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作物学报

作物学报 ›› 2025, Vol. 51 ›› Issue (9): 2266-2284.doi: 10.3724/SP.J.1006.2025.51028

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

小麦叶片气孔相关性状全基因组关联分析

李璐琪1,2(), 程宇坤1,2, 白斌3, 雷斌4, 耿洪伟1,2,*()   

  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丝绸之路经济带作物生物育种联合实验室, 新疆乌鲁木齐 830052
    3甘肃省农业科学院小麦研究所, 甘肃兰州 730070
    4新疆农业科学院, 新疆乌鲁木齐 830091
  • 收稿日期:2025-03-11 接受日期:2025-06-01 出版日期:2025-09-12 网络出版日期:2025-06-13
  • 通讯作者: *耿洪伟, E-mail: hw-geng@163.com
  • 作者简介:E-mail: 1455812032@qq.com
  • 基金资助:
    本研究由新疆维吾尔自治区科学技术厅自然科学基金杰出青年科学基金项目(2022D01E46);新疆维吾尔自治区重点研发计划(2023B02006);新疆维吾尔自治区青年科技拔尖人才专项(2022TSYCCX0079)

Genome-wide association analysis of stomatal-related traits in wheat leaves

LI Lu-Qi1,2(), CHENG Yu-Kun1,2, BAI Bin3, LEI Bin4, GENG Hong-Wei1,2,*()   

  1. 1College of Agronomy / Special High Quality Triticeae Crops Engineering and Technology Research Center, Xinjiang Agricultural University, Urumqi 830052, Xinjiang, China
    2Joint International Research Laboratory of Crop Biological Breeding along the Silk Road Economic Belt, Urumqi 830052, Xinjiang, China
    3Wheat Research Institute, Gansu Academy of Agricultural Sciences, Lanzhou 730070, Gansu, China
    4Xinjiang Academy of Agricultural Sciences, Urumqi 830091, Xinjiang, China
  • Received:2025-03-11 Accepted:2025-06-01 Published:2025-09-12 Published online:2025-06-13
  • Contact: *E-mail: hw-geng@163.com
  • Supported by:
    Xinjiang Uygur Autonomous Region Department of Science and Technology, the National Natural Science Fund for Distinguished Young Scholars(2022D01E46);Xinjiang Uygur Autonomous Region Key Research and Development Program(2023B02006);Xinjiang Uygur Autonomous Region Youth Science and Technology Talent Program(2022TSYCCX0079)

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摘要：

小麦气孔是调节光合作用和蒸腾作用的微观孔隙, 对产量有着关键作用, 了解小麦气孔性状的遗传机制, 挖掘关于小麦气孔相关性状的候选基因对于提高小麦产量具有重要意义。本研究以276份冬小麦品种(系)为研究对象, 在正常灌溉和干旱胁迫2个处理下, 对小麦抽穗期和灌浆期的气孔密度、气孔平均面积、气孔所占比等叶片气孔相关性状进行测定, 并结合90K小麦芯片, 对上述3个性状进行全基因组关联分析。研究结果显示, 在不同水分处理条件下, 小麦气孔相关性状在2个时期均表现出较大的表型变异, 变异系数介于0.06至0.28之间。全基因组关联结果显示, 共检测到88个位于除4D的其他20条染色体上与气孔相关性状显著关联的位点(P < 0.001), 其中与气孔平均面积显著相关的稳定遗传位点有4个, 分布于1B, 3A, 6A染色体上, 可解释遗传变异为2.78%~6.55%, 均在正常灌溉下的灌浆期检测到。在抽穗期和灌浆期同时检测到2个与气孔密度显著关联的位点, 其中在正常灌溉下6A染色体上检测到的位点Ex_c69429_328, 可解释表型变异为2.31%~3.06%; 在干旱胁迫下4A染色体上检测到的位点BS00064423_51, 可解释表型变异的4.40%~6.09%。在1A、1B、3A、4A、5A、6A、6D染色体上检测到8个“一因多效”遗传位点, 可解释表型变异的1.25%~7.31%。对贡献率大于5.00%且在2个环境或2个性状以上检测到位点进行单倍型分析, 发现与气孔平均面积和气孔所占比显著相关的wsnp_Ex_rep_c69627_68580121(R2=6.47%)位点存在Hap1、Hap2两个单倍型, 在276份品种(系)中含有Hap1(频率为81.20%)单倍型品种(系)的气孔平均面积显著小于含有Hap2(18.80%)单倍型品种(系)的气孔平均面积(P < 0.05), 不同单倍型在不同麦区的占比不同, 其中单倍型Hap1在西南冬麦区出现的频率最高, 单倍型Hap2则在北部冬麦区出现的频率最高。对2个环境检测到的及“一因多效”的位点进行候选基因的挖掘, 共筛选出9个与小麦气孔性状相关的候选基因, 这些基因在气孔发育、光合作用和抗逆性等方面具有潜在功能, 可作为进一步研究气孔相关性状的重要基因。

