小麦旗叶形态相关性状全基因组关联分析

doi:10.3724/SP.J.1006.2023.21085

Abstract

Abstract:

The flag leaf of wheat is the primary functional leaf for photosynthesis and contributes significantly to yield. Therefore, it is essential to investigate the genetic process of flag leaf morphology and identify the candidate genes for flag leaf morphology-related features. We combined 90K SNP gene chips and 300 wheat varieties (lines) for genome-wide association analysis of flag leaf length, width, and area under normal irrigation (NI), and drought stress (DS) conditions in five environments. The results showed that flag leaf length, width, and area exhibited significant differences between the two moisture treatments and displayed rich phenotypic variation with the coefficients of variation ranging from 0.07-0.23 in different environments (P<0.05). Moreover, genome-wide association study (GWAS) revealed that a total of 37 stable genetic loci were significantly associated with flag leaf length, width, and area. These loci were distributed on chromosomes 1D, 2A, 2B, 3A, 3D, 4A, 5A, 5B, 6A, 6B, 7A, and 7B, with individual SNP loci explaining 3.70%-9.05% of the genetic variation, including 22 stable genetic loci detected under normal irrigation and 15 stable genetic loci detected under drought stress. Eight stable genetic loci at the same time detected under both water treatments were discovered on chromosomes 2B, 3A, 5A, 6A, 7A, and 7B, while the five stable genetic loci related by several traits were simultaneously detected on chromosomes 2B, 3A, 6A, and 7A. By analyzing haplotypes at markers with stable inheritance and high contribution, it was found that the Kukri_c1406_275 (R²=9.05%) marker was significantly associated with flag leaf length, with three haplotypes of FLL-Hap1, FLL-Hap2, and FLL-Hap3, and the wsnp_bq170165A_Ta_1_1 (R²=7.88%) marker was also detected in three haplotypes, FLA-Hap1, FLA-Hap2, and FLA-Hap3. In combination with phenotypic analysis, the flag leaf length of 300 winter wheat varieties (lines) containing FLL-Hap1 (77.78% frequency of occurrence) or FLL-Hap2 (18.89%) haplotypes was significantly higher than that of FLL-Hap3 (3.33%) haplotypes. The flag leaf area was significantly higher in haplotypes containing FLA-Hap1 (48.19%) than in haplotypes containing FLA-Hap2 (30.80%) or FLA-Hap3 (21.01%) (P<0.05). Different haplotypes were distributed differently in different winter wheat varieties (lines). Haplotype FLL-Hap1 was more frequently distributed in foreign varieties (lines), while haplotypes FLL-Hap2 and FLL-Hap3 were more frequently distributed in the northern winter wheat region and the southwestern winter wheat region, respectively. Haplotypes FLA-Hap1 and FLA-Hap2 were more frequently distributed in the southwestern winter wheat region and the northern winter wheat region, respectively, while haplotypes FLA-Hap3 were no more frequently distributed in all winter wheat regions. Searching for stable genetic loci under both water treatments yielded and screening of five candidate genes associated with flag leaf morphology, which could be used as the important genes for flag leaf-related traits.

Key words: wheat, morphological characters of flag leaf, genome-wide association analysis, haplotype, candidate genes

WANG Rui, REN Yi, CHENG Yu-Kun, WANG Wei, ZHANG Zhi-Hui, GENG Hong-Wei. Genome-wide association analysis of morphological traits of flag leaf in wheat[J].Acta Agronomica Sinica, 2023, 49(11): 2886-2901.

Figures/Tables 8

Table 1

Table 2

Fig. 1

Fig. 2

Table 3

Flag leaf shape activity related Stable loci detected by SNP-GWAS"

