苗草专用型大麦品种鉴定及全基因组关联分析

doi:10.3724/SP.J.1006.2025.41028

作物学报 ›› 2025, Vol. 51 ›› Issue (1): 91-102.doi: 10.3724/SP.J.1006.2025.41028

• 作物遗传育种·种质资源·分子遗传学 • 上一篇下一篇

苗草专用型大麦品种鉴定及全基因组关联分析

马敏虎¹^,²(), 常华瑜¹^,², 陈朝燕², 仁增², 刘廷辉³, 邢国芳¹^,^*(), 郭刚刚²^,^*()

¹山西农业大学农学院 / 杂粮种质创新与分子育种山西省重点实验室, 山西晋中 030801
²中国农业科学院作物科学研究所 / 农业农村部粮食作物基因资源评价利用重点实验室 / 农作物基因资源与基因改良国家重大科学工程, 北京 100081
³甘孜藏族自治州农业科学研究所, 四川甘孜 626000

收稿日期:2024-04-16 接受日期:2024-09-18 出版日期:2025-01-12 网络出版日期:2024-10-10
通讯作者: *邢国芳, E-mail: sxauxgf@126.com; 郭刚刚, E-mail: guoganggang@caas.cn
作者简介:E-mail: mmh19984052889@163.com
基金资助:
国家重点研发计划项目(2022YFD2301301-1);山西省现代农业产业技术体系建设(杂粮, 2023CYJSTX03-19);杂粮种质创新与分子育种国家实验室(筹)(202204010910001-06);财政部和农业农村部国家现代农业产业技术体系建设专项(CARS-05)

Identification and genome-wide association study of specialized seedling grass barley cultivars

MA Min-Hu¹^,²(), CHANG Hua-Yu¹^,², CHEN Zhao-Yan², REN Zeng², LIU Ting-Hui³, XING Guo-Fang¹^,^*(), GUO Gang-Gang²^,^*()

¹College of Agronomy, Shanxi Agricultural University / Shanxi Key Laboratory of Germplasm Innovation and Molecular Breeding of Minor Crop, Jinzhong 030801, Shanxi, China
²Institute of Crop Sciences, Chinese Academy of Agricultural Sciences / Key Laboratory of Grain Crop Genetic Resources Evaluation and Utilization, Ministry of Agriculture and Rural Affairs / National Major Scienti?c Project of Crop Genetic Resources and Genetic Improvement, Beijing 100081, China
³Ganzi Tibetan Autonomous Prefecture Institute of Agricultural Sciences, Ganzi 626000, Sichuan, China

Received:2024-04-16 Accepted:2024-09-18 Published:2025-01-12 Published online:2024-10-10
Contact: *E-mail: sxauxgf@126.com; E-mail: guoganggang@caas.cn
Supported by:
National Key Research and Development Program of China(2022YFD2301301-1);Modern Agro-Industry Technology Research System of Shanxi Province(Minor Crops, 2023CYJSTX03-19);National Laboratory for Minor Crops Germplasm Innovation and Molecular Breeding (in Preparation)(202204010910001-06);China Agriculture Research System of MOF and MARA(CARS-05)

摘要/Abstract

摘要：

苗草工厂为实现草食动物的饲草周年供应提供了新的解决方案。针对苗草生产的品种需求, 本研究对124份中国大麦育成品种(系)与种质开展了苗草转化率鉴定和生物量调控位点发掘。结果显示, 大麦种子萌动后, 水培条件下苗草生物量呈指数增长趋势并在7 d后进入平台期。在植物工厂条件下, 鉴定出冬青16、扎青6号等10个高苗草转化率品种且分析发现苗草生物量与籽粒千粒重呈一定的负相关。进一步通过全基因组关联分析, 共鉴定到12个苗草生物量相关QTL位点, 从中预测出8个调控苗草生物量的候选基因。本研究不仅为大麦苗草工厂化生产筛选出高转化率品种, 同时也为苗草专用型大麦品种的遗传改良奠定了基础。

关键词: 大麦苗草, 生物量调控位点, 全基因组关联分析

Abstract:

