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作物学报

作物学报 ›› 2024, Vol. 50 ›› Issue (11): 2684-2698.doi: 10.3724/SP.J.1006.2024.41022

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

水旱条件下小麦持绿相关性状QTL定位

陈晨1(), 程宇坤1, 王伟1,2, 任毅1, 张海燕1, 陈慧波1, 耿洪伟1,*()   

  1. 1新疆农业大学农学院 / 新疆农业大学优质专用麦类作物工程技术研究中心, 新疆乌鲁木齐 830052
    2安阳工程学院计算机科学与信息工程系, 河南安阳 455000
  • 收稿日期:2024-03-14 接受日期:2024-06-20 出版日期:2024-11-12 网络出版日期:2024-07-18
  • 通讯作者: *耿洪伟, E-mail: hw-geng@163.com
  • 作者简介:E-mail: chenchen20210305@163.com
  • 基金资助:
    新疆自治区重点研发任务专项(2022B02001-3);新疆现代农业产业技术体系建设项目(小麦, XJARS-01)

QTL mapping of stay-green-related traits in wheat under drought condition

CHEN Chen1(), CHENG Yu-Kun1, WANG Wei1,2, REN Yi1, ZHANG Hai-Yan1, CHEN Hui-Bo1, GENG Hong-Wei1,*()   

  1. 1College of Agronomy, Xinjiang Agricultural University / Special High Quality Triticeae Crops Engineering and Technology Research Center, Xinjiang Agricultural University, Urumqi 830052, Xinjiang, China
    2Department of Computer Science and Information Engineering, Anyang Institute of Technology, Anyang 455000, Henan, China
  • Received:2024-03-14 Accepted:2024-06-20 Published:2024-11-12 Published online:2024-07-18
  • Contact: *E-mail: hw-geng@163.com
  • Supported by:
    Special Project of Key Research and Development Task of Xinjiang Autonomous Region(2022B02001-3);Xinjiang Agriculture Research System(Wheat, XJARS-01)

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摘要：

干旱是影响小麦生产中最重要的非生物胁迫。延长叶片持绿周期, 进而增加光合时间, 提高光合效率, 对小麦在旱胁迫下保证有机物积累, 最终达到小麦产量的稳定具有重要的意义。了解小麦旗叶持绿性保持的发育和遗传特征, 并探索与小麦持绿基因密切相关且不受环境影响的稳定的分子标记, 应用于育种可大大缩短抗旱育种周期。本研究以扬麦16和中麦895衍生的174个家系的DH群体为试验材料, 通过设置正常滴灌(NI)和干旱胁迫(DS)处理, 对2年2个水分处理下花后10 d、14 d、18 d、22 d、26 d和30 d旗叶绿色叶面积百分率(GLA)及叶绿素相对含量(SPAD值)进行表型测定, 运用Gompertz模型模拟出GLA (%)变化趋势。将达到最大衰老速度的时间(TMRS)、绿色叶面积持续期(GLAD)、平均衰老速率(ARS)、旗叶快速衰老开始的时间(T1)、旗叶快速衰老结束的时间(T2)和旗叶动态SPAD值等11个持绿相关性状进行QTL定位。 结果表明,水旱条件下, DH群体和亲本不同日龄下的GLA, 衰老特征参数和旗叶动态SPAD值均存在超亲分离现象, 并表现出一定差异。各家系和亲本不同日龄下的GLA均呈缓慢-快速-缓慢的动态下降趋势, 快速衰老主要集中在18 d、22 d和26 d。除ARS和GLA-10D呈负相关, 其余性状之间均呈现正相关。各性状遗传力范围在0.50~0.81之间。连锁分析在2个及以上环境中共鉴定到27个与小麦持绿相关的稳定位点, 其中11个与小麦衰老特征参数相关, 16个与旗叶动态SPAD值相关, 分布在1A、1B、4B、4D、5D、7B和7D染色体上, 分别解释表型变异3.86%~14.11%和2.99%~17.45%。在4B、4D和7B染色体上有1个调控T1和5个调控SPAD值的QTL在3个环境中被重复检测到, 分别解释8.97%~14.11%和6.85%~17.45%的表型变异。QTL聚合效应分析发现, 含有Rht-D1b和AA基因型的位点对SPAD-10D具有显著的增效作用和累加效应, 含有GG和TT基因型的位点对SPAD-18D具有显著的增效作用和累加效应。在4B、4D、7B和7D染色体上发现了4个效应明显的QTL簇, 位于4D染色体的QTL簇中存在不受水分环境影响的区段(18.80~28.58 Mb), 该区段内包含Rht-D1基因, 调控SPAD-0D和SPAD-10D的位点可能受到该基因的多效应影响。候选基因分析发现TraesCS4D01G054000、TraesCS1B01G434300和TraesCS7B01G010200等8个影响持绿相关性状的候选基因。以上研究结果为小麦持绿性分子标记辅助育种提供了理论依据。

