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Acta Agronomica Sinica ›› 2025, Vol. 51 ›› Issue (9): 2266-2284.doi: 10.3724/SP.J.1006.2025.51028

• CROP GENETICS & BREEDING·GERMPLASM RESOURCES·MOLECULAR GENETICS • Previous Articles     Next Articles

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 Online:2025-09-12 Published:2025-06-13
  • Contact: *E-mail: hw-geng@163.com E-mail:1455812032@qq.com;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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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

Table 1

Statistical analysis of phenotypic variation in the population across different environments"

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

Table 2

Population phenotypic variance analysis under different treatments"

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

Table S1

Distribution of wheat varieties (lines) based on BLUE values in different wheat-growing regions"

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

Table S2

Distribution and polymorphism of markers"

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

Fig. 1

Population structure analysis of 276 wheat varieties (lines) A: group structure diagram; B: the estimation of ?K value in population; C: principal component analysis; D: relationship analysis."

Table 3

Significance analysis of stomatal-related traits among different subpopulations"

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

Fig. 2

Manhattan plots of stomatal related traits with BLUE values under different treatments Abbreviations are the same as those given in Table 1."

Fig. S1

QQ plots of stomatal related traits with BLUE values under different treatments Abbreviations are the same as those given in Table 1."

Table 4

Stable loci significantly associated with stomatal traits in wheat"

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

Table 5

Genetic loci significantly associated with two stomatal traits"

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

Fig. 3

Haplotype analysis of wsnp_Ex_rep_c69627_68580121 A: LD regions associated with wsnp_Ex_rep_c69627_68580121 marker on chromosome 6A; B: haplotype distribution frequency of wsnp_Ex_rep_c69627_68580121 in different wheat regions. NWWR: the northern winter wheat region; FV: foreign varieties (lines); YHFWWR: Yellow and Huai winter wheat region; SWWR: southwestern winter wheat region; MLYWWR: middle and lower Yangtze River winter wheat region."

Table 6

Statistical analysis of haplotype results"

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

Table 7

Identification and information of candidate genes"

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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