冬小麦籽粒主要品质性状的全基因组关联分析

doi:10.3864/j.issn.0578-1752.2023.11.002

摘要/Abstract

摘要：

【目的】小麦籽粒品质是影响小麦加工品质和营养价值的重要因素。挖掘与小麦籽粒品质性状显著关联的位点及候选基因，为拓宽对小麦品质性状遗传机理的理解和分子标记辅助优质小麦育种提供依据。【方法】通过对来自国内外259份冬小麦品种（系）的蛋白质含量、湿面筋含量、淀粉含量、沉降值和籽粒硬度等5个品质性状进行测定，并结合90K SNP芯片进行全基因组关联分析，将定位到的显著性关联位点进行单倍型分析。【结果】5个性状均符合正态分布且在不同环境间均表现出丰富的变异，沉降值的变异系数最大（20.11%—24.42%）。各性状在基因型、环境、基因型×环境间均呈现出极显著差异（P<0.001），广义遗传力为0.77—0.84。通过全基因组关联分析，共检测到44个与5个性状显著关联（P<0.001）的位点，分布在除1D和3D染色体外的其他19个连锁群。在2个及以上的环境中均稳定存在的位点18个，涉及蛋白质含量（12个）、湿面筋含量（9个）、淀粉含量（11个）、沉降值（12个）和籽粒硬度（7个）等5个性状，能解释遗传变异的4.27%—10.98%。其中13个为多效应位点，与蛋白质含量、湿面筋含量、沉降值和淀粉含量等性状相关联的多效应位点最多（7个）。位于2B、2D和3A染色体的GENE-0762_631、IAAV7742和RAC875_c66845_466位点同时在2个环境和BLUP值下被检测到，表型贡献率的范围为4.32%—7.07%。通过对多环境下存在且表型贡献率高的多效应位点进行单倍型分析，在5D染色体的D_GDS7LZN02F4FP5_176位点发掘到与蛋白质含量、沉降值和淀粉含量等性状显著相关的Hap1、Hap2、Hap3和Hap4等4个不同单倍型，其中，Hap1是高淀粉含量单倍型（P<0.001），而Hap2和Hap3均为高蛋白质含量和沉降值的单倍型（P<0.05），4个单倍型分别占74.22%、16.21%、6.92%和2.65%。对不同来源冬小麦的单倍型分布频率进行分析，其中，高蛋白质含量和沉降值单倍型Hap2的分布频率由高到低为黄淮冬麦区>北部冬麦区>国外品种>长江中下游冬麦区>西南冬麦区。对稳定遗传的位点进行候选基因的挖掘，筛选到10个可能与小麦籽粒品质相关的候选基因。【结论】检测到18个与籽粒品质性状显著关联的稳定位点，鉴定到4个不同单倍型，筛选出10个与籽粒品质相关的候选基因。

关键词: 冬小麦, 品质, SNP标记, GWAS, 单倍型, 候选基因

Abstract:

【Objective】The quality of wheat grain was an important factor affecting the processing quality and nutritional. Mining loci and candidate genes significantly associated with wheat grain quality traits provided a basis for broadening the understanding of the genetic mechanism of quality traits and molecular marker-assisted quality. 【Method】By measuring five quality traits, including protein content (GPC), wet gluten content (WGC), starch content (GSC), settling value (SV) and grain hardness (GH), in 259 winter wheat varieties (lines) from domestic and abroad, and conducting genome-wide association analysis in combination with 90K SNP chip, the significant association loci located were subjected to haplotype analysis. 【Result】All five traits conformed to normal distribution and showed rich variation among different environments, and the coefficient of variation of sedimentation value was the largest (20.11%-24.42%). All traits have shown highly significant differences (P<0.001) among genotype, environment, and genotype×environment, with a broad-sense heritability of 0.77-0.84. A total of 44 loci significantly associated (P<0.001) with five traits were detected by genome-wide association analysis, distributed in 19 linkage groups other than chromosomes 1D and 3D. Eighteen loci were stable in two or more environments, involving all five traits including protein content (12), wet gluten content (9), starch content (11), sedimentation value (12) and grain hardness (7), explaining 4.27%-10.98% of the genetic variation. Thirteen of them were multi-effect loci, with the largest number of multi-effect loci (7) associated with traits such as protein content, wet gluten content, settling value and starch content. The GENE-0762_631, IAAV7742 and RAC875_c66845_466 loci located on 2B, 2D and 3A chromosomes were detected simultaneously at two environmental and BLUP values with a range of 4.32%-7.07% phenotypic contribution. Through haplotype analysis of multi-effect loci present in multiple environments with high phenotypic contribution, four different haplotypes, Hap1, Hap2, Hap3 and Hap4, which were significantly associated with traits such as protein content, sedimentation value and starch content, were uncovered at the D_GDS7LZN02F4FP5_176 locus of chromosome 5D, among them Hap1 was a high starch content haplotype (P<0.001), while Hap2 and Hap3 were both haplotypes with high protein content and sedimentation value (P<0.05), and the four haplotypes accounted for 74.22%, 16.21%, 6.92% and 2.65%, respectively. The distribution frequencies of haplotypes from different sources of winter wheat were analyzed, in which the distribution frequencies of haplotype Hap2 with high protein content and sedimentation value were from high to low in the Huanghuai winter wheat regions>northern winter wheat region>abroad varieties>middle and lower reaches of the Yangtze River winter wheat region>southwest winter wheat region. Candidate genes were mined for stable genetic loci, and 10 candidate genes that might be related to wheat grain quality were screened. 【Conclusion】In the study, 18 stable loci significantly associated with grain quality traits were detected, 4 different haplotypes were identified, and 10 candidate genes related to grain quality were screened.

