扬麦4号/偃展1号RIL群体株高QTL挖掘及其对赤霉病抗性的效应分析与验证

doi:10.3724/SP.J.1006.2023.31005

作物学报 ›› 2023, Vol. 49 ›› Issue (12): 3215-3226.doi: 10.3724/SP.J.1006.2023.31005

• 作物遗传育种·种质资源·分子遗传学 • 上一篇下一篇

扬麦4号/偃展1号RIL群体株高QTL挖掘及其对赤霉病抗性的效应分析与验证

赵蝶¹^,²(), 胡文静²^,³^,^*(), 程晓明², 王书平¹, 张春梅², 李东升², 高德荣²^,³^,^*()

¹长江大学农学院, 湖北荆州 434025
²江苏里下河地区农业科学研究所 / 农业农村部长江中下游小麦生物学与遗传育种重点实验室, 江苏扬州 225007
³扬州大学 / 江苏省粮食作物现代产业技术协同创新中心 / 江苏省植物功能基因组学重点实验室, 江苏扬州225009

收稿日期:2023-01-10 接受日期:2023-05-24 出版日期:2023-12-12 网络出版日期:2023-06-15
通讯作者: * 胡文静, E-mail: huren2008@126.com; 高德荣, E-mail: gdr@wheat.org.cn
作者简介:E-mail: zd2021720791@163.com
基金资助:
国家自然科学基金项目(31901544);泰州市科技计划项目(TN202117);江苏现代农业产业单项技术研发(CX(21)3063);江苏省重点研发项目(BE2021335)

Detection and verification of QTL for plant height in Yangmai 4/Yanzhan 1 recombinant inbred lines population and their genetic effects on Fusarium head blight resistance

ZHAO Die¹^,²(), HU Wen-Jing²^,³^,^*(), CHENG Xiao-Ming², WANG Shu-Ping¹, ZHANG Chun-Mei², LI Dong-Sheng², GAO De-Rong²^,³^,^*()

¹College of Agriculture, Yangtze University, Jingzhou 434025, Hubei, China
²Lixiahe Institute of Agriculture Sciences / Key Laboratory of Wheat Biology and Genetic Improvement for Low and Middle Yangtze Valley, Ministry of Agriculture and Rural Affairs, Yangzhou 225007, Jiangsu, China
³Jiangsu Co-Innovation Center for Modern Production Technology of Grain Crops / Jiangsu Key Laboratory of Crop Genomics and Molecular Breeding, Yangzhou University, Yangzhou 225009, Jiangsu, China

Received:2023-01-10 Accepted:2023-05-24 Published:2023-12-12 Published online:2023-06-15
Contact: * E-mail: huren2008@126.com; E-mail: gdr@wheat.org.cn
Supported by:
National Natural Science Foundation of China(31901544);Taizhou Science and Technology Project(TN202117);Jiangsu Modern Agricultural Industry Single Technology Research and Development(CX(21)3063);National Key Research and Development Program of Jiangsu(BE2021335)

摘要/Abstract

摘要：

小麦的株高(plant height, PH)性状与赤霉病(fusarium head blight, FHB)抗性的关系密切。本研究利用扬麦4号/偃展1号(YM4/YZ1)杂交组合衍生的重组自交系(recombinant inbred lines, RIL)群体, 利用55K单核苷酸多态性(single-nucleotide polymorphism, SNP)芯片数据, 结合3年共6个环境下RIL群体及其亲本的株高数据, 挖掘株高性状的遗传位点。同时利用土壤表面赤霉病麦粒抛撒法和单花滴注法鉴定株高位点对赤霉病抗侵染(Type I)和抗扩展(Type II)两种类型的效应。在染色体2D、4B、4D、5A和7D上检测到7个与株高相关的数量性状位点(quantitative trait loci, QTL), 经过比对, QPh.yas-2D.2可能是新的株高位点。QPh.yas-2D.1、QPh.yas-2D.2和QPh.yas-5A的矮秆效应来源于扬麦4号, 其余4个QTL的矮秆效应来源于偃展1号。QPh.yas-4D和QPh.yas-5A均在6个环境下被检测到, 表型变异贡献率(PVE)范围分别为19.48%~44.11%和10.48%~13.71%。研究发现Rht-D1和QPh.yas-7D.2位点上的高秆等位变异(YM4等位变异)分别降低侵染型平均病小穗率(average percentage of infected spikelets, PIS) 34.97%和19.09%, QPh.yas-2D.2和QPh.yas-5A位点上的矮秆等位变异(YM4等位变异)分别降低扩展型平均病小穗率(average percentage of diseased spikelets, PDS) 24.73%和14.56%。QPh.yas-5A的矮秆等位变异来源于阿夫。利用小麦中国春2.1版本的参考基因组信息分析QPh.yas-5A区间, 发现一共有146个有注释功能的高置信基因, 主要涉及合成细胞色素P450、脱水反应元件结合蛋白、乙烯响应转录因子、转录因子MYC2和细胞壁受体相关激酶等。进一步将QPh.yas-5A位点紧密连锁SNP标记转化成育种可用分子标记KASP-5A, 并在126份小麦品种(系)中初步验证其对株高和赤霉病抗性的效应。研究结果可为QPh.yas-5A的育种应用和精细定位奠定基础。

关键词: 小麦, 株高, 赤霉病, QTL, QPh.yas-5A, 候选基因

Abstract:

Plant height (PH) is associated with fusarium head blight (FHB) resistance in wheat. In this study, a recombinant inbred lines (RIL) population derived from the cross of Yangmai 4/Yanzhan 1 (YM4/YZ1) was used to mine the quantitative trait loci (QTL) of PH traits using 55K SNP (single-nucleotide polymorphism) chip data, combined with the PH data of RIL population and their parents in six environments for three continuous years. Soil surface inoculation method and single spikelet inoculation were used to identify the FHB resistance to infection (Type I) and spread (Type II), respectively. Seven QTLs related to PH were detected on chromosomes 2D, 4B, 4D, 5A, and 7D, and only QPh.yas-2D.2 may be a new QTL of PH after comparing with previous studies. The dwarfing effects of QPh.yas-2D.1, QPh.yas-2D.2, and QPh.yas-5A were derived from YM4, and the dwarfing effects of the other four QTLs were derived from YZ1. Both QPh.yas-4D and QPh.yas-5A could be detected in six environments, and phenotypic variation explained (PVE) rates ranged from 19.48%-44.11% and 10.48%-13.71%, respectively. The increasing alleles at the Rht-D1 and QPh.yas-7D.2 (YM4 allele) significantly reduced the average percentage of infected spikelets (PIS) by 34.97% and 19.09%, respectively. The dwarfing alleles at QPh.yas-2D.2 and QPh.yas-5A (YM4 allele) significantly reduced the average percentage of diseased spikelets (PDS) by 24.73% and 14.56%, respectively. The dwarfing allele of QPh.yas-5A was derived from Funo. Furthermore, we preliminary analyzed the genes within the physical interval of QPh.yas-5A using the reference genome information of wheat version 2.1. A total of 146 high-confidence annotated genes were detected in the target interval, which were mainly involved in the synthesis of cytochrome P450, dehydration response element-binding protein, ethylene response transcription factor, transcription factor MYC2, and cell wall receptor-associated kinases. The SNP marker closely linked to QPh.yas-5A was further converted into kompetitive allele-specific PCR marker KASP-5A, and its effect on plant height and FHB resistance was then verified in 126 wheat cultivars (lines). The results of this study could provide a solid foundation for future fine mapping QPh.yas-5A.

Key words: Triticum aestivum, plant height, fusarium head blight, QTL, QPh.yas-5A, candidate gene

赵蝶, 胡文静, 程晓明, 王书平, 张春梅, 李东升, 高德荣. 扬麦4号/偃展1号RIL群体株高QTL挖掘及其对赤霉病抗性的效应分析与验证[J]. 作物学报, 2023, 49(12): 3215-3226.

ZHAO Die, HU Wen-Jing, CHENG Xiao-Ming, WANG Shu-Ping, ZHANG Chun-Mei, LI Dong-Sheng, GAO De-Rong. Detection and verification of QTL for plant height in Yangmai 4/Yanzhan 1 recombinant inbred lines population and their genetic effects on Fusarium head blight resistance[J]. Acta Agronomica Sinica, 2023, 49(12): 3215-3226.

图/表 7

表1

图1

表2

图2

附表1

126个品种/品系株高、PIS和PDS相关QTL目标等位基因的分布"