Seedling grass factories represent an innovative strategy to ensure a consistent year-round forage supply for herbivores. This study was conducted to assess the conversion efficiency of seedling grass and to identify loci governing biomass in a diverse collection of 124 Chinese barley breeding varieties and germplasm. Our results indicate that following barley seed germination, the biomass of seedling grass in hydroponic systems undergoes exponential growth, leveling off after seven days. Within the plant factory setting, we have pinpointed ten varieties with superior conversion rate for seedling grass, including Dongqing 16 and Zaqing 6. A negative correlation was detected between the biomass of seedling grass and the thousand-kernel weight of the seeds. Genome-wide association studies revealed twelve QTL loci linked to seedling grass biomass, and eight candidate genes implicated in biomass regulation were predicted. This research not only identifies high-conversion varieties suitable for production in barley seedling grass factories but also lays the groundwork for genetically enhancing barley varieties specifically for seedling grass production.

Key words: barley seedling grass, biomass regulatory loci, genome-wide association study

马敏虎, 常华瑜, 陈朝燕, 仁增, 刘廷辉, 邢国芳, 郭刚刚. 苗草专用型大麦品种鉴定及全基因组关联分析[J]. 作物学报, 2025, 51(1): 91-102.

MA Min-Hu, CHANG Hua-Yu, CHEN Zhao-Yan, REN Zeng, LIU Ting-Hui, XING Guo-Fang, GUO Gang-Gang. Identification and genome-wide association study of specialized seedling grass barley cultivars[J]. Acta Agronomica Sinica, 2025, 51(1): 91-102.

图/表 11

附表1

参试大麦品种及其来源"