Key words: winter wheat, quality, SNP marker, GWAS, haplotype, candidate genes

董一帆, 任毅, 程宇坤, 王睿, 张志辉, 时晓磊, 耿洪伟. 冬小麦籽粒主要品质性状的全基因组关联分析[J]. 中国农业科学, 2023, 56(11): 2047-2063.

DONG YiFan, REN Yi, CHENG YuKun, WANG Rui, ZHANG ZhiHui, SHI XiaoLei, GENG HongWei. Genome-Wide Association Study of Grain Main Quality Related Traits in Winter Wheat[J]. Scientia Agricultura Sinica, 2023, 56(11): 2047-2063.

图/表 9

表1

表2

图1

图2

表3

与小麦籽粒品质性状显著关联的稳定位点"

性状 Trait	标记 Marker	染色体 Chr.	位置 Position (Mb)	P值 P-value	表型贡献率 R²	环境 Environment
蛋白质含量 GPC (%)	BobWhite_c1027_1127	1A	586.91—590.20	1.26E^-04-7.47E^-04	4.89—7.32	E2/E3
	wsnp_Ex_c5740_10081171	2B	733.71—742.81	4.60E^-04-8.98E^-04	4.63—5.14	E1/E3
	GENE-0762_631	2B	790.44—790.76	5.76E^-05-8.35E^-04	4.57—6.55	E1/E2/E3
	IAAV7742	2D	645.61	4.41E^-05-9.28E^-04	4.63—7.07	E2/E3
	RAC875_c66845_466	3A	54.46—61.31	1.47E^-04-9.65E^-04	4.32—5.78	E1/E2/E3
	wsnp_Ex_c8360_14085858	3B	5.67—5.95	6.13E^-04-8.42E^-04	4.55—7.67	E2/E3
	Excalibur_rep_c102020_253	4A	631.90—631.92	4.96E^-05-9.82E^-04	4.42—7.18	E1/E3
	Tdurum_contig33737_157	4B	37.69	3.04E^-04-7.19E^-04	4.59—5.02	E1/E3
	BS00077733_51	5B	693.87	3.25E^-04-9.88E^-04	4.27—5.82	E2/E3
	D_GDS7LZN02F4FP5_176	5D	559.92—562.12	2.12E^-04-8.58E^-04	4.53—6.26	E2/E3
	Kukri_c74599_85	7A	14.19—19.97	6.81E^-05-9.21E^-04	4.64—6.40	E1/E3
	RAC875_c80504_487	7A	30.09—35.60	8.05E^-04-9.89E^-04	5.61—8.55	E2/E3
湿面筋含量 WGC (%)	BobWhite_c1027_1127	1A	586.91	4.09E^-05-7.87E^-04	5.23—6.79	E2/E3
	wsnp_Ex_c5740_10081171	2B	742.81	5.86E^-04-9.77E^-04	4.46—5.02	E1/E3
	GENE-0762_631	2B	790.44—790.76	5.76E^-05-7.62E^-04	4.89—5.45	E1E3
	RAC875_c66845_466	3A	54.46	3.75E^-04-6.06E^-04	4.32—4.79	E1/E2/E3
	wsnp_Ex_c8360_14085858	3B	5.67—5.95	7.23E^-04-8.04E^-04	5.62—8.91	E2/E3
	Excalibur_rep_c102020_253	4A	631.90—631.92	3.35E^-04-7.28E^-04	4.69—6.36	E1/E3
	Tdurum_contig33737_157	4B	37.69	3.04E^-04-6.37E^-04	4.78—5.21	E1/E3
	BS00077733_51	5B	693.87	4.66E^-04-6.03E^-04	4.70—5.17	E2/E3
	Kukri_c74599_85	7A	14.19—19.97	4.02E^-04-6.23E^-04	4.74—5.08	E1/E3
沉降值 SV (mL)	BobWhite_c1027_1127	1A	586.91—593.30	5.25E^-04-5.79E^-04	4.66—7.36	E2/E3
	wsnp_Ex_c5740_10081171	2B	733.71—742.81	4.60E^-04-8.23E^-04	4.78—5.14	E1/E3
	GENE-0762_631	2B	790.44—790.76	6.26E^-04-7.09E^-04	5.02—6.19	E1/E2
	IAAV7742	2D	645.61	8.84E^-05-6.72E^-04	5.81—7.04	E1/E3
	RAC875_c66845_466	3A	54.46—61.31	4.03E^-04-9.58E^-04	4.25—5.02	E1/E3
	wsnp_Ex_c8360_14085858	3B	5.67—5.95	6.66E^-04-8.40E^-04	5.69—7.21	E2/E3
	Excalibur_rep_c102020_253	4A	631.90—631.92	5.30E^-04-6.78E^-04	4.66—5.19	E1/E3
	Tdurum_contig33737_157	4B	37.69	6.22E^-04-7.00E^-04	4.59—5.10	E1/E3
	BS00077733_51	5B	693.87	5.22E^-04-7.28E^-04	4.63—5.24	E2/E3
	D_GDS7LZN02F4FP5_176	5D	559.92—562.12	2.55E^-04-7.63E^-04	4.53—5.32	E2/E3
	Kukri_c67_1504	6A	73.72—75.02	2.86E^-04-9.14E^-04	4.44—5.44	E1/E3
	RAC875_c80504_487	7A	30.09—35.60	8.05E^-04-8.69E^-04	5.82—8.06	E2/E3
淀粉含量 GSC (%)	BobWhite_c1027_1127	1A	593.3	5.62E^-05-2.71E^-04	5.06—7.59	E2/E3
	wsnp_Ex_c5740_10081171	2B	733.71—742.81	4.60E^-04-7.67E^-04	4.46—5.11	E1/E3
	GENE-0762_631	2B	790.44—790.76	9.23E^-05-6.26E^-04	4.57—6.02	E2/E3
	IAAV7742	2D	645.61	7.78E^-05-4.36E^-04	5.03—6.58	E1/E2/E3
	RAC875_c66845_466	3A	54.46—61.31	2.06E^-04-7.26E^-04	4.65—5.28	E2/E3
	Excalibur_rep_c102020_253	4A	631.90—631.92	6.15E^-04-8.62E^-04	4.89—6.69	E1/E3
	Tdurum_contig33737_157	4B	37.69	5.32E^-04-5.98E^-04	4.63—5.63	E1/E3
	BS00077733_51	5B	693.87	4.24E^-04-6.06E^-04	4.27—5.82	E2/E3
	D_GDS7LZN02F4FP5_176	5D	559.92—562.12	4.62E^-04-7.31E^-04	5.26—7.08	E2/E3
	Kukri_c67_1504	6A	73.72—75.02	4.26E^-04-6.26E^-04	4.87—5.02	E1/E3
	RAC875_c80504_487	7A	35.6	8.05E^-04-9.08E^-04	6.23—7.03	E2/E3
籽粒硬度 GH (%)	GENE-1019_96	2A	2.48	1.51E^-04-4.90E^-04	4.96—5.83	E1/E2/E3
	BobWhite_c2002_100	2A	535.72	5.67E^-04-7.71E^-04	4.66—4.99	E1/E3
	BS00081871_51	2B	12.08—17.39	6.81E^-05-9.21E^-04	5.08—6.40	E1/E3
	wsnp_Ex_c8360_14085858	3B	5.67—5.95	6.18E^-04-6.41E^-04	5.60—7.99	E2/E3
	BS00000020_51	5D	3.61	4.06E^-07-1.82E^-05	7.75—10.98	E1/E2/E3
	Kukri_c74599_85	7A	14.19—19.97	5.22E^-05-7.26E^-04	5.02—6.21	E1/E3
	Kukri_c34147_152	7A	689.95—690.05	2.81E^-04-5.26E^-04	6.08—6.78	E2/E3