数目 Number	品种名称 Name	PIS^a	PDS^b	株高 Plant height (cm)	KASP_5A^c
1	鄂麦174 Emai 174	66.25	21.21	83.50	A
2	鄂麦12 Emai 12	61.86	17.18	87.00	A
3	鄂麦19 Emai 19	74.04	56.14	87.75	A
4	宁12046 Ning 12046	55.17	15.63	89.50	A
5	宁1529 Ning 1529	32.53	5.96	85.94	A
6	宁1616 Ning 1616	9.64	22.93	84.50	A
7	宁16399 Ning 16399	12.86	26.85	81.13	A
8	宁17097 Ning 17097	14.46	35.63	86.63	A
9	宁17320 Ning 17320	20.00	8.40	85.13	A
10	宁17329 Ning 17329	28.57	20.31	86.69	A
11	宁17342 Ning 17342	22.35	10.09	83.75	A
12	宁麦15 Ningmai 15	57.69	24.88	85.19	A
13	宁麦16 Ningmai 16	52.00	17.46	85.63	A
14	宁麦18 Ningmai 18	52.11	12.55	86.81	A
15	宁麦19 Ningmai 19	53.16	25.14	83.00	A
16	宁麦24 Ningmai 24	52.78	17.46	79.38	A
17	宁麦3号 Ningmai 3	57.58	33.55	93.25	A
18	宁麦9号 Ningmai 9	42.11	17.34	87.75	A
19	宁麦15318 Ningmai 15318	77.38	8.45	79.81	A
20	生选3号 Shengxuan 3	68.00	33.02	90.88	A
21	苏麦5号 Sumai 5	27.47	7.17	82.38	A
22	扬173046 Yang 173046	41.67	25.10	80.13	A
23	扬辐麦3048 Yangfumai 3048	15.05	5.34	81.81	A
24	扬辐麦4188 Yangfumai 4188	35.63	16.79	82.13	A
25	扬辐麦5242 Yangfumai 5242	18.75	36.67	85.50	A
26	扬辐麦2 Yangfumai 2	39.08	32.23	87.13	A
27	扬辐麦6 Yangfumai6	55.95	56.64	82.88	A
28	扬辐麦9 Yangfumai 9	26.60	5.45	81.06	A
29	扬麦12 Yangmai 12	54.22	15.68	87.13	A
30	扬麦158 Yangmai 158	56.00	25.32	89.88	A
31	扬麦18 Yangmai 18	27.17	7.69	85.25	A
32	扬麦20 Yangmai 20	44.94	19.68	84.50	A
33	扬麦21 Yangmai 21	37.97	11.21	84.50	A
34	扬麦3号 Yangmai 3	52.08	69.70	93.00	A
35	扬麦4号 Yangmai 4	35.00	22.77	90.38	A
36	扬麦5号 Yangmai 5	22.31	26.36	88.00	A
37	镇麦11 Zhenmai 11	72.94	13.38	84.38	A
38	镇麦12 Zhenmai 12	44.58	7.51	89.81	A
39	镇麦5号 Zhenmai 5	45.74	8.45	85.88	A
40	镇麦8号 Zhenmai 8	44.44	5.45	86.50	A
41	宁7840 Ning 7840	62.50	43.98	92.69	A
42	百农3217 Bainong 3217	57.53	64.60	82.50	A
43	宁麦7号 Ningmai 7	71.59	37.46	79.38	A
44	皖麦19 Wanmai 19	76.58	92.26	83.13	A
45	扬辐麦9311 Yangfumai 9311	52.22	32.78	81.31	A
46	扬麦13 Yangmai 13	48.15	38.01	83.99	A
47	豫麦18 Yumai 18	80.43	64.95	85.38	A
48	周麦30 Zhoumai 30	47.73	32.39	78.81	A
49	周麦32 Zhoumai 32	84.38	46.14	78.00	A
50	博爱7023 Boai 7023	51.25	79.81	90.88	B
51	鄂麦11 Emai 11	56.18	41.78	88.13	B
52	鄂麦17 Emai 17	40.43	20.43	88.50	B
53	鄂麦6号 Emai 6	61.05	40.38	118.13	B
54	淮麦22 Huaimai 22	63.64	50.45	90.19	B
55	淮麦302 Huaimai 302	13.54	62.89	83.94	B
56	淮麦33 Huaimai 33	56.52	24.09	81.25	B
57	济麦77248 Jimai 77248	25.00	4.98	96.75	B
58	宁12188 Ning 12188	42.17	9.77	88.00	B
59	宁15186 Ning 15186	26.47	13.77	86.13	B
60	宁1625 Ning 1625	16.67	25.84	89.38	B
61	宁17073 Ning 17073	75.49	15.24	88.38	B
62	生选6号 Shengxuan 6	35.44	13.20	86.13	B
63	苏麦6号 Sumai 6	42.67	48.90	87.88	B
64	皖麦31 Wanmai 31	38.04	46.67	101.63	B
65	皖麦32 Wanmai 32	44.71	28.44	91.25	B
66	万年2号 Wannian 2	27.27	46.15	119.69	B
67	西农511 Xinong 511	36.26	6.53	84.13	B
68	信阳12 Xinyang 12	42.72	55.61	93.00	B
69	徐麦31 Xumai 31	11.22	75.65	80.38	B
70	扬辐麦5059 Yangfumai 5059	9.52	6.25	83.38	B
71	扬辐麦6076 Yangfumai 6076	18.95	24.00	84.38	B
72	扬辐麦7291 Yangfumai 7291	27.85	5.17	80.69	B
73	扬麦1号 Yangmai 1	66.67	99.17	107.63	B
74	扬麦14 Yangmai 14	46.15	4.88	90.00	B
75	扬麦15 Yangmai 15	84.85	22.81	81.63	B
76	扬麦16 Yangmai 16	51.19	39.15	91.81	B
77	扬麦23 Yangmai 23	58.43	61.45	82.63	B
78	扬麦27 Yangmai 27	41.41	14.77	81.94	B
79	扬麦28 Yangmai 28	37.76	7.06	87.00	B
80	扬麦29 Yangmai 29	42.35	14.29	84.88	B
81	扬麦6号 Yangmai 6	24.73	21.94	81.25	B
82	扬麦9号 Yangmai 9	53.76	12.87	85.25	B
83	扬糯麦1号 Yangnuomai 1	62.79	9.35	87.75	B
84	镇13056 Zhen 13056	36.14	35.53	80.63	B
85	镇麦1 Zhenmai 1	50.00	48.28	93.00	B
86	镇麦10 Zhenmai 10	44.32	37.25	81.88	B
87	镇麦168 Zhenmai 168	56.99	12.54	87.38	B
88	矮抗58 Aikang 58	87.76	58.78	82.44	B
89	百农418 Bainong 418	79.38	26.91	84.13	B
90	百农4199 Bainong 4199	87.96	35.00	85.63	B
91	鄂麦133 Emai 133	61.63	10.06	84.31	B
92	鄂麦178 Emai 178	73.42	39.02	87.38	B
93	鄂麦15 Emai 15	28.21	60.68	86.50	B
94	鄂麦16 Emai 16	82.80	34.13	84.56	B
95	丰德存麦1 Fengdecunmai 1	92.13	64.36	85.88	B
96	淮麦18 Huaimai 18	55.56	12.54	80.94	B
97	淮麦20 Huaimai 20	42.00	30.98	83.13	B
98	淮麦25 Huaimai 25	28.05	61.86	87.63	B
99	济麦44 Jimai 44	72.73	87.09	89.25	B
100	济麦77231 Jimai 77231	22.62	49.50	90.13	B
101	皖麦33 Wanmai 33	43.33	55.56	83.75	B
102	伟隆169 Weilong 169	48.48	25.08	82.69	B
103	西农20 Xinong 20	45.56	27.96	84.00	B
104	西农979 Xinong 979	75.76	31.12	84.94	B
105	新乡197 Xinxiang 197	91.25	21.84	86.81	B
106	新乡280 Xinxiang 280	60.64	18.57	86.94	B
107	新乡289 Xinxiang 289	26.21	23.91	82.88	B
108	徐麦15098 Xumai 15098	29.29	93.92	83.38	B
109	徐麦16144 Xumai 16144	34.72	47.09	82.63	B
110	徐麦16196 Xumai 16196	17.91	27.76	82.75	B
111	徐麦17252 Xumai 17252	10.87	11.59	82.13	B
112	徐麦17258 Xumai 17258	27.63	21.84	81.75	B
113	徐麦178 Xumai 178	58.67	86.36	82.00	B
114	徐麦2023 Xumai 2023	35.06	42.44	83.25	B
115	徐麦32 Xumai 32	7.14	81.12	83.13	B
116	徐麦33 Xumai 33	12.07	31.12	82.50	B
117	徐农029 Xunong 029	13.10	33.91	84.50	B
118	烟农1212 Yannong 1212	71.74	95.63	85.75	B
119	烟农19 Yannong 19	37.74	83.77	86.50	B
120	郑1342 Zheng 1342	42.99	36.67	84.13	B
121	郑1860 Zheng 1860	43.56	31.29	83.13	B
122	郑9188 Zheng 9188	67.68	19.75	83.50	B
123	郑麦9023 Zhengmai 9023	56.00	36.84	85.75	B
124	镇麦6号 Zhenmai 6	45.16	9.58	84.31	B
125	周麦18 Zhoumai 18	66.34	53.96	85.75	B
126	周麦36 Zhoumai 36	60.82	72.15	83.38	B

附表1

图3

附表2

QPh.yas-5A区间的高置信基因"