关键词: 小麦, 持绿相关性状, QTL定位, 聚合效应, 候选基因

Abstract:

Drought is the primary abiotic stress that significantly affects wheat production. Extending the duration of leaf greenness, thereby increasing photosynthetic time and efficiency, is crucial for wheat in ensuring organic matter accumulation under drought stress and ultimately stabilizing wheat yield. Understanding the developmental and genetic characteristics of flag leaf greenness retention in wheat and identifying stable molecular markers closely associated with greenness-related genes, independent of environmental influences, can significantly accelerate the breeding process for drought resistance. In this study, we utilized a DH population consisting of 174 lines derived from Yangmai 16 and Zhongmai 895 as experimental materials. We conducted phenotypic evaluations of the percentage of green leaf area (GLA) and relative chlorophyll content (SPAD value) in flag leaves at 10 d, 14 d, 18 d, 22 d, 26 d, and 30 d post-flowering under two moisture environments: normal drip irrigation (NI) and drought stress (DS). We simulated the change in GLA using the Gompertz model and performed QTL mapping for eleven greenness-related traits, including time to maximum senescence rate (TMRS), green leaf area duration (GLAD), average senescence rate (ARS), time to the beginning of rapid senescence (T1), time to the end of rapid senescence (T2), and the dynamic SPAD value of the flag leaf. The results revealed a phenomenon of transgressive segregation in GLA, aging characteristic parameters, and dynamic SPAD values of the DH population and parents under both well-watered and drought conditions, demonstrating certain differences. A consistent trend of slow-rapid-slow decline in GLA was observed at different time points across all families and parents, with rapid senescence primarily occurring at 18 d, 22 d, and 26 d Except for the negative correlation between ARS and GLA-10D, positive correlations were observed among all other traits. The heritability of each trait ranged from 0.50 to 0.81. Linkage analysis identified a total of 27 stable loci associated with greenness retention in wheat across two or more environments. Among these, 11 loci were associated with wheat senescence characterization trait parameters, and 16 loci were associated with dynamic SPAD values of the flag leaf. These loci were distributed on chromosomes 1A, 1B, 4B, 4D, 5D, 7B, and 7D, explaining 3.86%-14.11% and 2.99%-17.45% of the phenotypic variance, respectively. One QTL regulating T1 and five SPAD values on chromosomes 4B, 4D, and 7B were consistently detected across the three environments, explaining 8.97%-14.11% and 6.85%-17.45% of the phenotypic variance, respectively. The results of QTL co-localization effect analysis revealed that loci containing Rht-D1b and AA genotypes exhibited significant enhancement and cumulative effects on SPAD-10D, while loci containing GG and TT genotypes showed significant enhancement and cumulative effects on SPAD-18D. Four QTL clusters with significant effects were identified on chromosomes 4B, 4D, 7B, and 7D. Notably, the segment of the QTL cluster on chromosome 4D (18.80-28.58 Mb) remained unaffected by water availability and contained the Rht-D1 gene. The loci regulating SPAD-0D and SPAD-10D might be influenced by the multiplicative effect of this gene. Candidate gene analysis identified eight candidate genes, such as TraesCS4D01G054000, TraesCS1B01G434300, and TraesCS7B01G010200, which are associated with greenness retention. These findings provide a theoretical foundation for molecular marker-assisted breeding aimed at improving greenness retention in wheat.