性状 Trait	标记名称 Marker name	环境 Environment	处理Treatment	染色体 Chr.	位置 Position (Mb）	P值 P-value	贡献率 R²(%)	先前已报道位点 Previously reported QTL
FLL	BS00067711_51	E1/E2/BLUE	NI	2A	36.93	2.61E^‒04-5.69E^‒04	4.09-4.57
	Tdurum_contig82812_213	E3/E4	NI	2A	74.11-78.33	2.17E^‒04-9.46E^‒04	3.81-4.72	Qfll2A-1^[31]
	Excalibur_c11863_289	E1/E2/BLUE	DS	2B	157.69	2.45E^‒04-8.93E^‒04	6.87-8.40
	IACX6324	E1/BLUE	DS	2B	747.18	1.37E^‒04-4.85E^‒04	4.26-4.96	qFll-2B.1^[11]
	wsnp_Ex_c18747_27625264	E1/E2//E4/BLUE	NI	3A	645.09-652.03	4.43E^‒04-8.60E^‒04	4.57-6.84
	wsnp_Ex_c18747_27625264	E2/E3/BLUE	DS	3A	645.09-646.85	4.02E^‒04-8.05E^‒04	3.94-4.28
	RAC875_c10194_673	E1/E2/BLUE	NI	3A	683.07-686.13	1.12E^‒04-7.50E^‒04	3.90-5.04	Qfla-3A^[26]
	wsnp_Ku_c42416_50159250	E3/E2/BLUE	NI	5A	610.22-617.98	2.51E^‒04-7.17E^‒04	5.15-5.72	EPQFll.nau-5A^[12]
	BS00022378_51	E1/E2/BLUE	NI	5A	663.91-664.48	4.76E^‒06-9.50E^‒04	4.71-7.17
	BS00022378_51	E1/E2/ BLUE	DS	5A	663.91-664.48	4.76E^‒06-9.69E^‒04	3.74-7.17
	BobWhite_c8266_227	E2/BLUE	NI	5A	698.51	8.40E^‒06-5.90E^‒04	4.12-6.75
	BobWhite_c8266_227	E2 /BLUE	DS	5A	698.51	5.31E^‒05-2.61E^‒04	4.66-5.75
	RFL_Contig2206_1694	E4/ BLUE	DS	6A	600.45-608.12	2.25E^‒05-9.64E^‒04	3.87-6.45
	tplb0059j12_800	E1/E2 /BLUE	DS	6B	186.70-188.19	1.94E^‒04-7.95E^‒04	3.94-4.71
	Kukri_c1406_275	E1/E2/BLUE	NI	7A	538.69-546.04	6.50E^‒07-7.75E^‒04	3.95-9.05
	Kukri_c1406_275	E3/BLUE	DS	7A	546.04	1.63E^‒04-4.01E^‒04	4.40-6.13
	GENE-4898_208	E1/E2/BLUE	NI	7A	567.58-572.87	1.67E^‒05-7.72E^‒04	3.78-6.42
	CAP7_rep_c5949_55	E1/BLUE	NI	7A	709.12-709.89	3.27E^‒04-4.89E^‒04	4.27-4.52
	BS00067530_51	E3/E4/	NI	7B	648.92-648.93	2.70E^‒05-9.32E^‒04	5.00-7.72
	BS00067530_51	E1/BLUE	DS	7B	648.11-648.93	2.40E^‒04-4.70E^‒04	4.10-5.60
	BobWhite_c14966_231	E1/BLUE	NI	7B	693.00	1.23E^‒04-4.36E^‒04	4.36-5.20	Qfll7B-1^[31]
FLW	CAP12_c46_333	E3/BLUE	NI	1D	15.32	8.92E^‒06-7.96E^‒04	4.14-7.45
	Ra_c19501_1510	E2/BLUE	NI	2B	11.19-14.83	7.15E^‒04	3.96-5.02
	Kukri_c783_1833	E1/BLUE	DS	3D	418.39	4.35E^‒05-3.65E^‒04	4.38-5.79
	BS00023151_51	E1/E2/BLUE	NI	4A	606.59	2.74E^‒04-7.94E^‒04	3.97-4.82
	Tdurum_contig29319_256	E3/BLUE	NI	5B	558.41-558.42	1.99E^‒05-6.10E^‒04	4.11-6.56
	BS00023192_51	E1/E2/BLUE	NI	6A	13.81-18.71	3.90E^‒04-9.76E^‒04	3.79-5.82
	RFL_Contig2206_1694	E2//BLUE	NI	6A	600.45-602.71	3.36E^‒04-9.64E^‒04	3.87-4.45
	RFL_Contig2206_1694	E2/BLUE	DS	6A	600.45	2.25E^‒05-1.31E^‒04	5.17-6.45
	tplb0024a09_2028	E1/BLUE	NI	7A	42.50-46.97	7.89E^‒04-8.19E^‒04	3.90-4.13
	Tdurum_contig66023_89	E3/E4	DS	7A	719.58-724.12	8.43E^‒06-7.80E^‒04	4.23-7.30	Qfla7A-1^[31]
FLA	Ra_c19501_1510	E1/BLUE	DS	2B	7.91-11.19	5.92E^‒05-7.84E^‒04	3.94-6.77
	wsnp_Ex_c18747_27625264	E1/E2/BLUE	NI	3A	645.09-646.85	2.26E^‒05-8.13E^‒04	3.85-6.42
	RAC875_c10194_673	E1/E2/BLUE	NI	3A	683.07	4.93E^‒04-9.55E^‒04	3.70-4.13	Qfla-3A^[26]
	wsnp_bq170165A_Ta_1_1	E1/E2/BLUE	NI	7A	553.39-553.99	9.01E^‒05-8.84E^‒04	3.81-7.88
	wsnp_bq170165A_Ta_1_1	E4/BLUE	DS	7A	553.39-553.99	2.11E^‒04-2.96E^‒04	4.50-4.80
	Tdurum_contig66023_89	E3/E4	DS	7A	721.92	3.32E^‒06-1.22E^‒04	5.29-7.62	Qfla7A-1^[31]