基因ID (CS 2.1) Gene ID (CS 2.1)	物理位置^a Physical location (Mb)^a	功能 Function	水稻的同源基因 Homologous gene in Oryza sativa	拟南芥的同源基因 Homologous gene in Arabidopsis thaliana
TraesCS5A03G0745700	516.05	Transcription factor MYC2 [UniProtKB/Swiss-Prot:Q39204]	Os09g0519100	NA
TraesCS5A03G0746000	517.21	Transcription factor MYC2 [UniProtKB/Swiss-Prot:Q39204]	Os09g0519100	NA
TraesCS5A03G0746100	517.36	Transcription factor MYC2 [UniProtKB/Swiss-Prot:Q39204]	Os09g0519100	NA
TraesCS5A03G0747200	519.16	Transcription factor MYC2 [UniProtKB/Swiss-Prot:Q39204]	Os09g0519100	NA
TraesCS5A03G0748400	520.44	Transcription factor MYC2 [UniProtKB/Swiss-Prot:Q39204]	Os09g0519100	NA
TraesCS5A03G0748500	520.60	DNA polymerase delta subunit 4 [UniProtKB/Swiss-Prot:Q3T0X9]	Os09g0520100	NA
TraesCS5A03G0748600	520.63	Probable 1-acylglycerol-3-phosphate O-acyltransferase [UniProtKB/Swiss-Prot:Q0J0A4]	Os09g0520200	AT4G24160
TraesCS5A03G0748700	520.64	Pentatricopeptide repeat-containing protein At2g36980, mitochondrial [UniProtKB/Swiss-Prot:Q9SJK9]	Os09g0520300	NA
TraesCS5A03G0748800	520.64	Cell division cycle 5-like protein [UniProtKB/Swiss-Prot:P92948]	Os04g0348300	AT1G09770
TraesCS5A03G0748900	520.65	Serine carboxypeptidase-like 18 [UniProtKB/Swiss-Prot:Q9C7Z9]	Os10g0101200	NA
TraesCS5A03G0749000	520.69	NA	Os11g0292050	NA
TraesCS5A03G0749700	520.84	Serine carboxypeptidase-like 18 [UniProtKB/Swiss-Prot:Q9C7Z9]	Os09g0462875	NA
TraesCS5A03G0749800	520.86	Probable acyl-activating enzyme 5, peroxisomal [UniProtKB/Swiss-Prot:Q9FFE6]	Os09g0555800	AT5G16370
TraesCS5A03G0750100	520.95	Premnaspirodiene oxygenase [UniProtKB/Swiss-Prot:A6YIH8]	Os09g0447300	NA
TraesCS5A03G0750200	520.98	Cytochrome P450 87A3 [UniProtKB/Swiss-Prot:Q7XU38]	Os09g0457100	NA
TraesCS5A03G0750300	521.12	NA	Os09g0498700	NA
TraesCS5A03G0750400	521.18	NA	NA	NA
TraesCS5A03G0750800	521.27	Ethylene-responsive transcription factor 1B [UniProtKB/Swiss-Prot:Q8LDC8]	Os09g0457900	NA
TraesCS5A03G0751000	521.40	NA	NA	NA
TraesCS5A03G0751100	521.42	RING-H2 finger protein ATL72 [UniProtKB/Swiss-Prot:Q9SG96]	Os09g0468300	NA
TraesCS5A03G0751300	521.47	RING-H2 finger protein ATL72 [UniProtKB/Swiss-Prot:Q9SG96]	Os09g0468300	NA
TraesCS5A03G0751600	521.74	Transcription factor RF2b [UniProtKB/Swiss-Prot:Q6S4P4]	Os09g0520400	NA
TraesCS5A03G0751700	521.74	NA	Os09g0520500	NA
TraesCS5A03G0751900	521.76	Ycf49-like protein [UniProtKB/Swiss-Prot:Q55720]	Os04g0379400	AT4G22830
TraesCS5A03G0752000	521.76	Probable sodium/metabolite cotransporter BASS5, chloroplastic [UniProtKB/Swiss-Prot:Q650U0]	Os09g0520600	AT4G22840
TraesCS5A03G0752100	521.83	Putative disease resistance protein RGA4 [UniProtKB/Swiss-Prot:Q7XA39]	Os09g0479500	NA
TraesCS5A03G0752600	522.02	Alpha-L-fucosidase 1 [UniProtKB/Swiss-Prot:Q8GW72]	Os09g0520800	NA
TraesCS5A03G0752700	522.03	DEAD-box ATP-dependent RNA helicase 7 [UniProtKB/Swiss-Prot:Q650T9]	Os09g0520700	AT5G62190
TraesCS5A03G0752800	522.03	Alpha-L-fucosidase 1 [UniProtKB/Swiss-Prot:Q8GW72]	Os09g0520800	NA
TraesCS5A03G0752900	522.09	3-ketoacyl-CoA synthase 4 [UniProtKB/Swiss-Prot:Q9LN49]	Os09g0521100	NA
TraesCS5A03G0753100	522.43	3-ketoacyl-CoA synthase 4 [UniProtKB/Swiss-Prot:Q9LN49]	Os09g0521100	NA
TraesCS5A03G0753200	522.48	Transcription factor PCF2 [UniProtKB/Swiss-Prot:Q6ZBH6]	Os09g0521300	AT5G51910
TraesCS5A03G0753600	522.50	F-box protein At1g52495 [UniProtKB/Swiss-Prot:Q9SSQ2]	Os09g0479100	NA
TraesCS5A03G0753700	522.59	NA	Os09g0517800	NA
TraesCS5A03G0753800	522.66	NA	Os09g0559500	NA
TraesCS5A03G0753900	522.66	ATPase GET3A [UniProtKB/Swiss-Prot:Q949M9]	Os09g0521500	AT1G01910
TraesCS5A03G0754100	522.78	SPX domain-containing membrane protein OsI_32082 [UniProtKB/Swiss-Prot:B8BDK8]	Os09g0521800	AT4G22990
TraesCS5A03G0754200	522.79	Flap endonuclease GEN-like 1 [UniProtKB/Swiss-Prot:Q64MA3]	Os09g0521900	NA
TraesCS5A03G0754300	522.79	Dehydration-responsive element-binding protein 1A [UniProtKB/Swiss-Prot:A2Z389]	Os09g0522200	AT5G51990
TraesCS5A03G0754400	522.81	Dehydration-responsive element-binding protein 1A [UniProtKB/Swiss-Prot:A2Z389]	Os09g0522000	AT5G51990
TraesCS5A03G0754500	522.83	Dehydration-responsive element-binding protein 1B [UniProtKB/Swiss-Prot:Q3T5N4]	Os09g0522000	AT5G51990
TraesCS5A03G0754700	522.84	NA	NA	NA
TraesCS5A03G0755100	522.85	Dehydration-responsive element-binding protein 1B [UniProtKB/Swiss-Prot:Q8GUW4]	Os09g0522000	AT5G51990
TraesCS5A03G0755300	522.95	Dehydration-responsive element-binding protein 1B [UniProtKB/Swiss-Prot:Q3T5N4]	Os09g0522000	AT5G51990
TraesCS5A03G0755500	523.08	Dehydration-responsive element-binding protein 1B [UniProtKB/Swiss-Prot:Q3T5N4]	Os09g0522000	AT5G51990