表3

表4

与2个及2个以上籽粒品质性状显著关联的稳定位点"

性状 Trait	标记 Marker	染色体 Chr.	位置 Position (Mb)	P值 P-value	表型贡献率 R²	环境 Environment
蛋白质含量、湿面筋含量、沉降值、淀粉含量 GPC, WGC, SV, GSC	BobWhite_c1027_1127	1A	586.91—593.30	4.09E^-05-7.87E^-04	4.66—7.59	E2/E3
蛋白质含量、湿面筋含量、沉降值、淀粉含量 GPC, WGC, SV, GSC	wsnp_Ex_c5740_10081171	2B	733.71—742.81	4.60E^-04-9.77E^-04	4.46—5.14	E1/E3
蛋白质含量、湿面筋含量、沉降值、淀粉含量 GPC, WGC, SV, GSC	GENE-0762_631	2B	790.44—790.76	5.76E^-05-8.35E^-04	4.57—6.55	E1/E2/E3
蛋白质含量、沉降值、淀粉含量 GPC, SV, GSC	IAAV7742	2D	645.61	4.41E^-05-9.28E^-04	4.63—7.07	E1/E2/E3
蛋白质含量、湿面筋含量、沉降值、淀粉含量 GPC, WGC, SV, GSC	RAC875_c66845_466	3A	54.46—61.31	1.47E^-04-9.65E^-04	4.32—5.78	E1/E2/E3
蛋白质含量、湿面筋含量、沉降值、籽粒硬度 GPC, WGC, SV, GH	wsnp_Ex_c8360_14085858	3B	5.67—5.95	6.13E^-04-8.42E^-04	4.55—8.91	E2/E3
蛋白质含量、湿面筋含量、沉降值、淀粉含量 GPC, WGC, SV, GSC	Excalibur_rep_c102020_253	4A	631.90—631.92	4.96E^-05-9.82E^-04	4.42—7.18	E1/E3
蛋白质含量、湿面筋含量、沉降值、淀粉含量 GPC, WGC, SV, GSC	Tdurum_contig33737_157	4B	37.69	3.04E^-04-7.19E^-04	4.60—5.21	E1/E3
蛋白质含量、湿面筋含量、沉降值、淀粉含量 GPC, WGC, SV, GSC	BS00077733_51	5B	693.87	3.25E^-04-9.88E^-04	4.27—5.82	E2/E3
蛋白质含量、沉降值、淀粉含量 GPC, SV, GSC	D_GDS7LZN02F4FP5_176	5D	559.92—562.12	2.12E^-04-8.58E^-04	4.53—7.08	E2/E3
沉降值、淀粉含量SV, GSC	Kukri_c67_1504	6A	73.72—75.02	2.86E^-04-9.14E^-04	4.44—5.44	E1/E3
蛋白质含量、湿面筋含量、籽粒硬度 GPC, WGC, GH	Kukri_c74599_85	7A	14.19—19.97	6.81E^-05-9.21E^-04	4.64—6.40	E1/E3
蛋白质含量、沉降值、淀粉含量 GPC, SV, GSC	RAC875_c80504_487	7A	30.09—35.60	8.05E^-04-9.89E^-04	5.61—8.55	E2/E3