Key words: wheat, stay-green-related traits, QTL localization, convergent effect, candidate genes

附表1

DH群体及亲本旗叶绿色叶面积比例统计分析"

性状
Trait		 环境
Environment		 处理
Treatment		 亲本 Parents DH家系 DH Lines
扬麦16
Yangmai 16		 中麦895
Zhongmai 895		 最小值
Min.		 最大值
Max.		 平均值
Mean		 标准差
SD		 变异系数
CV		 偏度
Skew.		 峰度
Kurt.
GLA-10D E1 NI 10.00 10.00 9.10 10.00 9.94* 0.17 0.02 ‒2.92 8.46
DS 9.80 9.80 7.70 10.00 9.72 0.47 0.05 ‒2.45 6.13
E2 NI 10.00 10.00 9.10 10.00 9.98 0.09 0.01 ‒6.80 54.47
DS 9.80 10.00 8.80 10.00 9.97 0.14 0.01 ‒6.21 43.19
BLUE NI 10.00 10.00 9.20 10.00 9.96* 0.11 0.01 ‒3.68 17.89
DS 9.80 9.90 8.50 10.04 9.85 0.26 0.03 ‒2.55 7.19
GLA-14D E1 NI 9.60 10.00* 7.20 10.00 9.51* 0.58 0.06 ‒1.82 3.23
DS 7.60 8.80 4.00 10.00 8.63 1.30 0.15 ‒1.50 2.17
E2 NI 9.90 10.00 8.70 10.00 9.94 0.18 0.02 ‒4.68 23.94
DS 9.70 10.00* 8.30 10.00 9.93 0.22 0.02 ‒4.51 24.31
BLUE NI 9.75 10.00* 8.60 10.00 9.73* 0.31 0.03 ‒1.74 2.76
DS 8.65 9.40* 6.95 10.09 9.28 0.67 0.07 ‒1.47 2.01
GLA-18D E1 NI 6.30 7.90* 3.00 10.00 7.80* 1.58 0.20 ‒0.87 0.16
DS 2.90 4.60* 0.50 9.20 4.55 2.26 0.50 0.11 ‒0.95
E2 NI 9.40 10.00* 6.60 10.00 9.72* 0.49 0.05 ‒3.35 14.61
DS 8.80 10.00* 7.50 10.00 9.61 0.55 0.06 ‒1.95 3.78
BLUE NI 7.85 8.95 6.35 10.00 8.76* 0.82 0.09 ‒0.86 0.08
DS 5.85 7.30 4.30 9.60 7.08 1.18 0.17 0.00 ‒0.78
GLA-22D E1 NI 2.80 4.00* 0.50 9.60 4.42* 2.14 0.49 0.40 ‒0.83
DS 0.90 1.50* 0.00 7.20 2.21 1.67 0.75 0.58 ‒0.55
E2 NI 8.50 10.00* 6.10 10.00 9.08* 0.86 0.09 ‒1.06 0.84
DS 7.80 9.70* 4.00 9.90 7.83 1.31 0.17 ‒0.53 ‒0.44
BLUE NI 5.65 7.00 3.80 9.47 6.75* 1.19 0.18 0.14 ‒0.75
DS 4.35 5.60 2.00 8.30 5.02 1.19 0.24 0.01 ‒0.35
GLA-26D E1 NI 0.50 1.70* 0.00 7.60 1.93* 1.44 0.75 1.27 2.04
DS 0.00 0.90* 0.00 2.80 0.41 0.62 1.54 1.80 2.94
E2 NI 5.70 6.80* 1.50 8.80 5.56* 1.00 0.18 0.25 1.65
DS 4.30 5.50* 1.00 7.40 3.54 1.71 0.48 0.09 ‒0.97
BLUE NI 3.10 4.25 1.80 7.07 3.75* 0.99 0.26 0.73 0.52
DS 2.15 3.20 0.50 4.75 1.97 1.00 0.51 0.23 ‒0.69
GLA-30D E2 NI 1.10 2.60* 0.00 4.80 1.64* 1.08 0.66 0.87 0.45
DS 0.90 1.20 0.00 4.60 0.82 0.64 0.78 1.48 6.11