TraesCS5A03G0755900	523.15	Dehydration-responsive element-binding protein 1B [UniProtKB/Swiss-Prot:Q3T5N4]	Os09g0522000	AT5G51990
TraesCS5A03G0756000	523.33	Dehydration-responsive element-binding protein 1B [UniProtKB/Swiss-Prot:Q3T5N4]	Os09g0522000	AT5G51990
TraesCS5A03G0756100	523.34	Dehydration-responsive element-binding protein 1A [UniProtKB/Swiss-Prot:A2Z389]	Os09g0457900	AT5G51990
TraesCS5A03G0756400	523.58	Dehydration-responsive element-binding protein 1A [UniProtKB/Swiss-Prot:Q64MA1]	Os09g0457900	AT5G51990
TraesCS5A03G0756500	523.59	Dehydration-responsive element-binding protein 1B [UniProtKB/Swiss-Prot:Q3T5N4]	Os09g0522000	AT5G51990
TraesCS5A03G0756700	523.60	Dehydration-responsive element-binding protein 1H [UniProtKB/Swiss-Prot:Q0J090]	Os09g0522100	AT5G51990
TraesCS5A03G0756900	523.67	Dehydration-responsive element-binding protein 1H [UniProtKB/Swiss-Prot:Q0J090]	Os09g0522100	AT5G51990
TraesCS5A03G0757200	523.70	NA	NA	NA
TraesCS5A03G0757300	523.73	Dehydration-responsive element-binding protein 1B [UniProtKB/Swiss-Prot:Q8GUW4]	Os09g0522200	AT5G51990
TraesCS5A03G0757600	523.79	NA	NA	NA
TraesCS5A03G0757800	523.85	Dehydration-responsive element-binding protein 1H [UniProtKB/Swiss-Prot:Q0J090]	Os09g0522100	AT5G51990
TraesCS5A03G0758000	523.86	Dehydration-responsive element-binding protein 1A [UniProtKB/Swiss-Prot:Q64MA1]	Os09g0522000	AT5G51990
TraesCS5A03G0758600	524.16	Dehydration-responsive element-binding protein 1H [UniProtKB/Swiss-Prot:Q0J090]	Os09g0522100	AT5G51990
TraesCS5A03G0758700	524.28	NA	NA	NA
TraesCS5A03G0758800	524.30	Dehydration-responsive element-binding protein 1H [UniProtKB/Swiss-Prot:Q0J090]	Os09g0522100	AT5G51990
TraesCS5A03G0759100	524.32	Dehydration-responsive element-binding protein 1A [UniProtKB/Swiss-Prot:Q64MA1]	Os09g0522100	AT5G51990
TraesCS5A03G0759200	524.33	Dehydration-responsive element-binding protein 1A [UniProtKB/Swiss-Prot:A2Z389]	Os09g0522200	AT5G51990
TraesCS5A03G0759500	524.45	NA	NA	NA
TraesCS5A03G0759600	524.53	Protein DETOXIFICATION 50 [UniProtKB/Swiss-Prot:Q9FJ87]	Os09g0524300	AT5G52050
TraesCS5A03G0759800	524.57	NA	NA	NA
TraesCS5A03G0759900	524.59	Ethylene-responsive transcription factor RAP2-11 [UniProtKB/Swiss-Prot:Q6J9S1]	Os09g0522200	AT5G52020
TraesCS5A03G0760100	524.72	BAG family molecular chaperone regulator 1 [UniProtKB/Swiss-Prot:Q0WUQ1]	Os09g0524800	AT5G52060
TraesCS5A03G0760200	524.73	Ribosomal RNA processing protein 36 homolog [UniProtKB/Swiss-Prot:A7SL20]	Os09g0525200	AT1G12650
TraesCS5A03G0760400	524.76	F-box protein PP2-A13 [UniProtKB/Swiss-Prot:Q9LEX0]	Os09g0525300	AT5G52120
TraesCS5A03G0760500	524.76	E3 ubiquitin-protein ligase BIG BROTHER [UniProtKB/Swiss-Prot:Q8L649]	Os09g0525400	AT5G52140
TraesCS5A03G0760800	524.92	Ethylene-responsive transcription factor ERF071 [UniProtKB/Swiss-Prot:O22259]	NA	AT5G52020
TraesCS5A03G0760900	524.93	Glucuronoxylan 4-O-methyltransferase 1 [UniProtKB/Swiss-Prot:Q6NMK1]	Os12g0204500	AT1G09610
TraesCS5A03G0761400	524.95	Protein YY1 [UniProtKB/Swiss-Prot:O23810]	Os09g0525500	AT5G52160
TraesCS5A03G0761600	524.95	Plant UBX domain-containing protein 8 [UniProtKB/Swiss-Prot:F4JPR7]	Os09g0525600	NA
TraesCS5A03G0761700	525.02	Germin-like protein 9-3 [UniProtKB/Swiss-Prot:Q652P9]	Os09g0568700	NA
TraesCS5A03G0761800	525.02	Protein HAPLESS 2 [UniProtKB/Swiss-Prot:F4JP36]	Os09g0525700	AT4G11720
TraesCS5A03G0762400	525.10	Germin-like protein 9-3 [UniProtKB/Swiss-Prot:Q652P9]	Os09g0568600	NA
TraesCS5A03G0762500	525.10	Protein HAPLESS 2 [UniProtKB/Swiss-Prot:F4JP36]	Os09g0525700	AT4G11720
TraesCS5A03G0762700	525.11	Acyl transferase 1 [UniProtKB/Swiss-Prot:Q7G4G7]	Os09g0543900	NA
TraesCS5A03G0762800	525.11	Acyl transferase 1 [UniProtKB/Swiss-Prot:Q7G4G7]	Os09g0543900	NA
TraesCS5A03G0763100	525.37	RHOMBOID-like protein 2 [UniProtKB/Swiss-Prot:Q9CAN1]	Os09g0525900	AT1G12750
TraesCS5A03G0763300	525.38	NA	NA	NA
TraesCS5A03G0763600	525.38	NA	NA	NA
TraesCS5A03G0763800	525.42	Protein ROOT PRIMORDIUM DEFECTIVE 1 [UniProtKB/Swiss-Prot:Q689D6]	Os09g0526000	NA
TraesCS5A03G0764000	525.61	Homeobox-leucine zipper protein ROC6 [UniProtKB/Swiss-Prot:Q7Y0V7]	Os09g0526300	AT5G52170
TraesCS5A03G0764300	525.68	NA	Os09g0526500	NA
TraesCS5A03G0764400	525.73	Ethylene-responsive transcription factor ERF071 [UniProtKB/Swiss-Prot:O22259]	Os09g0522200	AT1G12610
TraesCS5A03G0764500	525.78	Heat stress transcription factor B-2c [UniProtKB/Swiss-Prot:Q652B0]	Os09g0526600	AT5G62020
TraesCS5A03G0764700	525.82	UDP-glucose 4-epimerase 3 [UniProtKB/Swiss-Prot:Q652A8]	Os09g0526700	AT1G12780
TraesCS5A03G0764900	525.82	NA	Os09g0526800	AT1G01225