表4

图3

图4

表5

参考文献 46

[1]	贾光锋, 范丽霞, 王金水. 小麦面筋蛋白结构､功能性及应用. 粮食加工, 2004, 29(2): 11-13, 22.
	JIA G F, FAN L X, WANG J S. The structure, functional properties and using of wheat gluten protein. Grain Processing, 2004, 29(2): 11-13, 22. (in Chinese)
[2]	王一杰, 辛岭, 胡志全, 安晓宁. 我国小麦生产､消费和贸易的现状分析. 中国农业资源与区划, 2018, 39(5): 36-45.
	WANG Y J, XIN L, HU Z Q, AN X N. Current sittuation of production, consumption and trade of wheat in China. Chinese Journal of Agricultural Resources and Regional Planning, 2018, 39(5): 36-45. (in Chinese)
[3]	张爱民, 李欣, 刘冬成, 孙家柱, 阳文龙. 品质支撑农作物产业与未来发展. 中国农业科学, 2016, 49(22): 4265-4266. doi: 10.3864/j.issn.0578-1752.2016.22.001
	ZHANG A M, LI X, LIU D C, SUN J Z, YANG W L. Quality-The future of crop production. Scientia Agricultura Sinica, 2016, 49(22): 4265-4266. (in Chinese)
[4]	赵新, 王步军. 小麦蛋白质和淀粉性状与面包品质关系研究进展. 中国农学通报, 2008, 24(12): 124-127.
	ZHAO X, WANG B J. Advances in relationship of bread quality and characteristics of protein and starch of wheat. Chinese Agricultural Science Bulletin, 2008, 24(12): 124-127. (in Chinese)
[5]	刘建军, 何中虎, 赵振东, 宋建民, 刘爱峰. 小麦面条加工品质研究进展. 麦类作物学报, 2001, 21(2): 81-84.
	LIU J J, HE Z H, ZHAO Z D, SONG J M, LIU A F. Review of noodle industrial quality of wheat. Journal of Triticeae Crops, 2001, 21(2): 81-84. (in Chinese)
[6]	杜巍, 魏益民, 张国权, 欧阳韶辉, 胡新中. 小麦品质与面条品质关系的研究. 西北农林科技大学学报(自然科学版), 2001, 29(3): 24-28.
	DU W, WEI Y M, ZHANG G Q, OUYANG S H, HU X Z. Study on the relation of wheat property and noodle quality. Journal of Northwest A&F University (Natural Science Edition), 2001, 29(3): 24-28. (in Chinese)
[7]	陈锋, 陈东升, 钱森和, 张艳, 夏先春, 何中虎. Puroindoline基因对春小麦磨粉及馒头､面条品质的影响. 作物学报, 2006, 32(7): 980-986.
	CHEN F, CHEN D S, QIAN S H, ZHANG Y, XIA X C, HE Z H. Influence of Puroindoline gene on milling performance steamed bread and noodle qualities in spring wheat. Acta Agronomica Sinica, 2006, 32(7): 980-986. (in Chinese)
[8]	周艳华, 何中虎, 阎俊, 张艳, 王德森. 中国小麦品种磨粉品质研究. 中国农业科学, 2003, 36(6): 615-621.
	ZHOU Y H, HE Z H, YAN J, ZHANG Y, WANG D S. Characterization of milling quality in Chinese common wheat. Scientia Agricultura Sinica, 2003, 36(6): 615-621. (in Chinese)
[9]	WANG D W, ZHANG K P, DONG L L, DONG Z Y, LI Y W, HUSSAIN A, ZHAI H J. Molecular genetic and genomic analysis of wheat milling and end-use traits in China: Progress and perspectives. The Crop Journal, 2018, 6(1): 68-81. doi: 10.1016/j.cj.2017.10.001
[10]	严勇亮, 时晓磊, 张金波, 耿洪伟, 肖菁, 路子峰, 倪中福, 丛花. 春小麦籽粒主要品质性状的全基因组关联分析. 中国农业科学, 2021, 54(19): 4033-4047. doi: 10.3864/j.issn.0578-1752.2021.19.001
	YAN Y L, SHI X L, ZHANG J B, GENG H W, XIAO J, LU Z F, NI Z F, CONG H. Genome-wide association study of grain quality related characteristics of spring wheat. Scientia Agricultura Sinica, 2021, 54(19): 4033-4047. (in Chinese) doi: 10.3864/j.issn.0578-1752.2021.19.001
[11]	关二旗, 魏益民, 张波. 小麦籽粒品质与基因型及环境条件的关系. 麦类作物学报, 2010, 30(5): 963-969.
	GUAN E Q, WEI Y M, ZHANG B. Relationships between wheat kernel quality and genotype as well as environmental conditions. Journal of Triticeae Crops, 2010, 30(5): 963-969. (in Chinese)
[12]	ROZBICKI J, CEGLIŃSKA A, GOZDOWSKI D, JAKUBCZAK M, CACAK-PIETRZAK G, MĄDRY W, GOLBA J, PIECHOCIŃSKI M, SOBCZYŃSKI G, STUDNICKI M, DRZAZGA T. Influence of the cultivar, environment and management on the grain yield and bread-making quality in winter wheat. Journal of Cereal Science, 2015, 61: 126-132. doi: 10.1016/j.jcs.2014.11.001
[13]	PU Z E, YE X L, LI Y, SHI B X, GUO Z, DAI S F, MA J, LIU Z H, JIANG Y F, LI W, JIANG Q T, CHEN G Y, WEI Y M, ZHENG Y L. Identification and validation of novel loci associated with wheat quality through a genome-wide association study. Journal of Integrative Agriculture, 2022, 21(11): 3131-3147. doi: 10.1016/j.jia.2022.08.085
[14]	CHEN F, ZHANG F Y, LI H H, MORRIS C F, CAO Y Y, SHANG X L, CUI D Q. Allelic variation and distribution independence of Puroindoline b-B2 variants and their association with grain texture in wheat. Molecular Breeding, 2013, 32(2): 399-409. doi: 10.1007/s11032-013-9879-z
[15]	GIROUX M J, MORRIS C F. Wheat grain hardness results from highly conserved mutations in the friabilin components puroindoline a and b. Proceedings of the National Academy of Sciences of the United States of America, 1998, 95(11): 6262-6266.
