013,53:1994-2005.
DOI:10.2135/cropsci2012.10.0584URL [本文引用: 2]
Perennial wheatgrasses (Thinopyrum spp.) are recognized sources of genetic variation for annual wheat (Triticum aestivum L.) improvement. Amphiploid lines made by crossing Thinopyrum spp. and T. aestivum (common wheat) can increase resilience of wheat to pathogens and abiotic stress. However, lack of pairing between chromosomes of Thinopyrum and Triticum species reduces genome stability, seed set, and perenniality. Fifty-two wheat-wheatgrass amphiploids from the perennials Thinopyrum intermedium (Host) Barkworth & D. R. Dewey, Thinopyrum ponticum (Podp.) Barkworth & D. R. Dewey, and Thinopyrum junceum (L.) A. Love crossed with the annuals T. aestivum, Triticum turgidum L. subsp. carthlicum (Nevski) A. Love & D. Love (syn. Triticum carthlicum Nevski), and Triticum turgidum subsp. durum (Desf.) Husn, were screened for wheat stem rust (caused by Puccinia graminis) and Fusarium head blight (FHB) (caused by Fusarium graminearum) reaction and evaluated for winter hardiness and perenniality. Twenty-four of 48 amphiploid lines were resistant to all stem rust races screened, including TTKSK (syn. Ug99), TRTTF, and common U. S. races. Of the 30 amphiploid lines point inoculated with F. graminearum, 21 were resistant based on the percentage of infected spikelets and the percent of visually scabby kernels. Three sources each of potentially novel stem rust and uncharacterized FHB resistance were identified and may be useful for wheat improvement. Two lines showed perenniality in Minnesota and may be valuable as cold-tolerant perennial wheat germplasm. Seven lines representing two families showed potential genetic stability based on chromosome counts and seed production.

[28] Fu S, Lv Z, Qi B, Guo X, Li J, Liu B, Han F . Molecular cytogenetic characterization of wheat-Thinopyrum elongatum addition, substitution and translocation lines with a novel source of resistance to wheat Fusariumhead blight
J Genet Genomics, 2013,39:103-110.
DOI:10.1016/j.jgg.2011.11.008URLPMID:22361509 [本文引用: 1]
Thinopyrum elongatum (2n=2x=14, EE), a wild relative of wheat, has been suggested as a potentially novel source of resistance to several major wheat diseases including Fusarium Head Blight (FHB). In this study, a series of wheat (cv. Chinese Spring, CS) substitution and ditelosomic lines, including Th. elongatum additions, were assessed for Type II resistance to FHB. Results indicated that the lines containing chromosome 7E of Th. elongatum gave a high level of resistance to FHB, wherein the infection did not spread beyond the inoculated floret. Furthermore, it was determined that the novel resistance gene(s) of 7E was located on the short-arm (7ES) based on sharp difference in FHB resistance between the two 7E ditelosomic lines for each arm. On the other hand, Th. elongatum chromosomes 5E and 6E likely contain gene(s) for susceptibility to FHB because the disease spreads rapidly within the inoculated spikes of these lines. Genomic in situ hybridization (GISH) analysis revealed that the alien chromosomes in the addition and substitution lines were intact, and the lines did not contain discernible genomic aberrations. GISH and multicolor-GISH analyses were further performed on three translocation lines that also showed high levels of resistance to FHB. Lines TA3499 and TA3695 were shown to contain one pair of wheat-Th. elongatum translocated chromosomes involving fragments of 7D plus a segment of the 7E, while line TA3493 was found to contain one pair of wheat-Th. elongatum translocated chromosomes involving the D- and A-genome chromosomes of wheat. Thus, this study has established that the short-arm of chromosome 7E of Th. elongatum harbors gene(s) highly resistant to the spreading of FHB, and chromatin of 7E introgressed into wheat chromosomes largely retained the resistance, implicating the feasibility of using these lines as novel material for breeding FHB-resistant wheat cultivars.