TraesCS5A03G0765400	526.30	Probable polyamine transporter At3g13620 [UniProtKB/Swiss-Prot:Q9LHN7]	Os03g0374900	AT3G13620
TraesCS5A03G0765500	526.40	Nuclear speckle RNA-binding protein A [UniProtKB/Swiss-Prot:A1A6K6]	Os09g0527100	NA
TraesCS5A03G0765700	526.41	NA	NA	NA
TraesCS5A03G0765900	526.43	NA	NA	NA
TraesCS5A03G0766000	526.43	Probable serine/threonine-protein kinase WNK5 [UniProtKB/Swiss-Prot:Q0D541]	Os07g0584100	NA
TraesCS5A03G0766100	526.44	NA	Os09g0527500	NA
TraesCS5A03G0766400	526.62	Probable polyamine transporter At3g13620 [UniProtKB/Swiss-Prot:Q9LHN7]	NA	NA
TraesCS5A03G0766800	527.09	Ras-related protein RABA3 [UniProtKB/Swiss-Prot:Q9LNK1]	Os09g0527600	AT1G01200
TraesCS5A03G0766900	527.09	Rhodanese-like domain-containing protein 14, chloroplastic [UniProtKB/Swiss-Prot:Q94A65]	Os09g0530000	AT4G27700
TraesCS5A03G0767000	527.10	2-keto-3-deoxy-L-rhamnonate aldolase [UniProtKB/Swiss-Prot:B7NN63]	Os09g0529900	AT4G10750
TraesCS5A03G0767200	527.23	Subtilisin-like protease SBT1.4 [UniProtKB/Swiss-Prot:Q9LVJ1]	Os09g0530800	NA
TraesCS5A03G0767400	527.28	Vacuolar protein sorting-associated protein 22 homolog 1 [UniProtKB/Swiss-Prot:Q5M759]	Os09g0529700	AT4G27040
TraesCS5A03G0767500	527.28	NA	Os09g0529400	NA
TraesCS5A03G0767600	527.29	NA	Os09g0529350	NA
TraesCS5A03G0767700	527.29	NA	Os09g0529300	NA
TraesCS5A03G0768000	527.36	Rab GTPase-activating protein 22 [UniProtKB/Swiss-Prot:Q94BY9]	Os09g0528800	AT5G41940
TraesCS5A03G0768200	528.19	Cytochrome P450 78A6 [UniProtKB/Swiss-Prot:Q9ZNR0]	Os09g0528700	AT3G61880
TraesCS5A03G0768800	528.54	Mediator of RNA polymerase II transcription subunit 10b [UniProtKB/Swiss-Prot:F4HPA7]	Os09g0528300	AT1G26665
TraesCS5A03G0768900	528.60	Homeobox-leucine zipper protein HOX6 [UniProtKB/Swiss-Prot:Q651Z5]	Os09g0528200	NA
TraesCS5A03G0769100	528.61	30S ribosomal protein S31, mitochondrial [UniProtKB/Swiss-Prot:P47909]	Os09g0528100	AT2G21290
TraesCS5A03G0769300	528.61	Kinesin-like protein KIN-7J [UniProtKB/Swiss-Prot:Q651Z7]	Os09g0528000	NA
TraesCS5A03G0769500	528.81	B-box zinc finger protein 19 [UniProtKB/Swiss-Prot:C0SVM5]	Os09g0527900	AT2G21320
TraesCS5A03G0769600	528.96	Auxin-responsive protein IAA26 [UniProtKB/Swiss-Prot:Q652A1]	Os09g0527700	NA
TraesCS5A03G0769700	528.97	U-box domain-containing protein 73 [UniProtKB/Swiss-Prot:Q6K762]	NA	NA
TraesCS5A03G0769800	528.98	Endoglucanase 23 [UniProtKB/Swiss-Prot:Q69NF5]	Os09g0530200	AT4G39010
TraesCS5A03G0769900	529.06	Cytochrome b561, DM13 and DOMON domain-containing protein At5g54830 [UniProtKB/Swiss-Prot:Q9FFU6]	Os09g0557700	AT5G54830
TraesCS5A03G0770000	529.18	Cytochrome P450 71A1 [UniProtKB/Swiss-Prot:P24465]	Os09g0530300	NA
TraesCS5A03G0770100	529.25	LRR receptor-like serine/threonine-protein kinase EFR [UniProtKB/Swiss-Prot:C0LGT6]	Os09g0479200	NA
TraesCS5A03G0770200	529.27	Transcription initiation factor TFIID subunit 5 [UniProtKB/Swiss-Prot:Q6S7B0]	Os06g0649500	AT5G25150
TraesCS5A03G0770400	529.30	NA	NA	NA
TraesCS5A03G0770800	529.58	Cytochrome P450 71A1 [UniProtKB/Swiss-Prot:P24465]	Os09g0530300	NA
TraesCS5A03G0770900	529.60	Cytochrome P450 71A25 [UniProtKB/Swiss-Prot:Q9STK8]	Os09g0530300	NA
TraesCS5A03G0771000	529.63	Zinc finger CCCH domain-containing protein 19 [UniProtKB/Swiss-Prot:Q9SIV5]	Os09g0530500	NA
TraesCS5A03G0771400	529.77	7-deoxyloganetin glucosyltransferase [UniProtKB/Swiss-Prot:F8WKW1]	Os02g0578300	AT1G22360
TraesCS5A03G0771500	529.78	Dolichol-phosphate mannosyltransferase subunit 1 [UniProtKB/Swiss-Prot:Q9LM93]	Os03g0824400	AT1G20575
TraesCS5A03G0771600	529.78	NEDD8-activating enzyme E1 regulatory subunit AXR1 [UniProtKB/Swiss-Prot:P42744]	Os03g0820100	AT1G05180
TraesCS5A03G0771700	529.99	NA	NA	NA
TraesCS5A03G0771800	530.00	NA	Os09g0530700	NA
TraesCS5A03G0771900	530.06	Subtilisin-like protease SBT3.6 [UniProtKB/Swiss-Prot:Q8L7I2]	Os09g0530800	AT1G32980
TraesCS5A03G0772000	530.06	NA	Os09g0530900	NA
TraesCS5A03G0772100	530.13	Protein MEI2-like 6 [UniProtKB/Swiss-Prot:Q652K6]	Os09g0531200	NA
TraesCS5A03G0773500	530.88	NA	NA	NA
TraesCS5A03G0773600	530.99	Protein SHORT INTERNODES [UniProtKB/Swiss-Prot:Q9XGX0]	Os09g0531600	NA
TraesCS5A03G0773800	531.70	Xylanase inhibitor protein 1 [UniProtKB/Swiss-Prot:Q8L5C6]	NA	NA
TraesCS5A03G0773900	531.76	PTI1-like tyrosine-protein kinase At3g15890 [UniProtKB/Swiss-Prot:Q9LSC2]	Os09g0531701	NA
TraesCS5A03G0774500	532.08	NA	NA	NA
TraesCS5A03G0774700	532.14	Haloacid dehalogenase-like hydrolase domain-containing protein At2g33255 [UniProtKB/Swiss-Prot:Q8RYE9]	Os02g0816100	AT2G33255
TraesCS5A03G0776500	533.41	Uncharacterized protein L728 [UniProtKB/Swiss-Prot:Q5UNY4]	Os09g0499500	AT5G13210