[16]	MORRIS C F. Puroindolines: The molecular genetic basis of wheat grain hardness. Plant Molecular Biology, 2002, 48(5): 633-647. doi: 10.1023/A:1014837431178
[17]	胡文静, 张晓, 刘巧, 方正武, 高德荣. 普通小麦籽粒硬度的全基因组关联分析. 麦类作物学报, 2021, 41(2): 157-163.
	HU W J, ZHANG X, LIU Q, FANG Z W, GAO D R. Genome-wide association study of grain hardness in common wheat. Journal of Triticeae Crops, 2021, 41(2): 157-163. (in Chinese)
[18]	LOU H Y, ZHANG R Q, LIU Y T, GUO D D, ZHAI S S, CHEN A Y, ZHANG Y F, XIE C J, YOU M S, PENG H R, LIANG R Q, NI Z F, SUN Q X, LI B Y. Genome-wide association study of six quality-related traits in common wheat (Triticum aestivum L.) under two sowing conditions. Theoretical and Applied Genetics, 2021, 134(1): 399-418. doi: 10.1007/s00122-020-03704-y
[19]	黄奇鹏, 武文斌, 李聪, 孟乐, 林冬华. 中国小麦供需形势分析与对策. 现代面粉工业, 2018, 32(5): 39-42.
	HUANG Q P, WU W B, LI C, MENG L, LIN D H. Analysis and countermeasures of wheat supply and demand situation in China. Modern Flour Milling Industry, 2018, 32(5): 39-42. (in Chinese)
[20]	马冬云, 朱云集, 郭天财, 王晨阳. 基因型和环境及其互作对河南省小麦品质的影响及品质稳定性分析. 麦类作物学报, 2002, 22(4): 13-18.
	MA D Y, ZHU Y J, GUO T C, WANG C Y, Effects of genotype, environment and G×E interaction on wheat quality of Henan province and the stability analysis. Journal of Triticeae Crops, 2002, 22(4): 13-18. (in Chinese)
[21]	张萮, 张海涛, 王伟. 不同储藏方式对小麦品质的影响. 粮食科技与经济, 2017, 42(2): 68-70.
	ZHANG Y, ZHANG H T, WANG W. Effects of different storage methods on wheat quality. Grain Science and Technology and Economy, 2017, 42(2): 68-70. (in Chinese)
[22]	吴新元, 芦静, 张新忠, 黄天荣, 李建疆, 周安定, 梁晓东, 曹俊梅, 高永红, 曾潮武. 新疆小麦品质生态区划研究. 新疆农业科学, 2017, 54(8): 1373-1383. doi: 10.6048/j.issn.1001-4330.2017.08.001
	WU X Y, LU J, ZHANG X Z, HUANG T R, LI J J, ZHOU A D, LIANG X D, CAO J M, GAO Y H, ZENG C W. Study of ecological division for wheat quality in Xinjiang. Xinjiang Agricultural Sciences, 2017, 54(8): 1373-1383. (in Chinese) doi: 10.6048/j.issn.1001-4330.2017.08.001
[23]	李鸿恩, 张玉良, 吴秀琴, 李宗智. 我国小麦种质资源主要品质特性鉴定结果及评价. 中国农业科学, 1995, 28(5): 29-37.
	LI H E, ZHANG Y L, WU X Q, LI Z Z. Determination and evaluation on the main quality characters of wheat germplasm resources in China. Scientia Agricultura Sinica, 1995, 28(5): 29-37. (in Chinese)
[24]	孟智鹏, 张靖卓. 优质专用强筋和弱筋小麦生产现状､问题和对策: 基于河南等省调研分析. 农学学报, 2019, 9(3): 89-94. doi: 10.11923/j.issn.2095-4050.cjas18120012
	MENG Z P, ZHANG J Z. Current situation, problems and countermeasures of high-gluten and low-gluten wheat with premium quality for special purposes: An investigation in Henan and other provinces. Journal of Agriculture, 2019, 9(3): 89-94. (in Chinese)
[25]	关二旗, 魏益民, 张波, 郭进考, 张国权, 刘彦军, 罗勤贵, 班进福. 黄淮冬麦区部分区域小麦品种构成及品质性状分析. 中国农业科学, 2012, 45(6): 1159-1168. doi: 10.3864/j.issn.0578-1752.2012.06.014
	GUAN E Q, WEI Y M, ZHANG B, GUO J K, ZHANG G Q, LIU Y J, LUO Q G, BAN J F. Analysis of the variety composition and quality properties of wheat in a part of the Yellow-Huai river zone. Scientia Agricultura Sinica, 2012, 45(6): 1159-1168. (in Chinese) doi: 10.3864/j.issn.0578-1752.2012.06.014
[26]	CHEN J H, ZHANG F Y, ZHAO C J, LÜ G G, SUN C W, PAN Y B, GUO X Y, CHEN F. Genome-wide association study of six quality traits reveals the association of the TaRPP13L1gene with flour colour in Chinese bread wheat. Plant Biotechnology Journal, 2019, 17(11): 2106-2122. doi: 10.1111/pbi.v17.11
[27]	HAO S Y, LOU H Y, WANG H W, SHI J H, LIU D, BAOGERILE, TAO J G, MIAO S M, PEI Q C, YU L L, WU M, GAO M, ZHAO N H, DONG J C, YOU M S, XIN M M. Genome-wide association study reveals the genetic basis of five quality traits in Chinese wheat. Frontiers in Plant Science, 2022, 13: 835306. doi: 10.3389/fpls.2022.835306
[28]	RATHAN N D, KRISHNA H, ELLUR R K, SEHGAL D, GOVINDAN V, AHLAWAT A K, KRISHNAPPA G, JAISWAL J P, SINGH J B, SV S, AMBATI D, SINGH S K, BAJPAI K, MAHENDRU-SINGH A. Genome-wide association study identifies loci and candidate genes for grain micronutrients and quality traits in wheat (Triticum aestivum L.). Scientific Reports, 2022, 12(1): 1-15. doi: 10.1038/s41598-021-99269-x
[29]	MASSA A N, MORRIS C F. Molecular evolution of the puroindoline-a, puroindoline-b, and grain softness protein-1 genes in the tribe triticeae. Journal of Molecular Evolution, 2006, 63(5): 718. doi: 10.1007/s00239-006-8292-1