[29] Liu Z H, Xu M, Xian Z P, Li X, Chen W Q, Luo P G . Registration of the novel wheat lines L658, L693, L696, and L699, with resistance toFusarium head blight, stripe rust, and powdery mildew
J Plant Regist, 2015,9:121-124.
DOI:10.3198/jpr2014.01.0003crgURL [本文引用: 1]

[30] Zhang X L, Shen X R, Hao Y F, Cai J J, Ohm H W, Kong LR . A genetic map ofLophopyrum ponticum chromosome 7E, harboring resistance genes to Fusarium head blight and leaf rust
Theor Appl Genet, 2011,122:263-270.
DOI:10.1007/s00122-010-1441-3URL [本文引用: 1]
The leaf rust resistance gene Lr19 and Fusarium head blight (FHB) resistance quantitative trait loci (QTL) derived from the wild wheatgrass Lophopyrum ponticum have been located on chromosome 7E. The main objectives of the present study were to develop a genetic map of chromosome 7E and map the two resistance loci using a population of 237 F(7:8) recombinant inbred lines (RILs) derived from a cross between two Thatcher-L. ponticum substitution lines, K11463 (7el(1)(7D)) and K2620 (7el(2)(7D)). 532 G-SSR, E-SSR and STS markers from wheat chromosome group 7 were screened in the parent lines. Of these, 118 markers were polymorphic, with a polymorphism frequency of 22.2%. A genetic map of L. ponticum chromosome 7E was constructed with 64 markers, covering 95.76 cM, with an average genetic distance of 1.47 cM between markers. The major FHB resistance locus, temporarily assigned as FhbLoP, was mapped to the very distal region of the long arm of chromosome 7E within a 3.71 cM interval flanked by Xcfa2240 and Xswes19, which accounts for 30.46% of the phenotypic variance. Lr19 was bracketed by Xwmc273 and XBE404744, with a map distance of 1.54 and 1.43 cM from either side, respectively. The closely linked markers identified in this study will be helpful for marker-assisted introgression of the L. ponticum-derived FhbLoP and Lr19 genes into elite cultivars of wheat, and the development of a genetic map will accelerate the map-based cloning of these two genes.

[31] 史文琦, 杨立军, 冯洁, 张旭, 曾凡松, 向礼波, 汪华, 喻大昭 . 小麦赤霉病流行区镰刀菌致病种及毒素化学型分析
植物病理学报, 2011,41:486-494.
URL [本文引用: 1]
为从分子水平上明确小麦赤霉病流行区镰刀菌致病种及其B 型毒素化学型的分布特点,本研究对2008 年度采自四川、重庆、湖北、安徽、江苏、河南6 省33 县市的赤霉病穗上分离获得的433 个镰刀菌单孢菌株,用鉴定种和鉴定B 型毒素化学型的特异性引物进行了鉴定分析。致病种检测结果表明,四川病穗检测到Fusarium asiaticum、F. graminearum、F.avenaceum 和F. meridionale 4 个镰刀菌种,重庆、湖北、安徽和江苏病穗检测到F. asiaticum 和F. graminearum 2 个种,河南病穗仅检测到F. graminearum 1 个种。毒素化学型检测结果表明,Nivalenol(NIV)是四川和重庆镰刀菌主要毒素化学型,Deoxynivalenol(DON)是湖北、河南、安徽和江苏镰刀菌主要毒素化学型;将DON 化学型进一步划分为3-AcDON 和15-AcDON 显示,四川、湖北、江苏镰刀菌毒素以3-AcDON 为主,安徽镰刀菌毒素为3-AcDON 和15-AcDON 两者参半,河南镰刀菌全部产生15-AcDON。结果揭示,F. asiaticum 是四川、重庆、湖北和江苏等赤霉病流行麦区的优势致病种;镰刀菌产生的DON 和NIV 毒素化学型存在明显的地域分布,长江上游的麦区以NIV 为优势化学型,长江中下游麦区以DON 为优势化学型;镰刀菌致病种与DON 毒素的化学型间存在一定关系。
Shi W Q, Yang L J, Feng J, Zhang X, Zeng F S, Xiang L B, Wang H, Yu D Z . Analysis on the population structure of Fusarium pathogenic spp. and its mycotoxin chemotypes in Fusarium head blight epidemic region
Acta Phytopathol Sin, 2011,41:486-494 (in Chinese with English abstract).
URL [本文引用: 1]
为从分子水平上明确小麦赤霉病流行区镰刀菌致病种及其B 型毒素化学型的分布特点,本研究对2008 年度采自四川、重庆、湖北、安徽、江苏、河南6 省33 县市的赤霉病穗上分离获得的433 个镰刀菌单孢菌株,用鉴定种和鉴定B 型毒素化学型的特异性引物进行了鉴定分析。致病种检测结果表明,四川病穗检测到Fusarium asiaticum、F. graminearum、F.avenaceum 和F. meridionale 4 个镰刀菌种,重庆、湖北、安徽和江苏病穗检测到F. asiaticum 和F. graminearum 2 个种,河南病穗仅检测到F. graminearum 1 个种。毒素化学型检测结果表明,Nivalenol(NIV)是四川和重庆镰刀菌主要毒素化学型,Deoxynivalenol(DON)是湖北、河南、安徽和江苏镰刀菌主要毒素化学型;将DON 化学型进一步划分为3-AcDON 和15-AcDON 显示,四川、湖北、江苏镰刀菌毒素以3-AcDON 为主,安徽镰刀菌毒素为3-AcDON 和15-AcDON 两者参半,河南镰刀菌全部产生15-AcDON。结果揭示,F. asiaticum 是四川、重庆、湖北和江苏等赤霉病流行麦区的优势致病种;镰刀菌产生的DON 和NIV 毒素化学型存在明显的地域分布,长江上游的麦区以NIV 为优势化学型,长江中下游麦区以DON 为优势化学型;镰刀菌致病种与DON 毒素的化学型间存在一定关系。