TraesCS5A03G0777400	533.69	Polygalacturonate 4-alpha-galacturonosyltransferase [UniProtKB/Swiss-Prot:Q9LE59]	Os09g0531800	AT3G61130
TraesCS5A03G0777500	533.74	Protein STAY-GREEN, chloroplastic [UniProtKB/Swiss-Prot:Q652K1]	Os09g0532000	AT4G22920
TraesCS5A03G0777600	533.75	7-deoxyloganetin glucosyltransferase [UniProtKB/Swiss-Prot:F8WKW1]	NA	NA
TraesCS5A03G0777800	533.95	60S ribosomal protein L36a [UniProtKB/Swiss-Prot:O23290]	Os07g0523150	AT3G23390
TraesCS5A03G0777900	533.96	NDR1/HIN1-like protein 1 [UniProtKB/Swiss-Prot:Q9SRN0]	Os09g0532200	NA
TraesCS5A03G0778000	533.96	Two-component response regulator-like PRR95 [UniProtKB/Swiss-Prot:Q689G6]	Os09g0532400	NA
TraesCS5A03G0778100	534.01	Pentatricopeptide repeat-containing protein At4g01400, mitochondrial [UniProtKB/Swiss-Prot:Q8LDU5]	Os09g0532800	NA
TraesCS5A03G0778300	534.15	NA	Os09g0532700	NA
TraesCS5A03G0778700	534.22	Protein ODORANT1 [UniProtKB/Swiss-Prot:Q50EX6]	Os09g0532900	NA
TraesCS5A03G0779300	534.61	Pantothenate kinase 2 [UniProtKB/Swiss-Prot:Q0J035]	Os09g0533100	AT4G32180
TraesCS5A03G0779400	534.62	Glucan endo-1,3-beta-glucosidase, acidic isoform [UniProtKB/Swiss-Prot:P49237]	Os09g0533200	NA
TraesCS5A03G0779500	534.63	NA	NA	NA
TraesCS5A03G0779800	534.66	NA	NA	NA
TraesCS5A03G0780000	534.68	Probable inactive purple acid phosphatase 29 [UniProtKB/Swiss-Prot:Q9FMK9]	Os09g0533300	AT5G63140
TraesCS5A03G0780200	534.73	Probable inactive purple acid phosphatase 29 [UniProtKB/Swiss-Prot:Q9FMK9]	Os09g0533300	NA
TraesCS5A03G0780600	534.78	Probable inactive purple acid phosphatase 29 [UniProtKB/Swiss-Prot:Q9FMK9]	Os09g0533300	AT5G63140
TraesCS5A03G0780700	534.78	Chaperone protein ClpD1, chloroplastic [UniProtKB/Swiss-Prot:Q6H795]	Os02g0526400	AT5G51070
TraesCS5A03G0781100	534.91	Probable serine/threonine-protein kinase WNK9 [UniProtKB/Swiss-Prot:Q2QXC6]	Os09g0511000	NA
TraesCS5A03G0781200	534.91	Lon protease homolog 2, peroxisomal [UniProtKB/Swiss-Prot:B8BDV1]	Os09g0533400	AT5G47040
TraesCS5A03G0781300	534.94	Succinate dehydrogenase subunit 8A, mitochondrial [UniProtKB/Swiss-Prot:Q0DF13]	Os06g0124900	NA
TraesCS5A03G0781900	535.12	NA	NA	NA
TraesCS5A03G0782000	535.13	O-acyltransferase WSD1 [UniProtKB/Swiss-Prot:Q93ZR6]	NA	NA
TraesCS5A03G0782100	535.24	O-acyltransferase WSD1 [UniProtKB/Swiss-Prot:Q93ZR6]	NA	NA
TraesCS5A03G0782400	535.29	O-acyltransferase WSD1 [UniProtKB/Swiss-Prot:Q93ZR6]	NA	NA
TraesCS5A03G0782600	535.48	NA	NA	NA
TraesCS5A03G0782800	535.48	Probable serine/threonine-protein kinase PBL19 [UniProtKB/Swiss-Prot:Q9LTC0]	Os09g0533600	NA
TraesCS5A03G0783700	535.65	Pectin acetylesterase 3 [UniProtKB/Swiss-Prot:O80731]	NA	AT2G46930
TraesCS5A03G0783800	535.68	Chlorophyll a-b binding protein, chloroplastic [UniProtKB/Swiss-Prot:Q10HD0]	Os03g0592500	AT3G27690
TraesCS5A03G0783900	535.69	Putative HVA22-like protein g [UniProtKB/Swiss-Prot:Q9LR09]	Os09g0533800	AT5G42560
TraesCS5A03G0784200	535.80	Endoglucanase 24 [UniProtKB/Swiss-Prot:Q69SG5]	Os09g0533900	AT1G75680
TraesCS5A03G0784300	535.82	Putative cyclin-F1-2 [UniProtKB/Swiss-Prot:Q6K8S5]	NA	NA
TraesCS5A03G0784500	535.88	Seipin-2 [UniProtKB/Swiss-Prot:F4I340]	Os09g0534000	NA
TraesCS5A03G0784600	535.88	NA	NA	NA
TraesCS5A03G0784700	535.90	NA	NA	NA
TraesCS5A03G0784800	535.90	Peroxidase 40 [UniProtKB/Swiss-Prot:O23474]	Os08g0302000	AT4G16270
TraesCS5A03G0784900	535.90	Putative cyclin-F1-2 [UniProtKB/Swiss-Prot:Q6K8S5]	Os09g0466100	NA
TraesCS5A03G0785000	536.01	ER lumen protein-retaining receptor erd-2.2 [UniProtKB/Swiss-Prot:Q09473]	Os09g0534200	AT1G75760
TraesCS5A03G0785100	536.02	Transcription initiation factor IIB [UniProtKB/Swiss-Prot:Q8W0W3]	Os09g0534800	AT3G10330
TraesCS5A03G0785300	536.11	Putative serpin-Z8 [UniProtKB/Swiss-Prot:Q53P09]	NA	NA
TraesCS5A03G0785400	536.16	NA	NA	NA
TraesCS5A03G0785600	536.18	Wall-associated receptor kinase 5 [UniProtKB/Swiss-Prot:Q9LMN7]	Os09g0561450	NA
TraesCS5A03G0785700	536.19	Wall-associated receptor kinase 5 [UniProtKB/Swiss-Prot:Q9LMN7]	Os09g0561450	NA
TraesCS5A03G0786000	536.21	Wall-associated receptor kinase 5 [UniProtKB/Swiss-Prot:Q9LMN7]	Os09g0561000	NA
TraesCS5A03G0786100	536.22	NA	Os11g0616200	NA
TraesCS5A03G0786200	536.23	Triosephosphate isomerase, chloroplastic [UniProtKB/Swiss-Prot:P46225]	Os09g0535000	AT2G21170
TraesCS5A03G0786400	536.31	NA	NA	NA
TraesCS5A03G0786500	536.32	NA	Os12g0182800	AT1G48170
TraesCS5A03G0786600	536.33	NA	NA	NA
TraesCS5A03G0787400	536.36	NA	NA	NA
TraesCS5A03G0787700	536.39	NA	NA	NA
TraesCS5A03G0789200	536.44	NA	NA	NA
TraesCS5A03G0789500	536.53	UDP-glycosyltransferase 91B1 [UniProtKB/Swiss-Prot:Q9LSM0]	Os09g0518200	NA
TraesCS5A03G0789600	536.54	NA	Os03g0566600	AT2G26680