[30]	MUQADDASI Q H, BRASSAC J, EBMEYER E, KOLLERS S, KORZUN V, ARGILLIER O, STIEWE G, PLIESKE J, GANAL M W, RÖDER M S. Prospects of GWAS and predictive breeding for European winter wheat's grain protein content, grain starch content, and grain hardness. Scientific Reports, 2020, 10(1): 12541. doi: 10.1038/s41598-020-69381-5 pmid: 32719416
[31]	YANG Y, CHAI Y M, ZHANG X, LU S, ZHAO Z C, WEI D, CHEN L, HU Y G. Multi-locus GWAS of quality traits in bread wheat: Mining more candidate genes and possible regulatory network. Frontiers in Plant Science, 2020, 11: 1091. doi: 10.3389/fpls.2020.01091 pmid: 32849679
[32]	LI C L, BAI G H, CHAO S, CARVER B, WANG Z H. Single nucleotide polymorphisms linked to quantitative trait loci for grain quality traits in wheat. The Crop Journal, 2016, 4(1): 1-11. doi: 10.1016/j.cj.2015.10.002
[33]	WANG J Y, YANG C K, ZHAO W J, WANG Y, QIAO L, WU B B, ZHAO J J, ZHENG X W, WANG J L, ZHENG J. Genome-wide association study of grain hardness and novel Puroindoline alleles in common wheat. Molecular Breeding, 2022, 42(7): 40. doi: 10.1007/s11032-022-01303-x
[34]	陈玲玲, 刘亭萱, 谷勇哲, 宋健, 王俊, 邱丽娟. 大豆叶柄夹角相关基因GmILPA1单倍型分析. 植物遗传资源学报, 2021, 22(6): 1698-1707. doi: 10.13430/j.cnki.jpgr. 20210419003
	CHEN L L, LIU T X, GU Y Z, SONG J, WANG J, QIU L J. Haplotype analysis of petiole angle related gene GmILPA1 in soybean. Journal of Plant Genetic Resources, 2021, 22(6): 1698-1707. (in Chinese)
[35]	刘国圣, 张大乐. 功能性分子标记在小麦育种中的应用. 生物技术通报, 2016, 32(11): 18-29. doi: 10.13560/j.cnki.biotech.bull.1985.2016.11.003
	LIU G S, ZHANG D L. The application of the functional molecular marker in wheat breeding. Biotechnology Bulletin, 2016, 32(11): 18-29. (in Chinese)
[36]	JIN X F, FENG B, XU Z B, FAN X L, LIU J, LIU Q, ZHU P, WANG T. TaAAP6-3B, a regulator of grain protein content selected during wheat improvement. BMC Plant Biology, 2018, 18(1): 71. doi: 10.1186/s12870-018-1280-y pmid: 29685104
[37]	何中虎, 庄巧生, 程顺和, 于振文, 赵振东, 刘旭. 中国小麦产业发展与科技进步. 农学学报, 2018, 8(1): 99-106.
	HE Z H, ZHUANG Q S, CHENG S H, YU Z W, ZHAO Z D, LIU X. Wheat production and technology improvement in China. Journal of Agriculture, 2018, 8(1): 99-106. (in Chinese)
[38]	何中虎, 夏先春, 陈新民, 庄巧生. 中国小麦育种进展与展望. 作物学报, 2011, 37(2): 202-215.
	HE Z H, XIA X C, CHEN X M, ZHUANG Q S. Progress of wheat breeding in China and the future perspective. Acta Agronomica Sinica, 2011, 37(2): 202-215. (in Chinese) doi: 10.3724/SP.J.1006.2011.00202
[39]	李琳, 丁峰, 潘介春, 张树伟, 黄幸, 王金英, 王颖, 李浩然, 徐炯志, 彭宏祥, 何新华. 植物锌指蛋白转录因子家族研究进展. 热带农业科学, 2020, 40(2): 65-75.
	LI L, DING F, PAN J C, ZHANG S W, HUANG X, WANG J Y, WANG Y, LI H R, XU J Z, PENG H X, HE X H. Research progress on family of plant zinc-finger protein transcription factors. Chinese Journal of Tropical Agriculture, 2020, 40(2): 65-75. (in Chinese)
[40]	MIZUTANI M, SATO F. Unusual P450 reactions in plant secondary metabolism. Archives of Biochemistry and Biophysics, 2011, 507(1): 194-203. doi: 10.1016/j.abb.2010.09.026 pmid: 20920462
[41]	冯蕾, 张海文, 黄荣峰. 植物LRR类受体蛋白激酶的研究进展. 中国农业科技导报, 2012, 14(6): 43-48.
	FENG L, ZHANG H W, HUANG R F. Research progress on LRR receptor-like protein kinase in plant. Journal of Agricultural Science and Technology, 2012, 14(6): 43-48. (in Chinese)
[42]	JIA Q, XIAO Z X, WONG F L, SUN S, LIANG K J, LAM H M. Genome-wide analyses of the soybean F-box gene family in response to salt stress. International Journal of Molecular Sciences, 2017, 18(4): 818. doi: 10.3390/ijms18040818
[43]	杨乐, 齐妍, 刘生祥, 张双喜, 李连城, 陈明, 徐兆师, 马有志. 植物抗逆相关蛋白激酶的结构与功能. 植物遗传资源学报, 2013, 14(4): 659-667.
	YANG L, QI Y, LIU S X, ZHANG S X, LI L C, CHEN M, XU Z S, MA Y Z. Structure and function of stress-related protein kinases in plants. Journal of Plant Genetic Resources, 2013, 14(4): 659-667. (in Chinese)
[44]	胡梦芸, 张正斌, 徐萍. 植物光合产物转运蛋白及其生物学功能. 植物生理学通讯, 2008, 44(1): 1-6.
	HU M Y, ZHANG Z B, XU P. Photoassimilate transport proteins and biology function in plant. Plant Physiology Communications, 2008, 44(1): 1-6. (in Chinese)
[45]	KEEGSTRA K, RAIKHEL N. Plant glycosyltransferases. Current Opinion in Plant Biology, 2001, 4(3): 219-224. pmid: 11312132
[46]	SOSSO D, LUO D P, LI Q B, SASSE J, YANG J L, GENDROT G, SUZUKI M, KOCH K E, MCCARTY D R, CHOUREY P S, ROGOWSKY P M, ROSS-IBARRA J, YANG B, FROMMER W B. Seed filling in domesticated maize and rice depends on SWEET-mediated hexose transport. Nature Genetics, 2015, 47(12): 1489-1493. doi: 10.1038/ng.3422