关键词: 小麦, 气孔, 全基因组关联分析, 单倍型, 候选基因

Abstract:

Wheat stomata are microscopic pores that regulate photosynthesis and transpiration, playing a critical role in determining yield. Understanding the genetic mechanisms underlying stomatal traits and identifying candidate genes associated with these characteristics are essential for improving wheat productivity. In this study, 276 winter wheat varieties (lines) were evaluated under two irrigation treatments—normal irrigation and drought stress. Stomatal traits, including stomatal density, average stomatal area, and stomatal pore proportion, were measured at both the heading and filling stages. A genome-wide association study (GWAS) was conducted using a 90K wheat SNP chip to analyze these traits. The results revealed significant phenotypic variation in stomatal traits across the two irrigation regimes and developmental stages, with coefficients of variation ranging from 0.06 to 0.28. GWAS identified 88 loci significantly associated with stomatal traits across 20 chromosomes, excluding chromosome 4D (P < 0.001). Among these, four stable genetic loci related to average stomatal area were identified on chromosomes 1B, 3A, and 6A, explaining 2.78% to 6.55% of the phenotypic variance. These loci were all detected during the filling stage under normal irrigation. Two loci associated with stomatal density were identified at both the heading and filling stages: Ex_c69429_328 on chromosome 6A under normal irrigation explained 2.31% to 3.06% of the variation, while BS00064423_51 on chromosome 4A under drought stress accounted for 4.40% to 6.09%. Additionally, eight loci with pleiotropic effects were identified on chromosomes 1A, 1B, 3A, 4A, 5A, 6A, and 6D, explaining 1.25% to 7.31% of the phenotypic variation. Haplotype analysis was performed on loci with a contribution rate greater than 5.00% that were detected in at least two environments or associated with two or more traits. Notably, the wsnp-Ex_rep_c69627_68580121 locus (R2 = 6.47%), significantly associated with both average stomatal area and stomatal pore proportion, exhibited two haplotypes—Hap1 and Hap2. Among the 276 wheat lines, those carrying Hap1 (frequency: 81.20%) had a significantly smaller average stomatal area than those with Hap2 (frequency: 18.80%) (P < 0.05). Haplotype distribution varied across wheat-growing regions, with Hap1 most prevalent in the southwest winter wheat region, while Hap2 was more common in northern regions. A total of nine candidate genes associated with stomatal traits were identified based on loci detected in multiple environments and those with pleiotropic effects. These genes are likely involved in stomatal development, photosynthesis, and stress responses, offering promising targets for further research and genetic improvement of wheat stomatal traits.

Key words: wheat, stomata, genome-wide association analysis, haplotype, candidate genes

表1

不同环境下群体表型变异统计分析"

