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作物学报 ›› 2024, Vol. 50 ›› Issue (2): 265-279.doi: 10.3724/SP.J.1006.2024.34131

• 综述 •    下一篇

谷子中矮秆资源创制、遗传解析和育种利用

刁现民1,*(), 王立伟1, 智慧1, 张俊1, 李顺国2, 程汝宏2   

  1. 1作物基因资源与育种全国重点实验室 / 中国农业科学院作物科学研究所, 北京 100081
    2河北省农林科学院谷子研究所, 河北石家庄 050035
  • 收稿日期:2023-07-20 接受日期:2023-09-28 出版日期:2024-02-12 网络出版日期:2023-10-11
  • 通讯作者: *刁现民, E-mail: diaoxianmin@caas.cn
  • 基金资助:
    国家自然科学基金项目(32241042);国家自然科学基金项目(31771807);国家重点研发计划项目(2021YFF1000103);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-06-04);中国农业科学院创新工程项目资助

Development, genetic deciphering, and breeding utilization of dwarf lines in foxtail millet

DIAO Xian-Min1,*(), WANG Li-Wei1, ZHI Hui1, ZHANG Jun1, LI Shun-Guo2, CHENG Ru-Hong2   

  1. 1State Key Laboratory of Crop Gene Resources and Breeding / Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081, China
    2Institute of Millet Crops, Hebei Academy of Agricultural and Forestry Sciences, Shijiazhuang 050035, Hebei, China
  • Received:2023-07-20 Accepted:2023-09-28 Published:2024-02-12 Published online:2023-10-11
  • Contact: *E-mail: diaoxianmin@caas.cn
  • Supported by:
    National Natural Science Foundation of China(32241042);National Natural Science Foundation of China(31771807);National Key Research and Development Program of China(2021YFF1000103);China Agricultural Research System of MOF and MARA(CARS-06-04);Agricultural Science and Technology Innovation Program of the Chinese Academy of Agricultural Sciences

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摘要：

谷子是旱作生态农业的主栽作物和应对气候变化的战略储备作物, 也是新时期功能基因研究的模式作物。谷子品种的中矮秆化是产业发展的必然趋势。国内外目前已报道的谷子矮秆材料达70多个, 从形态上谷子矮秆种质可以分为叶穗直立的紧凑型和叶穗下披的常规型2类; 赤霉素(GA)敏感性测定发现4个材料对GA不敏感, 其他均是GA敏感型; 显隐性分析表明, 多数矮秆材料为隐性单基因控制, 但矮88株高为多基因控制。利用自然群体关联分析和双亲杂交分离群体合计发掘了109个控制株高的QTL, 精细定位了7个矮秆基因, 半显性矮秆材料84133的矮化基因SiDW1是唯一克隆并进行功能分析的矮化基因。对谷子中矮秆品种选育的梳理表明, 利用矮88和其衍生系为亲本已培育出139个中矮品种(系), 实现了谷子株高的显著矮化, 满足了机械化收获对抗倒伏降株高的要求。本文综述了国内外谷子株高矮化基因的研究进展, 梳理了已经定位和克隆的谷子矮秆基因, 讨论了谷子矮秆遗传和育种研究中的问题, 展望了未来发展方向。

关键词: 谷子, 株高, 矮秆, GA敏感性, 矮88

Abstract:

Foxtail millet (Setaria italica) is an important cereal crop in northern China’s arid and semiarid dry land agriculture, which has recently been proposed as a novel model for functional genomics. Breeding dwarf varieties is the development trend of foxtail millet industry. To date, more than 70 foxtail millet dwarf lines have been developed and reported worldwide. According to morphological characteristics, foxtail millet dwarf lines can be divided into two types: compact type with erect leaves and conventional type with droopy leaves. Gibberellins (GA) sensitivity assay indicated that four materials were not sensitive to GA and the others were sensitive. Genetic analyses detected that most of the dwarf phenotype lines were controlled by recessive genes, but the height phenotype of Ai 88 was controlled by multi-dwarf-gene. So far, 79 QTL related with plant height regulation were detected by natural population GWAS or linkage analysis. Among seven genes or QTL fine mapped in foxtail millet, the semi-dominant dwarfism gene SiD1 in 84113 was the only one cloned and functionally characterized. In the history of breeding dwarf foxtail millet variety, Ai 88 was a backbone line, which had been utilized to develop 139 cultivars with reducing plant height to meet the requirement of logging resistance and mechanized harvest. In this study, we reviewed systematically the research progress of dwarfing gene in foxtail millet, sorted out the dwarfing genes that had been located and cloned, discussed the problems in the research of dwarfing genetics and breeding, and prospected the future development in foxtail.