编号 Number	品种 Cultivar	类型 Type	皮裸 Hull/naked	棱型 Row	原产地 Origin
1	皖饲麦1号 Wansimai 1	饲用大麦 Feed barley	皮 Hull	6	安徽 Anhui
2	皖饲麦2号 Wansimai 2	饲用大麦 Feed barley	皮 Hull	6	安徽 Anhui
3	中饲麦1号 Zhongsimai 1	饲用大麦 Feed barley	皮 Hull	6	北京 Beijing
4	甘青6号 Ganqing 6	青稞 Qingke barley	裸 Naked	6	甘肃 Gansu
5	黄青1号 Huangqing 1	青稞 Qingke barley	裸 Naked	6	甘肃 Gansu
6	垦啤2号 Kenpi 2	啤用大麦 Malting barley	皮 Hull	2	甘肃 Gansu
7	垦啤3号 Kenpi 3	啤用大麦 Malting barley	皮 Hull	2	甘肃 Gansu
8	甘啤2号 Ganpi 2	啤用大麦 Malting barley	皮 Hull	2	甘肃 Gansu
9	甘啤3号 Ganpi 3	啤用大麦 Malting barley	皮 Hull	2	甘肃 Gansu
10	甘啤5号 Ganpi 5	啤用大麦 Malting barley	皮 Hull	2	甘肃 Gansu
11	甘啤6号 Ganpi 6	啤用大麦 Malting barley	皮 Hull	2	甘肃 Gansu
12	甘垦啤7号 GanKenpi 7	啤用大麦 Malting barley	皮 Hull	2	甘肃 Gansu
13	甘青9 Ganqing 9	食饲兼用Barley for food and feed	裸 Naked	6	甘肃 Gansu
14	垦啤麦2号 Kenpimai 2	啤用大麦 Malting barley	皮 Hull	6	黑龙江 Heilongjiang
15	垦啤麦3号 Kenpimai 3	啤用大麦 Malting barley	皮 Hull	2	黑龙江 Heilongjiang
16	垦啤麦5号 Kenpimai 5	啤用大麦 Malting barley	皮 Hull	6	黑龙江 Heilongjiang
17	垦啤麦6号 Kenpimai 6	啤用大麦 Malting barley	皮 Hull	2	黑龙江 Heilongjiang
18	垦啤麦9号 Kenpimai 9	啤用大麦 Malting barley	皮 Hull	6	黑龙江 Heilongjiang
19	龙啤麦2号 Longpimai 2	啤用大麦 Malting barley	皮 Hull	2	黑龙江 Heilongjiang
20	龙啤麦4号 Longpimai 4	啤用大麦 Malting barley	皮 Hull	6	黑龙江 Heilongjiang
21	华大麦2号 Huadamai 2	啤用大麦 Malting barley	皮 Hull	2	湖北 Hubei
22	华大麦6号 Huadamai 6	啤用大麦 Malting barley	皮 Hull	2	湖北 Hubei
23	华大麦7号 Huadamai 7	啤用大麦 Malting barley	皮 Hull	2	湖北 Hubei
24	鄂大麦9号 Edamai 9	饲用大麦 Feed barley	皮 Hull	2	湖北 Hubei
25	华大麦3号 Huadamai 3	饲用大麦 Feed barley	皮 Hull	6	湖北 Hubei
26	华大麦8号 Huadamai 8	饲用大麦 Feed barley	皮 Hull	2	湖北 Hubei
27	华大麦9号 Huadamai 9	饲用大麦 Feed barley	皮 Hull	2	湖北 Hubei
28	鄂大麦507 Edamai 507	食饲兼用 Barley for food and feed	裸 Naked	2	湖北 Hubei
29	华大麦1号 Huadamai 1	食饲兼用 Barley for food and feed	裸 Naked	2	湖北 Hubei
30	鄂大麦8号 Edamai 8	啤饲兼用 Barley for malting and feed	皮 Hull	2	湖北 Hubei
31	扬农啤2号 Yangnongpi 2	啤用大麦 Malting barley	皮 Hull	2	江苏 Jiangsu
32	扬农啤4号 Yangnongpi 4	啤用大麦 Malting barley	皮 Hull	2	江苏 Jiangsu
33	扬农啤7号 Yangnongpi 7	啤用大麦 Malting barley	皮 Hull	2	江苏 Jiangsu
34	盐引1号 Yanyin 1	啤用大麦 Malting barley	皮 Hull	2	江苏 Jiangsu
35	盐麦3号 Yanmai 3	啤用大麦 Malting barley	皮 Hull	2	江苏 Jiangsu
36	扬农啤10号 Yangnongpi 10	啤用大麦 Malting barley	皮 Hull	2	江苏 Jiangsu
37	扬农啤11号 Yangnongpi 11	啤用大麦 Malting barley	皮 Hull	2	江苏 Jiangsu
38	扬农啤12号 Yangnongpi 12	啤用大麦 Malting barley	皮 Hull	2	江苏 Jiangsu
39	泰兴元麦 Taixingyuanmai	—	裸 Naked	2	江苏 Jiangsu
40	盐选一号 Yanxuanyihao	—	皮 Hull	2	江苏 Jiangsu
41	盐选50 Yanxuan 50	—	皮 Hull	2	江苏 Jiangsu