图1

2个环境下的2种水分环境中双亲及DH群体的曲线拟合结果 E1: 2021-2022玛纳斯; E2: 2022-2023三坪; 蓝线: 正常滴灌条件下预测值; 红线: 干旱胁迫条件下预测值; 蓝点: 正常滴灌条件下实测值; 红点: 干旱胁迫条件下实测值。"

表1

DH群体及亲本衰老参数值统计分析"

性状
Trait		 环境
Environ-
ment		 处理
Treatment		 亲本 Parents DH家系 DH line
扬麦16
Yangmai 16		 中麦895
Zhongmai 895		 最小值
Min.		 最大值
Max.		 平均值
Mean		 标准差
SD		 变异系数
CV		 偏度
Skew.		 峰度
Kurt.
TMRS (d) E1 NI 20.65 22.50* 13.93 31.27 22.71* 2.81 0.12 0.22 0.94
DS 16.84 18.82* 8.71 25.19 18.95 2.64 0.14 -0.42 0.74
E2 NI 27.69 29.02* 24.08 30.81 27.83* 1.16 0.04 0.37 0.36
DS 26.03 27.61* 22.29 30.92 25.93 1.55 0.06 -0.05 -0.29
BLUE NI 24.17 25.76 20.08 30.10 25.27* 1.67 0.07 0.28 0.48
DS 21.43 23.21 16.86 27.35 22.45 1.76 0.08 -0.22 0.11
GLAD (d) E1 NI 26.95 29.39* 19.74 40.53 28.76* 3.60 0.13 0.87 1.37
DS 23.38 25.33* 16.39 31.99 23.95 3.15 0.13 0.02 -0.80
E2 NI 32.28 33.39 27.28 39.02 32.32* 2.01 0.06 0.29 0.63
DS 31.14 31.67 25.20 37.11 30.07 2.16 0.07 0.10 -0.14
BLUE NI 29.61 31.39 25.67 38.44 30.55* 2.28 0.07 0.59 0.78
DS 27.26 28.50 21.33 33.00 27.01 2.09 0.08 -0.02 -0.29
ARS (% d-1) E1 NI -0.41 -0.38* -0.67 -0.27 -0.38* 0.06 0.15 -1.35 4.70
DS -0.52 -0.45* -1.30 -0.32 -0.47 0.11 0.24 -3.99 22.73
E2 NI -0.31 -0.31 -0.37 -0.26 -0.32* 0.02 0.06 -0.13 0.12
DS -0.32 -0.32 -0.41 -0.27 -0.34 0.03 0.07 -0.06 0.10
BLUE NI -0.36 -0.34 -0.52 -0.28 -0.35* 0.03 0.09 -1.08 4.13
DS -0.42 -0.39 -0.75 -0.32 -0.41 0.06 0.14 -3.16 15.50
T1 (d) E1 NI 18.93 20.61* 10.08 29.36 21.06* 2.99 0.14 -0.23 1.22
DS 15.05 17.04* 4.59 23.91 17.58 2.84 0.16 -0.85 2.20
E2 NI 26.43 27.82* 22.97 29.75 26.60* 1.06 0.04 0.35 1.25
DS 24.62 26.50* 20.94 29.23 24.79 1.49 0.06 -0.09 -0.39
BLUE NI 22.68 24.22 18.01 28.85 23.82* 1.70 0.07 0.01 0.57
DS 19.84 21.77 15.30 25.82 21.20 1.79 0.08 -0.40 0.43
T2 (d) E1 NI 22.38 24.39* 17.78 33.18 24.37* 2.81 0.12 0.59 0.94
DS 18.63 20.60* 12.82 26.46 20.32 2.60 0.13 -0.10 -0.31
E2 NI 28.95 30.22* 25.20 32.32 29.06* 1.33 0.05 0.33 -0.05
DS 27.43 28.72* 23.43 32.61 27.06 1.66 0.06 -0.01 -0.27
BLUE NI 25.66 27.30 22.15 32.39 26.71* 1.74 0.07 0.46 0.47
DS 23.03 24.66 18.41 28.90 23.70 1.78 0.08 -0.09 -0.09