Key words: foxtail millet, plant height, dwarf, GA sensitivity, Ai 88

表1

谷子矮秆材料的显隐性"

编号
No.		 材料名称
Material name		 显隐性
Gene action		 编号
No.		 材料名称
Material name		 显隐性
Gene action		 编号
No.		 材料名称
Material name		 显隐性
Gene action
1 安矮15号 隐性 22 矮秆27-21 隐性 43 小野谷子 待定
An’ai 15 Recessive Aigan 27-21 Recessive Xiaoyeguzi Undetermined
2 红谷-1 隐性 23 志试897 隐性 44 创119-11 隐性
Honggu 1 Recessive Zhishi 8971 Recessive Chuang 119-11 Recessive
3 红谷-2 待定 24 矮秆2号 隐性 45 05-461 隐性
Honggu 2 Undetermined Aigan 2 Recessive Recessive
4 呼早谷-1 隐性 25 张矮10号 待定 46 延矮2号 隐性
Huzaogu 1 Recessive Zhang’ai 10 Undetermined Yan’ai 2 Recessive
5 赤矮9号 隐性 26 豫谷8号 待定 47 1066A 隐性
Chi’ai 9 Recessive Yugu 8 Undetermined Recessive
6 宽九 隐性 27 安矮13 隐性 48 麦谷1号 待定
Kuanjiu Recessive An’ai 13 Recessive Maigu 1 Undetermined
7 矮协1号 隐性 28 安矮3号 隐性 49 延096 隐性
Aixie 1 Recessive An’ai 3 Recessive Yan 096 Recessive
8 济矮5号 隐性 29 75原406 待定 50 郑矮2号 隐性
Ji’ai 5 Recessive 75 yuan 406 Undetermined Zheng’ai 2 Recessive
9 济矮6号 隐性 30 矮88号 隐性 51 7516 隐性
Ji’ai 6 Recessive Ai 88 Recessive Recessive
10 济矮9号 隐性 31 耧里莠 待定 52 84133 显性
Ji’ai 9 Recessive Loulixiu Undetermined Dominant
11 济矮10号 隐性 32 安矮8号 隐性 53 T22 隐性
Ji’ai 10 Recessive An’ai 8 Recessive Recessive
12 济矮11 隐性 33 安矮9号 待定 54 T25 隐性
Ji’ai 11 Recessive An’ai 9 Undetermined Recessive
13 济矮12 隐性 34 安矮17 隐性 55 T539 隐性
Ji’ai 12 Recessive An’ai 17 Recessive Recessive
14 晋汾矮4号 隐性 35 矮宁黄 待定 56 Co2 待定
Jinfen’ai 4 Recessive Aininghuang Undetermined Undetermined
15 晋汾矮5号 隐性 36 矮丰1号 隐性 57 sic24 显性
Jinfen’ai 5 Recessive Aifeng 1 Recessive Dominant
16 晋汾矮6号 隐性 37 毛毛斗芝麻良 隐性 58 sic25 显性
Jinfen’ai 6 Recessive Maomaodouzhimaliang Recessive Dominant
17 矮秆竹叶青4n 隐性 38 鲁热朗代斯 隐性 59 sic26 显性
Aiganzhuyeqing 4n Recessive Lurelangdaisi Recessive Dominant
18 延4直 隐性 39 延矮1号 隐性 60 吨谷 待定
Yan4zhi Recessive Yan’ai 1 Recessive Dungu Undetermined
19 延023 待定 40 大青秸 隐性 61 623C 待定
Yan 023 Undetermined Daqingju Recessive Undetermined
20 延029 待定 41 C193 待定 62 d93090 待定
Yan 029 Undetermined Undetermined Undetermined
21 延035 隐性 42 白米谷子 待定
Yan 035 Recessive Baimiguzi Undetermined

图1

谷子半显性矮秆基因DWARF1的图位克隆 A: 左侧为正常高秆豫谷1号, 中间为矮秆基因杂合的杂种F1, 右侧为纯合显性矮秆的84133, 说明其半显性特征。B: 84133的矮秆转录本D1-TT的超表达转基因功能验证。C: 张矮10和84133杂交后代剩余杂交体茎秆切片, 展示其细胞长度的缩短。"

表2

多环境中检测到的稳定的谷子株高QTL"