Table 3

Fig. 3

Table 4

Table 5

References 46

[1]	Ray D K, Mueller N D, West P C, Foley J A, Hart J P. Yield trends are insufficient to double global crop production by 2050. PLoS One, 2013, 8: e66428. doi: 10.1371/journal.pone.0066428
[2]	赵广才, 常旭虹, 王德梅, 杨玉双, 冯金凤. 中国小麦生产发展潜力研究报告. 作物杂志, 2012, (3): 1-5.
	Zhao G C, Chang X H, Wang D M, Yang Y S, Feng J F. Research report on development of China’s wheat production potential. Crops, 2012, (3): 1-5 (in Chinese with English abstract).
[3]	Liu Y X, Tao Y, Wang Z Q, Guo Q L, Wu F K, Yang X L, Deng M, Ma J, Chen G D, Wei Y M, Zheng Y L. Identification of QTL for flag leaf length in common wheat and their pleiotropic effects. Mol Breed, 2018, 38: 11. doi: 10.1007/s11032-017-0766-x
[4]	Sharma S N, Sain R S, Sharma R K. The genetic control of flag leaf length in normal and late sown durum wheat. J Agric Sci, 2003, 141: 323-331. doi: 10.1017/S0021859603003642
[5]	王义芹, 杨兴洪, 李滨, 童依平, 李振声. 小麦叶面积及光合速率与产量关系的研究. 华北农学报, 2008, 23(增刊2): 10-15.
	Wang Y Q, Yang X H, Li B, Tong Y P, Li Z S. Study on the relationship between leaf area, photosynthetic rate and yield of wheat. Acta Agric Boreali-Sin, 2008, 23(S2): 10-15 (in Chinese with English abstract).
[6]	王敏, 张从宇. 小麦旗叶性状与产量因素的相关与回归分析. 种子, 2004, (3): 17-18.
	Wang M, Zhang C Y. Correlation and regression analysis of flag leaf traits and yield components in wheat. Seeds, 2004, (3): 17-18 (in Chinese with English abstract).
[7]	景蕊莲. 作物抗旱节水研究进展. 中国农业科技导报, 2007, (1): 1-5.
	Jing R L. Advances of research on drought resistance and water use efficiency in crop plants. J Agric Sci Technol, 2007, (1): 1-5 (in Chinese with English abstract).
[8]	Biswal A K, Kohli A. Cereal flag leaf adaptations for grain yield under drought: knowledge status and gaps. Mol Breed, 2013, 31: 749-766. doi: 10.1007/s11032-013-9847-7
[9]	Liu K Y, Xu H, Liu G, Guan P F, Zhou X Y, Peng H R, Yao Y Y, Ni Z F, Sun Q X, Du J K. QTL mapping of flag leaf-related traits in wheat (Triticum aestivum L.). Theor Appl Genet, 2018, 131: 839-849. doi: 10.1007/s00122-017-3040-z
[10]	吕学莲, 白海波, 董建力, 惠建, 孙亚宁, 蔡正云, 李树华. 春小麦旗叶大小相关性状的QTL定位分析. 麦类作物学报, 2016, 36: 1587-1593.
	Lyu X L, Bai H B, Dong J L, Hui J, Sun Y N, Cai Z Y, Li S H. QTL mapping for size traits of flag leaf in spring wheat. J Triticeae Crops, 2016, 36: 1587-1593 (in Chinese with English abstract).
[11]	Fan X L, Cui F, Zhao C H, Zhang W, Yang L J, Zhao X Q, Han J, Su Q N, Ji J, Zhao Z W, Tong Y P, Li J M. QTLs for flag leaf size and their influence on yield-related traits in wheat (Triticum aestivum L.). Mol Breed, 2015, 35: 1-16. doi: 10.1007/s11032-015-0202-z