附表2

DH群体及亲本SPAD值统计分析"

性状
Trait		 环境
Environment		 处理
Treatment		 亲本 Parents DH家系 DH Lines
扬麦16
Yangmai 16		 中麦895
Zhongmai 895		 最小值
Min.		 最大值
Max.		 平均值
Mean		 标准差
SD		 变异系数
CV		 偏度
Skew.		 峰度
Kurt.
SPAD-0D E1 NI 59.21 59.97 54.92 67.10 61.43 2.45 0.04 ‒0.45 0.26
DS 58.49 62.09* 54.99 66.86 61.19 2.31 0.04 ‒0.04 ‒0.05
E2 NI 59.66 60.42 53.36 64.38 59.33* 1.96 0.03 ‒0.09 0.32
DS 57.20 58.39 51.62 63.60 58.86 2.17 0.04 ‒0.58 0.51
BLUE NI 59.43 60.19 54.92 63.99 60.39 1.91 0.03 ‒0.60 0.20
DS 57.85 60.24* 53.30 63.94 60.02 1.94 0.03 ‒0.43 0.09
SPAD-10D E1 NI 59.63 59.49 54.69 68.00 62.48 2.63 0.04 ‒0.33 ‒0.11
DS 61.59 67.49* 55.17 70.06 62.72 2.76 0.04 0.03 ‒0.09
E2 NI 60.31 63.15* 55.60 65.71 60.89* 2.18 0.04 0.03 ‒0.49
DS 58.14 62.35* 55.70 65.29 60.26 2.00 0.03 ‒0.05 ‒0.37
BLUE NI 59.97 61.32 56.79 65.69 61.69* 2.04 0.03 ‒0.10 ‒0.78
DS 59.87 64.92* 56.73 67.52 61.48 2.02 0.03 0.19 0.06
SPAD-14D E1 NI 57.24 62.31* 52.54 70.12 61.76* 3.27 0.05 ‒0.18 0.11
DS 57.78 58.75 48.51 67.22 59.28 3.88 0.07 ‒0.39 ‒0.10
E2 NI 59.64 62.21 54.21 65.83 60.26 2.46 0.04 ‒0.19 ‒0.37
DS 59.90 63.29* 52.85 67.15 60.36 2.91 0.05 ‒0.26 ‒0.59
BLUE NI 58.44 62.26* 54.96 67.06 61.01 2.54 0.04 ‒0.14 ‒0.37
DS 58.84 61.02 51.90 65.19 59.82 2.79 0.05 ‒0.34 ‒0.47
SPAD-18D E1 NI 48.60 57.59* 40.34 67.61 56.29* 5.04 0.09 ‒0.59 0.41
DS 28.86 43.55* 7.68 65.41 48.49 9.49 0.20 ‒0.81 1.33
E2 NI 57.90 59.93* 50.93 65.89 59.03* 2.63 0.04 ‒0.10 ‒0.35
DS 56.68 60.49 50.61 64.82 58.18 2.55 0.04 ‒0.27 ‒0.22
BLUE NI 53.25 58.76* 48.66 64.23 57.68* 3.34 0.06 ‒0.38 ‒0.34
DS 42.77 52.02 35.59 63.45 53.38 5.22 0.10 ‒0.54 0.20
SPAD-22D E1 NI 32.36 43.49* 16.52 65.24 44.34* 9.65 0.22 ‒0.43 ‒0.09
DS 8.26 15.36* 4.30 54.44 25.04 11.43 0.46 0.26 ‒0.75
E2 NI 49.19 54.03* 37.76 63.07 53.53* 4.82 0.09 ‒0.66 0.37
DS 38.04 52.33* 11.29 61.09 45.88 11.88 0.26 ‒0.98 0.38
BLUE NI 40.77 48.76 30.56 62.94 48.97* 6.26 0.13 ‒0.39 ‒0.07
DS 23.15 33.85 9.40 54.56 35.46 9.43 0.27 ‒0.34 ‒0.11
SPAD-26D E1 NI 6.45 11.92* 5.52 47.54 16.27* 7.63 0.47 1.36 3.13
DS 3.84 5.38* 2.97 25.72 8.30 3.39 0.41 1.57 4.09
E2 NI 33.14 43.16* 17.11 57.05 33.86* 8.58 0.25 0.32 ‒0.25
DS 27.06 35.05* 6.24 56.54 18.99 13.07 0.69 1.15 0.27
BLUE NI 19.80 27.54 12.74 48.81 25.10* 6.55 0.26 0.66 1.02
DS 15.45 20.22 3.12 35.17 13.63 7.21 0.53 1.12 0.33
SPAD-30D E2 NI 6.20 11.81* 3.27 44.51 11.29* 6.44 0.57 1.94 5.22
DS 4.27 8.09 1.90 31.35 7.61 3.95 0.52 3.10 14.24