性状 Traits	环境 Environment	范围 Range	平均值±标准差 Mean±SD	变异系数 CV (%)	偏度 Ske.	峰度 Kur.	F值F value			遗传力 h²
性状 Traits	环境 Environment	范围 Range	平均值±标准差 Mean±SD	变异系数 CV (%)	偏度 Ske.	峰度 Kur.	基因型 Genotype	环境 Environment	基因型×环境 G×E	遗传力 h²
蛋白质含量 GPC (%)	E1	13.52—19.22	16.38±1.15	7.01	-0.10	-0.70	16.96***	3101.12***	4.29***	0.79
蛋白质含量 GPC (%)	E2	12.77—18.67	15.55±1.21	7.77	0.40	-0.26	16.96***	3101.12***	4.29***	0.79
湿面筋含量 WGC (%)	E1	24.78—37.93	31.75±2.88	9.07	-0.03	-0.76	17.50***	2995.32***	4.38***	0.79
湿面筋含量 WGC (%)	E2	22.58—38.28	28.98±2.87	9.91	0.60	0.29	17.50***	2995.32***	4.38***	0.79
沉降值 SV (mL)	E1	24.05—68.55	48.18±9.69	20.11	-0.11	-0.80	13.10***	4212.30***	3.67***	0.77
沉降值 SV (mL)	E2	16.95—67.50	40.27±9.83	24.42	0.50	0.01	13.10***	4212.30***	3.67***	0.77
淀粉含量 GSC (%)	E1	57.89—62.94	60.88±0.79	1.29	-0.64	1.10	17.84***	2858.98***	4.39***	0.79
淀粉含量 GSC (%)	E2	59.94—64.14	62.40±0.75	1.20	-0.42	0.13	17.84***	2858.98***	4.39***	0.79
籽粒硬度 GH (%)	E1	41.22—77.87	65.21±7.28	11.17	-0.62	-0.18	17.10***	7352.58***	3.03***	0.84
籽粒硬度 GH (%)	E2	36.07—76.42	59.20±8.12	13.71	-0.16	-0.31	17.10***	7352.58***	3.03***	0.84