QTL 染色体
Chr.		 区间
Interval		 环境数目
Number of environments		 参考文献
Reference
qPH1-1 1 1402842-4203850 2 Feldman et al.[34]
qPH1-2 1 8565051-37364207 28 Feldman et al. [34]; Mauro-Herrera and Doust [35]; Zhang et al. [36]; He et al. [39]
qPH1-3 1 38288357-41090626 2 Feldman et al. [34]; He et al. [39]
qPH2-1 2 169785-8284383 23 Feldman et al. [34]
qPH2-2 2 26040234-32043521 2 Mauro-Herrera and Doust [35]; He et al. [39]
qPH2-3 2 32717606-47811870 87 Feldman et al. [34]; Mauro-Herrera and Doust [35]
qPH3-1 3 1634-4766697 14 Feldman et al. [34]; Mauro-Herrera and Doust [35]; He et al. [39]
qPH3-2 3 4766697-12575474 11 Feldman et al. [34]; He et al. [39]
qPH3-3 3 12821696-21388568 5 Feldman et al. [34]; He et al. [39]
qPH4-1 4 4242694-5211174 2 He et al. [39]
qPH4-2 4 6963002-30351975 5 Feldman et al. [34]
qPH4-3 4 31349528-34519109 2 Zhang et al. [36]; He et al. [39]
qPH4-4 4 36731228-37718913 3 Feldman et al. [34]
qPH5-1 5 5107998-30425775 15 Feldman et al. [34]; Mauro-Herrera and Doust [35]
qPH5-2 5 36050389-44465736 113 Feldman et al. [34]; Mauro-Herrera and Doust [35]; Zhang et al. [36]; He et al. [39]
qPH6-1 6 1449746-4132666 3 Feldman et al. [34]; He et al. [39]
qPH6-2 6 7309479-30048845 2 He et al. [39]
qPH6-3 6 32307267-35817005 4 Zhang et al. [36]; He et al. [39]
qPH7 7 12773905-33438039 42 Feldman et al. [34]; Mauro-Herrera and Doust [35]; Zhang et al. [36]; He et al. [39]
qPH8-1 8 596198-10222496 4 Feldman et al. [34]
qPH8-2 8 1742176-31662547 24 Fan et al. [14]; Feldman et al. [34]; He et al. [39]
qPH9-1 9 90984-10589367 3 Feldman et al. [34]
qPH9-2 9 3452335-11604590 63 Feldman et al. [34]; Mauro-Herrera and Doust [35]
qPH9-3 9 35624887-55796987 24 Feldman et al. [34]; Mauro-Herrera and Doust [35]; He et al. [39]

图2

矮88株高控制基因的遗传解析 A: 谷子矮秆骨干亲本矮88株型特征。B: 利用“矮88×辽谷1号” RIL群体在13个环境下定位的谷子控制株高的QTL位点, 合计有26个QTL, 其中1号染色体上qPH1.3在9个环境中检测到, 9号染色体上qPH9.2和qPH9.5均能在7个环境中检测到。"

表3

矮88育成的一级优质谷子品种株高及区试信息"

品种
Variety		 品质
Quality		 株高
Plant height (cm)		 区试
Regional trial		 产量较对照±%
Yeild compared with CK		 位次
Rank		 适宜区域
Farming area
冀谷19
Jigu 19		 一级
First grade		 113.70 2002-2003国家华北区试
2002-2003 National North China regional trial		 13.24 1 华北夏谷区
North China Summer Sowing region
冀谷40
Jigu 40		 一级
First grade		 119.90 2014-2015国家华北区试
2014-2015 National North China regional trial		 12.8 1 华北夏谷区
North China Summer Sowing region
2016-2017国家西北区试
2016-2017 National Northwest China regional trial		 7.7 6 西北春谷区
Northwest China Spring Sowing region
2017-2018吉林省谷子区试
2017-2018 Jilin regional trial		 2.28 4 东北春谷区
Northeast China Spring Sowing region
中谷1号
Zhonggu 1		 一级
First grade		 121.18 2011-2012国家华北区试
2011-2012 National North China regional trial		 10.43 1 华北夏谷区
North China Summer Sowing region
中谷2号
Zhonggu 2		 一级
First grade		 120.77 2014-2015国家华北区试
2014-2015 National North China regional trial		 13.99 2 华北夏谷区
North China Summer Sowing region
2016-2017国家东北区试
2016-2017 National Northeast China regional trial		 5.59 3 东北春谷区
Northeast China Spring Sowing region
长生13
Changsheng 13		 一级
First grade		 125.60 2015-2016国家西北区试
2015-2016 National Northwest China regional trial		 14.19 1 西北春谷中晚熟区
Northwest China Spring Sowing and Mid-late-maturing region
豫谷25
Yugu 25		 一级
First grade		 121.00 2014-2015国家华北区试
2014-2015 National North China regional trial		 14.51 1 华北夏谷区
North China Summer Sowing region
公矮2号
Gongai 2		 一级
First grade		 108.00 2002-2003吉林省谷子区试
2002-2003 Jilin regional trial		 13.55 1 东北春谷区
Northeast China Spring Sowing region
赤谷K3
Chigu K3		 一级
First grade		 109.00 2018-2019内蒙古谷子区试
2018-2019 Inner Mongolia regional trial		 4.94 2 西北春谷早熟区
Northwest China Spring Sowing and eraly-maturing region

图3

谷子矮秆品种公谷85及其大面积生产 A: 吉林省农业科学院作物资源研究所培育的矮秆品种公谷85, 以及紧邻的传统高秆品种。B: 吉林省农业科学院作物资源研究所培育的矮秆品种公谷88的生产大田表现。"

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