附表2

参考文献 69

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[2]	Su Z, Jin S, Lu Y, Zhang G R, Chao S M, Bai G H. Single nucleotide polymorphism tightly linked to a major QTL on chromosome 7A for both kernel length and kernel weight in wheat. Mol Breed, 2016, 36: 1-11. doi: 10.1007/s11032-015-0425-z
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[4]	Peng J R, Richards D E, Hartley N M, Murphy G P, Devos K M, Flintham J E, Beales J, Fish L J, Worland A J, Pelica F, Sudhakar D, Christou P, Snape J W, Gale M D, Harberd N P. ‘Green revolution’ genes encode mutant gibberellin response modulators. Nature, 1999, 400: 256-261. doi: 10.1038/22307
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环境 Enviroment	亲本 Parents		RIL群体 RIL population			遗传力 Heritability (h²)
环境 Enviroment	YM4	YZ1	最小值 Min.	最大值 Max.	平均值 Mean	遗传力 Heritability (h²)
E1	99.47	74.89^**	68.50	124.00	97.42	0.81
E2	100.00	73.38^**	65.25	130.75	99.14
E3	99.38	75.72^**	63.50	123.00	97.63
E4	98.73	80.32^**	65.75	124.00	101.73
E5	92.65	70.47^**	57.75	121.00	88.31
E6	97.00	76.50^**	64.50	127.50	98.43
BLUE	98.40	75.21^**	65.62	119.98	97.14

连锁群 Linkage group	环境 Environment	位点 QTL	物理位置 Physical interval (Mb)	侧翼标记 Flanking markers	LOD	贡献率 PVE (%)	加性效应 Additive
2D	E3, E4, BLUE	QPh.yas-2D.1	18.40-20.42	AX 109242530-AX108988107	5.69-6.56	3.72-7.06	-3.6 to -2.07
	E3, E4, BLUE	QPh.yas-2D.2	511.67-515.21	AX110552619-GWM539	5.25-7.03	8.04-8.64	-2.30 to -2.04
4B	E1, E2, E3, E6, BLUE	QPh.yas-4B	589.85-590.54	AX111577080-AX108807261	4.36-9.43	3.16-10.76	2.22 to 3.69
4D	E1, E2, E3, E4, E5, E6, BLUE	QPh.yas-4D	19.19-19.59	Rht-D1_SNP-AX110572006	15.50-35.95	19.48-44.11	5.94 to 10.01
5A	E1, E2, E3, E4, E5, E6, BLUE	QPh.yas-5A	516.44-536.32	AX109341178-AX111618105	8.31-12.78	10.48-13.71	-4.84 to -3.73
7D	E1, E3, E4, E5, E6, BLUE	QPh.yas-7D.1	62.23-68.26	AX110468744-AX111061288	3.93-13.10	4.13-11.09	2.32 to 4.63
	E1, E2, E5, E6, BLUE	QPh.yas-7D.2	147.70-164.48	AX109847755-AX111460143	4.54-11.18	3.60-9.75	2.14 to 4.73

扬麦4号/偃展1号RIL群体株高QTL挖掘及其对赤霉病抗性的效应分析与验证

Detection and verification of QTL for plant height in Yangmai 4/Yanzhan 1 recombinant inbred lines population and their genetic effects on Fusarium head blight resistance

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