图2

不同水分环境中小麦持绿相关性状相关性分析 *表示P < 0.05水平差异显著; **表示P < 0.01水平差异显著; ***表示P < 0.001水平差异显著; 左下角为正常滴灌, 右上角为干旱胁迫。"

附表3

持绿相关性状方差分析及遗传力"

性状
Trait		 均方值Mean Square
基因型
Genotype (G)		 环境
Environment (E)		 G×E互作
G×E		 H²
%GLA-10D 0.22*** 5.07*** 0.11*** 0.60
%GLA-14D 1.56*** 131.19*** 0.86*** 0.60
%GLA-18D 6.49*** 2017.33*** 3.15*** 0.60
%GLA-22D 8.65*** 3441.3*** 3.51*** 0.66
%GLA-26D 5.39*** 1698.58*** 2.36*** 0.64
%GLA-30D 2.04*** 115.07*** 1.09*** 0.57
SPAD-0D 25.05*** 575.43*** 4.81 0.81
SPAD-10D 26.64*** 494.54*** 6.52*** 0.78
SPAD-14D 47.01*** 353.41*** 10.88*** 0.79
SPAD-18D 126.52*** 7855.07*** 40.73*** 0.74
SPAD-22D 372.19*** 51022.18*** 129.11*** 0.73
SPAD-26D 266.73*** 39912.29*** 116.02*** 0.68
SPAD-30D 62.44*** 2345.09*** 50.92*** 0.50
TMRS 18.13*** 5199.24*** 5.87*** 0.72
GLAD 24.98*** 4316.17*** 12.16*** 0.57
ARS 0.00*** 1.58*** 0.01*** 0.56
T1 18.79*** 5521.38*** 6.65*** 0.69
T2 18.56*** 4912.08*** 6.16*** 0.71

表2

不同水分环境中衰老特征参数和SPAD值稳定QTL位点"