42	蒙啤麦1号 Mengpimai 1	啤用大麦 Malting barley	皮 Hull	2	内蒙古Inner Mongolia
43	蒙啤麦3号 Mengpimai 3	啤用大麦 Malting barley	皮 Hull	6	内蒙古Inner Mongolia
44	蒙啤麦4号 Mengpimai 4	啤用大麦 Malting barley	皮 Hull	2	内蒙古Inner Mongolia
45	蒙古国草Mengguguocaomai	—	皮 Hull	2	内蒙古Inner Mongolia
46	北青1号 Beiqing 1	青稞 Qingke barley	裸 Naked	6	青海 Qinghai
47	北青2号 Beiqing 2	青稞 Qingke barley	裸 Naked	6	青海 Qinghai
48	北青3号 Beiqing 3	青稞 Qingke barley	裸 Naked	6	青海 Qinghai
49	北青4号 Beiqing 4	青稞 Qingke barley	裸 Naked	6	青海 Qinghai
50	北青5号 Beiqing 5	青稞 Qingke barley	裸 Naked	6	青海 Qinghai
51	北青6号 Beiqing 6	青稞 Qingke barley	裸 Naked	6	青海 Qinghai
52	北青7号 Beiqing 7	青稞 Qingke barley	裸 Naked	6	青海 Qinghai
53	北青8号 Beiqing 8	青稞 Qingke barley	裸 Naked	6	青海 Qinghai
54	东青1号 Dongqing 1	青稞 Qingke barley	裸 Naked	6	青海 Qinghai
55	互青2号 Huqing 2	青稞 Qingke barley	裸 Naked	6	青海 Qinghai
56	昆仑13号 Kunlun 13	青稞 Qingke barley	裸 Naked	6	青海 Qinghai
57	昆仑10号 Kunlun 10	—	皮 Hull	6	青海 Qinghai
58	济皮02 Jipi 02	—	皮 Hull	2	山东 Shandong
59	花22 Hua 22	啤用大麦 Malting barley	皮 Hull	2	上海 Shanghai
60	沪麦4号 Humai 4	啤用大麦 Malting barley	皮 Hull	2	上海 Shanghai
61	申麦2号 Shenmai 2	—	皮 Hull	2	上海 Shanghai
62	沪麦8号 Humai 8	—	皮 Hull	2	上海 Shanghai
63	西大麦2号 Xidamai 2	饲用大麦 Feed barley	皮 Hull	6	四川 Sichuan
64	西大麦3号 Xidamai 3	饲用大麦 Feed barley	皮 Hull	6	四川 Sichuan
65	藏青148 Zangqing 148	青稞 Qingke barley	裸 Naked	6	西藏 Xizang
66	藏青690 Zangqing 690	青稞 Qingke barley	裸 Naked	6	西藏 Xizang
67	冬青8号 Dongqing 8	青稞 Qingke barley	裸 Naked	6	西藏 Xizang
68	冬青16 Dongqing 16	青稞 Qingke barley	裸 Naked	6	西藏 Xizang
69	新啤1号 Xinpi 1	啤用大麦 Malting barley	皮 Hull	2	新疆 Xinjiang
70	新啤2号 Xinpi 2	啤用大麦 Malting barley	皮 Hull	2	新疆 Xinjiang
71	新啤6号 Xinpi 6	啤用大麦 Malting barley	皮 Hull	2	新疆 Xinjiang
72	新啤4号 Xinpi 4	啤用大麦 Malting barley	皮 Hull	2	新疆 Xinjiang
73	云稞1号 Yunke 1	青稞 Qingke barley	裸 Naked	6	云南 Yunnan
74	云啤2号 Yunpi 2	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
75	凤大麦6号 Fengdamai 6	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
76	云啤9号 Yunpi 9	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
77	云啤11号 Yunpi 11	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
78	云啤12号 YunPi 12	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
79	云啤14号 YunPi 14	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
80	云玉麦1号 Yunyumai 1	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
81	云啤15号 Yunpi 15	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
82	云啤17号 Yunpi 17	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