[32] 俞刚, 陈利锋, 谢卫平, 柴一秋 . 禾谷镰孢的产毒与致病性
南京农业大学学报, 2001,24(4):19-23.
[本文引用: 1]

Yu G, Chen L F, Xie W P, Chai Y Q . Correlation of trichothecene producing potential to pathogenicity of Fusarium graminearum on wheat
J Nanjing Agric Univ, 2001,24(4):19-23 (in Chinese with English abstract).
[本文引用: 1]

[33] 刘易科, 佟汉文, 朱展望, 陈泠, 邹娟, 张宇庆, 焦春海, 高春保 . 小麦赤霉病抗性改良研究进展
麦类作物学报, 2016,36:51-57.
[本文引用: 1]

Liu Y K, Tong H W, Zhu Z W, Chen L, Zou J, Zhang Y Q, Jiao C H, Gao C B . Review on improvement of Fusarium head blight resistance in wheat
J Triticeae Crops, 2016,36:51-57 (in Chinese with English abstract).
[本文引用: 1]

[34] 唐洪, 彭恒, 刘明龙, 杨建华, 吴吉林 . 小麦赤霉病田间病情与抽穗扬花期气象条件和病粒率关系
中国植保导刊, 2012,32(7):10-12.
[本文引用: 1]

Tang H, Peng H, Liu M L, Yang J H, Wu J L . Relationships among field condition of wheat scab and climate and percentage of Fusarium infected kernels at the heading and flowering stage
China Plant Protect, 2012,32:10-12 (in Chinese with English abstract).
[本文引用: 1]

[35] Mesterhazy A . Types and components of resistance toFusarium head blight of wheat
Plant breed, 1995,114:377-386.
DOI:10.1007/s00122-003-1272-6URLPMID:12768240 [本文引用: 1]
Fusarium head blight (FHB, scab) causes severe yield and quality losses, but the most serious concern is the mycotoxin contamination of cereal food and feed. The cultivation of resistant varieties may contribute to integrated control of this fungal disease. Breeding for FHB resistance by conventional selection is feasible, but tedious and expensive. The aim of this work was to detect QTLs for combined type I and type II resistance against FHB and estimate their effects in comparison to the QTLs identified previously for type II resistance. A population of 364, F1 derived doubled-haploid (DH) lines from the cross 'CM-82036' (resistant)/'Remus' (susceptible) was evaluated for components of FHB resistance during 2 years under field conditions. Plants were inoculated at anthesis with a conidial suspension of Fusarium graminearum or Fusarium culmorum. The crop was kept wet for 20 h after inoculation by mist-irrigation. Disease severity was assessed by visual scoring. Initial QTL analysis was performed on 239 randomly chosen DH lines and extended to 361 lines for putative QTL regions. Different marker types were applied, with an emphasis on PCR markers. Analysis of variance, as well as simple and composite interval mapping, revealed that two genomic regions were significantly associated with FHB resistance. The two QTLs on chromosomes 3B (Qfhs.ndsu-3BS) and 5A (Qfhs.ifa-5A) explained 29 and 20% of the phenotypic variance, respectively, for visual FHB severity. Qfhs.ndsu-3BS appeared to be associated mainly with resistance to fungal spread, and Qfhs.ifa-5A primarily with resistance to fungal penetration. Both QTL regions were tagged with flanking SSR markers. These results indicate that FHB resistance was under the control of two major QTLs operating together with unknown numbers of minor genes. Marker-assisted selection for these two major QTLs appears feasible and should accelerate the development of resistant and locally adapted wheat cultivars.