位点
QTL		 环境
Environment		 处理
Treatment		 左标记
Left marker		 右标记
Right marker		 置信区间
Confidence interval (cM)		 位置
Position (Mb)		 LOD值
LOD value		 表型贡献率
PVE (%)		 加性效应
Additive effect
QTMRS.xjau-4B E1/BLUE NI AX-110194463 AX-111651924 120.5‒122.0 7.21‒19.03 4.72‒6.06 10.34‒11.46 0.53‒1.04
QTMRS.xjau-5D E1/BLUE NI AX-111162962 AX-110717870 163.5‒168.5 459.07‒465.11 3.68‒4.28 7.26‒7.68 0.46‒0.83
QTMRS.xjau-4D E2/BLUE DS AX-108944764 AX-94558069 52.5‒53.5 15.91‒16.59 7.35‒8.85 11.46‒11.93 ‒0.65‒-0.59
QTMRS.xjau-7D E2/BLUE DS AX-109167420 AX-111918348 109.5‒112.5 72.30‒77.23 3.17‒3.62 3.86‒5.31 0.37‒0.41
QGLAD.xjau-4D E2/BLUE DS AX-108944764 Rht-D1 52.5‒55.5 15.91‒18.80 6.39‒10.61 6.97‒10.72 ‒0.90‒-0.72
QT1.xjau-4B E1/BLUE NI AX-86170717 AX-111651924 120.5‒122.0 7.21‒22.87 5.01‒6.11 10.13‒11.68 0.55‒1.10
QT1.xjau-5D E1/BLUE NI AX-111162962 AX-110035093 161.5‒168.5 459.07‒464.13 4.75‒4.97 8.16‒9.32 0.53‒0.92
QT1.xjau-4D E1/E2/BLUE DS AX-95659047 AX-94558069 49.5‒53.5 12.77‒16.59 7.47‒9.55 8.97‒14.11 -0.99‒-0.55
QT1.xjau-7D E2/BLUE DS AX-109167420 AX-111918348 109.5‒112.5 72.30‒77.23 5.01‒5.51 6.56‒7.00 0.47‒0.52
QT2.xjau-4D E2/BLUE DS AX-108944764 AX-94558069 52.5‒53.5 15.91‒16.59 9.38‒9.41 6.34‒12.58 -0.72‒-0.66
QT2.xjau-7B E1/BLUE DS AX-111478329 AX-108883282 1.5‒2.5 1.21‒4.47 3.11‒5.80 5.94‒7.74 -0.65‒-0.56
QSPAD-0D.xjau-4D.1 E2/BLUE NI Rht-D1 AX-109445215 56.5‒60.5 18.80‒28.58 4.24‒5.10 3.64‒11.65 -0.73‒-0.59
QSPAD-0D.xjau-4D.2 E2/BLUE DS AX-89421921 Rht-D1 54.5‒55.5 18.80‒19.29 5.35‒7.80 9.37‒12.20 -0.90‒-0.63
QSPAD-10D.xjau-4D.1 E1/BLUE NI AX-89421921 Rht-D1 54.5‒55.5 18.80‒19.29 6.15‒10.69 9.06‒14.86 -0.95‒-0.91
QSPAD-10D.xjau-1A E1/BLUE DS AX-94629887 AX-109406391 0.0‒0.5 8.64‒20.09 4.30‒5.82 4.86‒8.37 0.56‒0.94
QSPAD-10D.xjau-4B E1/E2/BLUE DS AX-95685381 AX-94912823 43.5‒46.5 171.62‒172.72 5.59‒10.27 8.79‒14.06 0.86‒0.89
QSPAD-10D.xjau-4D.2 E1/BLUE DS AX-89421921 Rht-D1 54.5‒55.5 18.80‒19.29 7.13‒13.05 10.72‒17.18 -0.98‒-0.98
QSPAD-14D.xjau-4B E2/BLUE NI AX-95685381 AX-108888649 43.5‒44.5 171.62‒263.38 8.39‒11.51 12.81‒17.04 1.22‒1.39
QSPAD-14D.xjau-4D E2/BLUE NI AX-111071805 AX-111662392 61.5‒63.5 27.15‒33.06 6.03‒10.24 9.64‒15.73 -1.12‒-0.80
QSPAD-18D.xjau-4B E1/E2/BLUE NI AX-95685381 AX-108888649 43.5‒44.5 171.62‒263.38 6.27‒8.18 9.75‒13.49 1.13‒1.69
QSPAD-18D.xjau-4D E1/E2/BLUE NI AX-95659047 AX-108944764 47.5‒52.5 12.77‒15.91 6.75‒13.27 10.92‒17.45 -2.18‒-1.01
QSPAD-18D.xjau-7B E1/BLUE DS AX-111581943 AX-110629026 0.0‒1.5 6.07‒7.00 3.33‒4.66 2.99‒7.05 ‒2.36‒‒0.90
QSPAD-22D.xjau-4D E1/E2/BLUE DS AX-108944764 AX-109478820 52.5‒57.5 15.91‒30.66 5.26‒9.05 7.21‒12.81 -4.45‒-3.75
QSPAD-22D.xjau-7D E2/BLUE DS AX-110941147 AX-111918348 111.5‒112.5 76.62‒77.23 4.50‒6.25 5.02‒5.94 2.57‒3.78
QSPAD-26D.xjau-7B E1/E2/BLUE NI AX-111581943 AX-110629026 0.0‒1.5 6.07‒7.00 3.42‒8.74 6.85‒13.27 -2.70‒-2.50
QSPAD-26D.xjau-1B E2/BLUE DS AX-110534755 AX-111590092 128.5‒129.5 653.94‒665.05 2.90‒4.64 6.60‒10.30 -3.05‒-2.39
QSPAD-26D.xjau-4D E2/BLUE DS AX-89421921 Rht-D1 54.5‒57.5 18.80‒19.29 4.21‒5.48 8.06‒9.76 -3.41‒-2.35