性状 Traits	统计参数 Statistical parameters	国内品种Domestic varieties				国外品种 Abroad varieties
性状 Traits	统计参数 Statistical parameters	黄淮冬麦区 Huanghuai 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	国外品种 Abroad varieties
蛋白质含量 GPC (%)	平均值±标准差 Mean±SD	16.17±0.78	16.06±0.95	15.87±0.77	15.54±0.65	15.69±0.93
	范围Range	13.94—18.15	14.46—17.98	14.32—17.10	14.62—16.89	13.61—17.6
	变异系数CV (%)	4.85	5.93	4.86	4.16	5.91
湿面筋含量 WGC (%)	平均值±标准差 Mean±SD	31.58±3.66	30.91±4.35	29.56±3.75	28.03±2.63	28.73±4.04
	范围Range	19.66—41.18	23.93—39.74	19.02—35.89	24.26—34.83	21.01—38.15
	变异系数CV (%)	11.60	14.09	12.69	9.37	14.05
沉降值 SV (mL)	平均值±标准差 Mean±SD	46.18±6.98	44.38±7.50	43.98±6.27	40.84±4.63	41.55±8.21
	范围Range	29.26—61.74	29.81—62.44	30.68—53.98	34.6—51.71	24.07—61.02
	变异系数CV (%)	15.11	16.91	14.26	11.33	19.76
淀粉含量 GSC (%)	平均值±标准差 Mean±SD	61.55±0.59	61.68±0.64	61.77±0.42	62.12±0.43	61.56±0.51
	范围Range	60.17—62.80	60.07—62.8	60.89—62.76	61.20—62.89	60.01—62.78
	变异系数CV (%)	0.95	1.04	0.67	0.69	0.83
籽粒硬度 GH (%)	平均值±标准差 Mean±SD	63.67±4.41	62.38±5.33	60.21±4.64	58.41±4.77	61.47±5.37
	范围Range	50.49—73.57	52.04—74.33	51.44—70.51	50.91—69.03	44.95—73.44
	变异系数CV (%)	6.92	8.55	7.71	8.17	8.74