性状
Trait		 时期
GS		 环境
Environment		 处理
Treatment		 最小值
Min.		 最大值
Max.		 均值
Mean		 标准差
SD		 变异系数
CV		 偏度
Skewness		 峰度
Kurtosis
气孔密度
SD (No. mm-2)		 抽穗期
HS		 E1 NI 47.71 79.17 60.63 5.21 0.09 0.56 0.60
DS 44.79 90.42 60.35 7.62 0.13 0.97 1.20
E2 NI 50.42 94.38 68.82 a 5.87 0.09 0.05 1.00
DS 44.79 90.42 71.96 b 5.64 0.08 -0.35 2.18
BLUE NI 50.42 82.61 64.72 a 4.37 0.07 0.27 0.78
DS 44.79 90.42 66.15 b 5.07 0.08 0.42 2.72
灌浆期
FS		 E1 NI 47.92 78.33 60.90 a 5.63 0.09 0.15 -0.37
DS 52.50 100.83 69.63 b 7.17 0.10 0.55 0.95
E2 NI 54.17 92.08 71.82 a 5.48 0.08 0.27 1.34
DS 52.50 100.83 75.78 b 6.21 0.08 0.40 1.29
BLUE NI 54.17 79.90 66.36 a 4.22 0.06 0.27 0.63
DS 52.50 100.83 72.70 b 5.40 0.07 0.58 3.05
气孔平均面积
ASA (μm2)		 抽穗期
HS		 E1 NI 1258.58 4179.35 2130.16 416.53 0.20 1.44 4.40
DS 1230.41 4940.16 2061.60 529.40 0.26 2.27 7.83
E2 NI 1276.98 4314.03 1965.63 a 392.69 0.20 2.02 6.94
DS 1113.76 4643.17 2063.41 b 450.51 0.22 1.79 5.37
BLUE NI 1393.89 3573.64 2047.90 285.90 0.14 1.13 3.07
DS 1366.31 4395.09 2078.81 432.98 0.21 2.38 8.15
灌浆期
FS		 E1 NI 1126.62 4687.50 2144.75 a 564.62 0.26 1.13 1.94
DS 1454.86 4313.81 2319.45 b 419.54 0.18 0.87 1.55
E2 NI 1561.86 4142.42 2333.74 439.86 0.19 0.87 1.06
DS 1309.23 4963.83 2357.83 566.99 0.24 1.29 2.69
BLUE NI 1452.04 4142.42 2239.25 379.54 0.17 1.03 2.49
DS 1536.35 3246.55 2294.21 316.03 0.14 0.46 0.04
气孔所占比
SPP (%)		 抽穗期
HS		 E1 NI 8.37 23.47 12.77 2.46 0.19 1.31 2.94
DS 7.85 23.40 12.46 2.81 0.23 1.25 1.67
E2 NI 7.73 26.21 13.47 a 2.86 0.21 1.42 4.04
DS 7.80 26.08 14.56 b 2.74 0.19 1.24 2.69
BLUE NI 9.30 19.96 13.12 a 1.87 0.14 0.82 1.17
DS 8.91 22.33 13.51 b 2.00 0.15 0.98 1.59
灌浆期
FS		 E1 NI 6.61 25.75 12.97 a 3.62 0.28 0.87 0.69
DS 9.93 25.35 15.73 b 2.69 0.17 0.50 0.34
E2 NI 9.62 23.40 16.35 2.66 0.16 0.18 -0.23
DS 10.18 25.24 16.78 2.63 0.16 0.50 0.31
BLUE NI 9.48 22.41 14.66 a 2.34 0.16 0.54 0.49
DS 12.03 23.53 16.25 b 1.98 0.12 0.35 0.21

表2

不同处理下群体表型方差分析"

性状
Trait		 时期
Growth stage		 处理
Treatment		 均方值 Mean square 广义遗传力H2
基因型
Genotype (G)		 环境
Environment (E)		 基因型×环境
G×E
气孔密度SD 抽穗期HS NI 76.29*** 18,518.82*** 46.79*** 0.53
DS 102.79*** 37,173.04*** 76.72*** 0.49
灌浆期FS NI 71.08*** 32,919.59*** 52.46*** 0.45
DS 116.64*** 10,469.03*** 63.18*** 0.57
气孔平均面积ASA 抽穗期HS NI 326,951.03*** 7,471,329.00*** 328,452.19*** 0.28
DS 822,431.25*** 399,662.53*** 681,708.94*** 0.20
灌浆期FS NI 576,210.81*** 9,858,272.00*** 448,330.50*** 0.43
DS 399,508.41*** 703,669.13*** 312,814.47*** 0.34
气孔所占比SPP 抽穗期HS NI 13.99*** 135.53*** 14.39*** 0.27
DS 15.96*** 1217.26*** 14.85*** 0.31
灌浆期FS NI 21.94*** 3161.25*** 18.44*** 0.45
DS 15.75*** 299.87*** 12.64*** 0.39

附表1

基于BLUE值筛选的小麦品种(系)在不同麦区的分布"