83	云啤18号 Yunpi 18	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
84	云靖麦2号 Yunjingmai 2	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
85	凤大麦9号 Fengdamai 9	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
86	凤大麦10号 Fengdamai 10	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
87	凤大麦11号 Fengdamai 11	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
88	凤大麦13号 Fengdamai 13	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
89	凤啤麦1号 Fengpimai 1	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
90	凤啤麦2号 Fengpimai 2	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
91	云大麦2号 Yundamai 2	饲用大麦 Feed barley	皮 Hull	2	云南 Yunnan
92	凤大麦7号 Fengdamai 7	饲用大麦 Feed barley	皮 Hull	2	云南 Yunnan
93	凤03-39 Feng03-39	饲用大麦 Feed barley	皮 Hull	6	云南 Yunnan
94	云饲麦1号 Yunsimai 1	饲用大麦 Feed barley	皮 Hull	6	云南 Yunnan
95	云饲麦3号 Yunsimai 3	饲用大麦 Feed barley	皮 Hull	6	云南 Yunnan
96	云饲麦4号 Yunsimai 4	饲用大麦 Feed barley	皮 Hull	6	云南 Yunnan
97	云饲麦8号 Yunsimai 8	饲用大麦 Feed barley	皮 Hull	6	云南 Yunnan
98	云饲麦9号 Yunsimai 9	饲用大麦 Feed barley	皮 Hull	6	云南 Yunnan
99	凤大麦12号 Fengdamai 12	饲用大麦 Feed barley	皮 Hull	2	云南 Yunnan
100	保大麦8号 Baodamai 8	饲用大麦 Feed barley	皮 Hull	6	云南 Yunnan
101	保大麦13号 Baodamai 13	饲用大麦 Feed barley	皮 Hull	6	云南 Yunnan
102	保大麦14号 Baodamai 14	饲用大麦 Feed barley	皮 Hull	6	云南 Yunnan
103	保大麦15号 Baodamai 15	饲用大麦 Feed barley	皮 Hull	6	云南 Yunnan
104	保大麦16号 Baodamai 16	饲用大麦 Feed barley	皮 Hull	6	云南 Yunnan
105	保大麦18号 Baodamai 18	饲用大麦 Feed barley	皮 Hull	6	云南 Yunnan
106	保大麦19号 Baodamai 19	饲用大麦 Feed barley	皮 Hull	6	云南 Yunnan
107	保大麦20号 Baodamai 20	饲用大麦 Feed barley	皮 Hull	6	云南 Yunnan
108	迪青4号 Diqing 4	青稞 Qingke barley	裸 Naked	6	云南 Yunnan
109	迪青1号 Diqing 1	粮用大麦 Grain barley	裸 Naked	6	云南 Yunnan
110	扎青6号 Zhaqing 6	—	裸 Naked	6	云南 Yunnan
111	楚08YD-4 Chu08YD-4	—	皮 Hull	6	云南 Yunnan
112	楚09YD-6 Chu08YD-6	—	皮 Hull	2	云南 Yunnan
113	楚09YD-9 Chu08YD-9	—	皮 Hull	2	云南 Yunnan
114	凤大麦8号 Fengdamai 8	—	皮 Hull	2	云南 Yunnan
115	保大麦22号 Baodamai 22	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
116	保大麦23号 Baodamai 23	啤用大麦 Malting barley	皮 Hull	2	云南 Yunnan
117	凤啤麦4号 Fengpimai 4	—	皮 Hull	2	云南 Yunnan
118	浙皮3号 Zhepi 3	啤用大麦 Malting barley	皮 Hull	2	浙江 Zhejiang
119	浙皮8号 Zhepi 8	啤用大麦 Malting barley	皮 Hull	2	浙江 Zhejiang
120	浙皮7号 Zhepi 7	饲用大麦 Feed barley	皮 Hull	2	浙江 Zhejiang
121	浙原18 Zheyuan 18	饲用大麦 Feed barley	皮 Hull	2	浙江 Zhejiang
122	浙秀22 Zhexiu 22	饲用大麦 Feed barley	皮 Hull	2	浙江 Zhejiang
123	浙秀12 Zhexiu 12	啤饲兼用Malting and Feed barley	皮 Hull	2	浙江 Zhejiang
124	浙皮9号 Zhepi 9	—	皮 Hull	2	浙江 Zhejiang