图3

DH群体中衰老相关性状QTL位点在染色体上的位置"

图4

稳定QTL位点间的综合效应分析"

表3

筛选获得的候选基因"

位点
QTL		 左标记
Left marker		 右标记
Right marker		 物理位置
Position (Mb)		 候选基因
Gene ID		 功能注释
Functional annotation
QTMRS.xjau-4B AX-110194463 AX-111651924 15.18 TraesCS4B01G021200 基本螺旋-环-螺旋(BHLH) DNA结合超家族蛋白
Basic helix-loop-helix (BHLH) DNA-binding superfamily protein
QTMRS.xjau-4B AX-110194463 AX-111651924 13.30 TraesCS4B01G018400 衰老相关家族蛋白, 推测(DUF581)
Senescence-associated family protein, putative (DUF581)
QT1.xjau-4D AX-95659047 AX-94558069 16.30 TraesCS4D01G037000 半胱氨酸蛋白酶, 推测
Cysteine protease, putative
QSPAD-10D.xjau-4B AX-95685381 AX-94912823 171.62 TraesCS4B01G131400 蛋白质锌诱导的促进因子1
Protein zinc induced facilitator-like 1
QSPAD-14D.xjau-4D AX-111071805 AX-111662392 30.00 TraesCS4D01G054000 GRAS转录因子
GRAS transcription factor
QSPAD-18D.xjau-4D AX-95659047 AX-108944764 15.52 TraesCS4D01G033900 锌指家族蛋白
Zinc finger family protein
QSPAD-26D.xjau-1B AX-110534755 AX-111590092 658.91 TraesCS1B01G434300 WRKY家族转录因子
WRKY family transcription factor
QSPAD-26D.xjau-7B AX-111581943 AX-110629026 6.40 TraesCS7B01G010200 叶绿体POR1的蛋白伴侣样蛋白Protein chaperone-like protein of POR1, chloroplastic
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