性状
Trait		 时期
GS		 处理
Treatment		 国内品种(系)Domestic varieties (lines) 国外品种
Abroad varieties
黄淮冬麦区
Huang-Huai winter wheat region		 北部冬麦区
Northern winter wheat region		 长江中下游冬麦区
Middle and lower reaches of the Yangtze River winter wheat region		 西南冬麦区
Southwest winter wheat region
气孔密度
SD
(No. mm-2)		 抽穗期HS NI 豫麦34, 豫麦18, 山农20, 豫麦35
Yumai 34, Yumai 18, Shannong 20, Yumai 35		 洋小麦, 中麦415, 石4185, 观35, CA1133, 北京841, 新麦37
Yangxiaomai, Zhongmai 415, Shi 4185, Guan 35, CA1133, Beijing 841, Xinmai 37		 华2459, 淮麦18, 扬麦9号, 淮麦21
Hua 2459, Huaimai 18, Yangmai 9, Huaimai 21		 绵农4号
Miannong 4		 Nidera Baguette 10, STARSHINA, Lovrin10, Abodanza
DS 豫麦34, 泰山5号, 济南13, 周麦11
Yumai 34, Taishan 5, Jinan13, Zhoumai 11		 洋小麦, 北京841, 衡观33, CA1133, CA0958
Yangxiaomai, Beijing 841, Hengguan 33, CA1133, CA0958		 连麦2号, 皖麦50, 皖麦29
Lianmai 2, Wanmai 50, Wanmai 29		 绵农4号, 川麦49, 川育23, 川农23, 绵阳26
Miannong 4, Chuanmai 49, Chuanyu 23, Chuannong 23, Mianyang 26		 Lampo, Abodanza, TX03A0148
灌浆期FS NI 豫麦34, 85中33, 鲁麦9号, 豫麦21, 丰产3号, 兰考906
Yumai 34, 85 Zhong 33, Lumai 9, Yumai21, Fengchan 3, Lankao 906		 丰抗2号, CA1090, 北京841, 中麦415, 石4185
Fengkang 2, CA1090, Beijing 841, Zhongmai 415, Shi 4185		 川麦49
Chuanmai 49		 C39, MV05-08, MV LAURA, Norin 67, Klein Jabal 1, Nidera Baguette 20, F498U1-1021 / BOEMA, Festin
DS 碧蚂1号, 丰产3号
Nima 1, Fengchan 3		 洋小麦, CA0958, 观35, 北京841, 农大211
Yangxiaomai, CA0958, Guan 35, Beijing 841, Nongda211		 镇麦168, 皖麦38, 淮麦18, 鄂麦11
Zhenmai 168,Wanmai 38, Huaimai 18, Emai11		 绵农4号, 川麦49, 川农23, 川育23
Miannong 4, Chuanmai 49, Chuannong 23,Chuanyu 23		 Festin, Abodanza, Aca 801, Lovrin13, KNIISH 46
气孔平均面积ASA (μm2) 抽穗期HS NI 85中33, 陕715, 淄选2号, 中麦895, 周麦16, 矮丰3号, 豫麦50, 豫麦34, 豫麦18
85 Zhong 33, Shaan 715, Zixuan 2, Zhongmai 895, Zhoumai 16, Aifeng 3,Yumai 50, Yumai 34, Yumai 18		 科衡6654
Keheng 6654
 皖麦29
Wanmai 29		 川麦49, 科成麦1号, 绵阳26
Chuanmai 49, Kechengmai 1, Mianyang 26		 C39, Fr03711, Abodanza, Kitanokaori, MV LAURA, Aca 601
DS 豫麦7号, 周麦13, 碧蚂1号, 泰山5号, 豫麦34, 85中33
Yumai 7, Zhoumai 13, Bima1, Taishan 5, Yumai 34, 85 Zhong 33		 科衡6654, 邯6172, 洋小麦
Keheng 6654, Han 6172, Yangxiaomai		 扬麦10号, 皖麦29, 皖麦50
Yangmai 10, Wanmai 29, Wanmai 50		 川麦49, 绵农4号, 绵阳26