附表1

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表2

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表3

表4

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环境 Environment	平均值 Mean	标准差 SE	最小值 Min.	最大值 Max.	偏度 Skewness	峰度 Kurtosis	变异系数 CV (%)
2022-BJ	6.1	1.1	3.5	8.3	-0.081	-0.686	17.61
2022-GS	6.2	1.0	2.9	8.4	-0.208	0.936	15.91
2023-BJ	7.9	1.3	3.8	10.5	-0.601	0.229	16.18
2023-GS	7.5	1.2	3.5	10.2	-0.391	0.271	15.72

品种 Cultivar	苗草转化率 GSR				千粒重 1000-grain weight (g)	发芽率Germination rate (%)	综合排名 Comprehensive ranking
品种 Cultivar	2022-BJ	2022-GS	2023-BJ	2023-GS	千粒重 1000-grain weight (g)	发芽率Germination rate (%)	综合排名 Comprehensive ranking
冬青16 Dongqing 16	7.2	6.9	10.2	8.9	37.7±3.7	75.2±15.2	1
扎青6号 Zhaqing 6	7.1	6.6	9.3	9.9	44.2±6.0	76.6±5.4	2
华大麦9号 Huadamai 9	6.1	7.8	9.4	10.0	48.3±5.2	76.0±6.2	3
凤大麦12号 Fengdamai 12	7.0	7.1	9.9	8.2	55.4±3.3	78.8±11.5	4
昆仑13号 Kunlun 13	7.7	7.5	8.6	8.4	39.2±6.1	85.7±10.0	5
凤啤麦4号 Fengpimai 4	6.8	7.6	8.5	9.0	43.1±5.0	77.3±7.7	6
垦啤麦2号 Kenpimai 2	6.8	7.7	9.2	8.0	41.4±5.4	83.5±9.1	7
云啤12号 Yunpi 12	7.1	5.8	9.2	8.9	50.3±7.4	81.4±14.1	8
北青7号 Beiqing 7	7.1	5.4	9.2	9.7	43.5±8.1	83.6±9.5	9
云饲麦1号 Yunsimai 1	7.3	7.9	9.4	5.7	49.3±5.1	60.3±20.4	10

染色体 Chr.	QTL	位置 Position (Mb)	标记 Marker	环境 Environment	-log₁₀P	PVE (%)
1H	qgsr1H.1	426.4-430.4	rs1_428421759	2022-BJ	4.240	12.33
2H	qgsr2H.1	413.9-417.9	rs2_415944523	2022-GS	4.366	12.73
2H	qgsr2H.2	457.5-490.1	rs2_459543120, rs2_488125716	2022-BJ	4.343	11.66-12.65
3H	qgsr3H.1	108.6-112.6	rs3_110575889	2023-GS	4.004	11.60
			rs3_109862073	2023-BJ	3.985	11.54
	qgsr3H.2	615.3-619.3	rs3_617299505	2023-GS	4.354	12.69
4H	qgsr4H.1	17.1-21.1	rs4_19065845	2023-GS	4.274	12.44
4H	qgsr4H.2	362.9-366.9	rs4_364937809	2023-BJ	4.140	12.02
5H	qgsr5H.1	294.4-298.4	rs5_296385024	2023-GS	4.445	12.97
5H	qgsr5H.2	520.3-524.3	rs5_522318778	2023-BJ	4.780	14.01
6H	qgsr6H.1	289.2-293.2	rs6_291151063	2022-BJ	5.546	16.35
	qgsr6H.2	304.2-308.2	rs6_306192909	2023-BJ	4.178	12.14
	qgsr6H.3	435.4-439.4	rs6_437361512	2022-GS	4.007	11.60

QTL	基因名称 Gene ID	起始-终止 Start-end	注释信息 Annotation information
qgrs1H.1	HORVU.MOREX.r3.1HG0065130	427509599-427511636	NAC domain-containing protein
qgrs2H.2	HORVU.MOREX.r3.2HG0168310	475735626-475738921	Cellulose synthase family protein
	HORVU.MOREX.r3.2HG0170150	489043031-489043723	Pseudo-response regulator 3
	HORVU.MOREX.r3.2HG0170210	489506920-489509541	Pentatricopeptide repeat-containing protein
qgrs4H.1	HORVU.MOREX.r3.4HG0337240	17745839-17748824	Glutamine synthetase
	HORVU.MOREX.r3.4HG0337970	20646444-20650610	Transcription factor Inducer of CBF expression 1
	HORVU.MOREX.r3.4HG0338080	20945447-20949766	Peptide transporter
qgrs5H.2	HORVU.MOREX.r3.5HG0509460	523994766-523997934	WRKY transcription factor

苗草专用型大麦品种鉴定及全基因组关联分析

Identification and genome-wide association study of specialized seedling grass barley cultivars

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