Chuanmai 49, Miannong 4, Mianyang 26		 BRUTA, Darius, Abodanza, Norin 61, Aztec
灌浆期FS NI 豫麦35, 中892, 淄麦12, 新麦19, 陕农78-59
Yumai 35, Zhong 892, Zimai 12, Xinmai 19, Shaannong 78-59		 衡观33, 金禾9123, 石家庄15
Hengguan 33, Jinhe 9123, Shijiazhuang 15		 扬麦10号, 鄂麦14, 鄂麦21, 扬麦9号, 宁麦13
Yangmai10, Emai 14, Emai 21, Yangmai 9, Ningmai 13		 川麦49, 川麦107, 川麦47, 科成麦1号, 绵阳26
Chuanmai 49, Chuanmai 107, Chuanmai 47, Kechengmai 1, Mianyang 26		 NSA09-3645, Festin
DS PH82-2, 临旱2号, 西农291, 85中33, 郑麦366, 丰产3号
PH82-2, Linhan 2, Xinong 291, 85 Zhong 33, Zhengmai 366, Fengchan 3		 邯6172, 宁冬10
Han 6172, Ningdong 10		 扬麦9号, 鄂麦14, 皖麦29, 皖麦19
Yangmai 9, Emai 14, Wanmai 29, Wanmai 19		 绵农4号, 川麦49, 川麦107, 绵阳26
Miannong 4, Chuanmai 49, Chuanmai 107, Mianyang 26		 Abodanza, Klein Jabal 1, Aca 801, Kitanokaori
气孔所占比SPP (%) 抽穗期HS NI 中麦895, 周麦16, 临抗12, 郑引1号, 85中33, 中麦871, 豫麦50, 临麦2号, 周麦18, 济南17, 矮抗58
Zhongmai 895, Zhoumai 16, Linkang 12, Zhengyin 1, 85 Zhong 33, Zhongmai 871, Yumai 50, Linmai 2,Zhoumai 18, Jinan 17, Aikang 58		 CA0958 皖麦29, 皖麦33, 皖麦50
Wanmai 29, Wanmai 33, Wanmai 50
 川麦49, 科成麦1号, 内麦9号
Chuanmai 49, Kechengmai 1, Neimai 9		 Aca 601, Fr03711
DS 陕优225, 周麦13, 临旱2号, 周麦18, 郑引1号, 矮抗58, 85中33, 洛麦21, 碧蚂1号, 西农979-005
Shaanyou 225, Zhoumai 13, Linhan 2, Zhoumai 18, Zhengyin 1, Aikang 58, 85 Zhong 33, Luomai 21, Bima1, Xinong 979-005		 京冬8号, 科衡6654, 宁冬10, 邯6172
Jingdong 8, Keheng 6654, Ningdong 10, Han 6172		 扬麦10号
Yangmai 10		 Aztec, Salmone, BRUTA, Darius, DONSKI-93
灌浆期FS NI 陕优225, 豫麦57, 中892, 济麦20, 豫麦35, 淄麦12, 豫麦49, 陕农78-59, 周麦28
Shaanyou 225, Yumai 57, Zhong 892, Jimai 20, Yumai 35, Zimai 12, Yumai 49, Shaannong 78-59, Zhoumai 28		 高优503, CA0816(白)
Gaoyou 503, CA0816 (white)		 扬麦10号, 鄂麦14, 皖麦52, 宁麦13
Yangmai 10, Emai 14, Wanmai 52, Ningmai 13		 川麦47, 川麦107, 科成麦1号
Chuanmai 47, Chuanmai 107, Kechengmai 1		 NSA09-3645, HK1/6/NVSR3/
5/BEZ/TVR/5/CFN/BEZ//
SU92/CI13645/3NAI60
DS PH82-2, 中麦895, 郑麦366, 洛麦21, 周麦18, 周麦16, 临旱2号, 豫麦57, 中麦871, 新麦9号, 11CA40
PH82-2, Zhongmai 895, Zhengmai 366, Luomai 21, Zhoumai 18, Zhoumai 16, Linhan 2, Yumai 57, Zhongmai 871, Xinmai 9, 11CA40		 皖麦19, 鄂麦14, 皖麦52, 皖麦29, 扬麦9号
Wanmai 19, Emai 14, Wanmai 52, Wanmai 29, Yangmai 9		 川麦47
Chuanmai 47		 Abodanza, Kitanokaori, Klein Jabal 1

附表2

标记的分布及多态性"

染色体
Chromosome		 标记数目
No. of marker		 染色体长度
Chromosome length (Mb)		 标记密度
Density of marker (Mb)		 遗传多样性
Genetic diversity		 多态信息量 PIC
平均值
Mean		 变异范围 Range
1A 1243 592.38 0.48 0.32 0.26 0.01-0.38
1B 1292 688.60 0.53 0.33 0.27 0.02-0.38
1D 563 495.14 0.88 0.31 0.25 0.02-0.38
2A 1120 780.46 0.70 0.30 0.25 0.02-0.38
2B 1245 799.62 0.64 0.32 0.26 0.01-0.38
2D 472 650.94 1.38 0.31 0.25 0.01-0.38
3A 874 748.99 0.86 0.32 0.26 0.01-0.38
3B 1080 829.32 0.77 0.33 0.27 0.01-0.38
3D 224 613.92 2.74 0.30 0.25 0.02-0.38
4A 693 741.73 1.07 0.30 0.25 0.01-0.38
4B 565 672.56 1.19 0.32 0.26 0.02-0.38
4D 98 508.58 5.19 0.31 0.25 0.09-0.37
5A 955 709.43 0.74 0.36 0.29 0.01-0.38
5B 1142 712.82 0.62 0.36 0.28 0.01-0.38
5D 238 563.41 2.37 0.31 0.25 0.01-0.38
6A 996 617.40 0.62 0.34 0.27 0.02-0.38
6B 1066 720.82 0.68 0.32 0.26 0.01-0.38
6D 318 472.61 1.49 0.31 0.25 0.02-0.38
7A 1144 736.44 0.64 0.31 0.25 0.01-0.38
7B 856 750.49 0.88 0.32 0.26 0.01-0.38
7D 257 637.17 2.48 0.29 0.24 0.04-0.38
A型总数
Total genome A		 7025 4926.82 0.70 0.32 0.26 0.01-0.38
B型总数
Total genome B		 7246 5174.23 0.71 0.33 0.27 0.01-0.38
Total genome D 2170 3941.77 1.82 0.31 0.25 0.01-0.38
总数
Total		 16,441 14,042.83 0.85 0.32 0.26 0.01-0.38

图1

276份小麦品种(系)群体结构分析 A: 群体结构示意图; B: 群体的Δk值; C: 主成分分析图; D: 亲缘关系分析。"

表3

气孔相关性状在不同亚群间的显著性分析"

性状
Trait		 亚群
Subpopulation		 抽穗期HS 灌浆期FS
正常灌溉NI 干旱胁迫DS 正常灌溉NI 干旱胁迫DS
气孔密度SD Group1 64.34 a 66.18 a 65.98 a 72.41 a
Group2 65.30 a 66.10 a 67.06 a 73.51 a
Group3 64.77 a 66.16 a 66.28 a 72.41 a
气孔平均面积ASA Group1 2026.00 b 2100.68 a 2204.66 a 2283.14 a
Group2 2116.72 a 2126.79 a 2321.59 a 2324.24 a
Group3 2017.84 b 2015.70 a 2215.55 a 2283.30 a
气孔所占比SPP Group1 12.87 b 13.75 a 14.32 b 16.09 a
Group2 13.65 a 13.44 a 15.35 a 16.65 a
Group3 12.00 b 13.22 a 14.54 b 16.14 a

图2

不同处理下BLUE值的气孔相关性状的曼哈顿图 缩写同表1。"

附图1

不同处理下BLUE值的气孔相关性状的QQ图 缩写同表1。"

表4

与小麦气孔相关性状显著关联的稳定位点"

性状
Trait		 时期
Growth stage		 处理
Treatment		 环境
Environ-
ment		 标记
Marker		 染色体
Chr.		 物理位置
Physical
Position (Mb)		 P值
P-value		 贡献率
R2 (%)
气孔密度
SD		 抽穗期/灌浆期HS/FS 干旱胁迫DS E1 BS00064423_51 4A 627.82 3.76E-04-
8.83E-04		 4.40-
6.09
抽穗期/灌浆期HS/FS 正常灌溉NI BLUE Ex_c69429_328 6A 2.40 7.99E-05-
7.33E-04		 2.31-
3.06
气孔平均
面积
ASA		 灌浆期FS 正常灌溉NI E1/BLUE Kukri_c13458_226 1B 452.28 1.62E-04-
4.15E-04		 5.87-
6.55
灌浆期FS 正常灌溉NI E1/BLUE Kukri_c1308_2236 3A 650.04 1.12E-04-
8.66E-04		 3.42-
3.94
灌浆期FS 正常灌溉NI E2/BLUE wsnp_Ex_rep_c102011_87270703 6A 108.07-109.99 1.58E-04-
7.90E-04		 2.78-
6.22
灌浆期FS 正常灌溉NI E2/BLUE wsnp_Ra_c16745_25482384 6A 112.58-115.02 2.46E-04-
8.94E-04		 3.31-
5.71

表5

与2个气孔相关性状显著关联位点"

性状
Trait		 处理
Treatment		 标记
Marker		 染色体
Chr.		 物理位置
Physical
position (Mb)		 P值
P-value		 贡献率
R2 (%)
气孔平均面积/气孔所占比ASA/SPP 正常灌溉/干旱胁迫NI/DS Kukri_c45512_193 1A 533.90 4.40E-04-
9.92E-04		 4.20-
4.88
气孔平均面积/气孔所占比ASA/SPP 正常灌溉NI Kukri_c13458_226 1B 452.28 1.62E-04-
6.89E-04		 4.02-
6.55
气孔平均面积/气孔所占比ASA/SPP 正常灌溉NI Kukri_c1308_2236 3A 650.04 1.12E-04-
8.66E-04		 1.25-
3.42
气孔密度/气孔所占比SD/SPP 正常灌溉/干旱胁迫NI/DS Kukri_c9259_421 4A 673.07 4.88E-04-
9.20E-04		 3.03-
5.50
气孔密度/气孔所占比SD/SPP 正常灌溉/干旱胁迫NI/DS tplb0043g09_926 5A 395.06 3.95E-04-
8.13E-04		 5.00-
7.31
气孔平均面积/气孔所占比ASA/SPP 正常灌溉/干旱胁迫NI/DS wsnp_Ex_rep_c69627_68580121 6A 108.07-109.99 1.58E-04-
8.46E-04		 2.78-
6.47
气孔平均面积/气孔所占比ASA/SPP 正常灌溉/干旱胁迫NI/DS wsnp_Ra_c16745_25482384 6A 115.02 2.46E-04-
9.49E-04		 3.70-
6.20
气孔密度/气孔所占比SD/SPP 正常灌溉/干旱胁迫NI/DS RAC875_c18340_661 6D 51.95 9.55E-04-
9.72E-04		 4.78-
6.55

图3

wsnp_Ex_rep_c69627_68580121单倍型分析 A: 6A染色体上wsnp_Ex_rep_c69627_68580121标记关联的连锁区域; B: 不同麦区中wsnp_Ex_rep_c69627_68580121标记的2个单倍型分布频率。NWWR: 北部冬麦区; FV: 国外品种(系); YHFWWR: 黄淮冬麦区; SWWR: 西南冬麦区; MLYWWR: 长江中下游冬麦区。"

表6

单倍型结果的统计分析"

性状
Trait		 标记
Marker		 单倍型Haplotype 等位基因Allele 频率
Frequency (%)		 ASA平均值
ASA mean		 SPP平均值
SPP mean
(%)
NI (μm2) DS (μm2)
气孔平均面积/
气孔所占比ASA/SPP		 wsnp_Ex_rep_c69627_68580121 Hap1 TT 81.20 2193.68 a 2259.09 a 14.37 a
Hap2 CC 18.80 2419.18 b 2426.91 b 15.84 b

表7

筛选获得候选基因信息"

染色体
Chr.		 位点
Marker name		 性状
Trait		 物理位置
Position (Mb)		 基因
Gene		 基因注释或编码蛋白
Gene annotation or coding protein
1A Kukri_c45512_193 ASA/SPP 533.90 TraesCS7B01G298100 肉桂酰辅酶A还原酶4
Cinnamoyl-CoA reductase 4
1B Kukri_c13458_226 ASA/SPP 452.28 TraesCS1B01G261200 GDSL 酯酶/脂肪酶
GDSL esterase/lipase
3A D_contig72157_427 SD 46.17 TraesCS3A01G074400 细胞色素P450家族蛋白, 表达
Cytochrome P450 family protein, expressed
4A BS00064423_51 SD 627.82 TraesCS4A01G351400 F-box蛋白家族F-box family protein
5A tplb0043g09_926 SD/SPP 395.06 TraesCS5A01G192600 F-box蛋白家族F-box family protein
6A Ex_c69429_328 SD 2.40 TraesCS6A01G007300 类Myb家族转录因子蛋白
Myb family transcription factor-like protein
6A wsnp_Ex_rep_c69627_68580121 ASA/SPP 108.07-109.99 TraesCS6A01G189300LC 60S核糖体蛋白L28
60S ribosomal protein L28
6A wsnp_Ra_c16745_25482384 ASA/SPP 112.58-115.02 TraesCS6A01G193800LC ABC转运蛋白ATP结合蛋白
ABC transporter ATP-binding protein
6D RAC875_c18340_661 SD/SPP 51.95 TraesCS5A01G038300 生长素响应因子Auxin response factor
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