[12]	Jia H, Wan H, Yang S, Zhang Z Z, Kong Z X, Xue S L, Zhang L X, Ma Z Q. Genetic dissection of yield-related traits in a recombinant inbred line population created using a key breeding parent in China’s wheat breeding. Theor Appl Genet, 2013, 126: 2123-2139. doi: 10.1007/s00122-013-2123-8
[13]	Xue S L, Xu F, Li G Q, Zhou Y, Lin M S, Gao Z X, Su X H, Xu X W, Jiang G, Zhang S, Jia H Y, Kong Z X, Zhang L X, Ma Z Q. Fine mapping TaFLW1, a major QTL controlling flag leaf width in bread wheat (Triticum aestivum L.). Theor Appl Genet, 2013, 126: 1941-1949. doi: 10.1007/s00122-013-2108-7
[14]	朱治, 李龙, 李超男, 毛新国, 郝晨阳, 朱婷, 王景一, 常建忠, 景蕊莲. 小麦转录因子TaMYB5-3B与株高和千粒重相关. 作物学报, 2023, 49: 906-916. doi: 10.3724/SP.J.1006.2023.21029
	Zhu Z, Li L, Li C N, Mao X G, Hao C Y, Zhu T, Wang J Y, Chang J Z, Jing R L. Transcription factor TaMYB5-3B is associated with plant height and 1000-grain weight in wheat. Acta Agron Sin, 2023, 49: 906-916.
[15]	刘朦朦, 张萌娜, 张倩倩, 刘锡建, 郭宇航. 小麦旗叶宽主效QTL qFlw-5B遗传效应解析. 麦类作物学报, 2019, 39: 1399-1405.
	Liu M M, Zhang M N, Zhang Q Q, Liu X J, Guo Y H. Genetic analysis of a major stable QTL qFlw-5B for wheat flag leaf width. J Triticeae Crops, 2019, 39: 1399-1405 (in Chinese with English abstract).
[16]	李浩然, 李慧玲, 王红光, 李东晓, 李瑞奇, 李雁鸣. 冬小麦叶面积测算方法的再探讨. 麦类作物学报, 2018, 38: 455-459.
	Li H R, Li H L, Wang H G, Li D X, Li R Q, Li Y M. Further study on the method of leaf area calculation in winter wheat. J Triticeae Crops, 2018, 38: 455-459 (in Chinese with English abstract).
[17]	Cheng Y K, Li J, Yao F J, Long L, Wang Y Q, Wu Y, Li J, Ye X L, Wang J R, Jiang Q T, Kang H Y, Li W, Qi P F, Liu Y X, Deng M, Ma J, Jiang Y F, Chen X M, Zheng Y L, Wei Y M, Chen G Y. Dissection of loci conferring resistance to stripe rust in Chinese wheat landraces from the middle and lower reaches of the Yangtze River via genome-wide association study. Plant Sci, 2019, 287: 110204. doi: 10.1016/j.plantsci.2019.110204
[18]	Li D D, Xu Z X, Gu R L, Wang P X, Lyle D, Xu J L, Zhang H W, Wang G Y. Enhancing genomic selection by fitting large-effect SNPs as fixed effects and a genotype-by-environment effect using a maize BC₁F_3:4 population. PLoS One, 2019, 14: e0223898. doi: 10.1371/journal.pone.0223898
[19]	Wang S X, Zhu Y L, Zhang D X, Shao H, Liu P, Hu J B, Zhang H, Zhang H P, Chang C, Lu J, Xia X C, Sun G L, Ma C X. Genome- wide association study for grain yield and related traits in elite wheat varieties and advanced lines using SNP markers. PLoS One, 2017, 12: e0188662. doi: 10.1371/journal.pone.0188662
[20]	Zhu C S, Gore M, Buckler E S, Yu J M. Status and prospects of association mapping in plants. Plant Genome, 2008, 1: 5-20.
[21]	Yu J M, Pressoir G, Briggs W H, Bi L V, Yamasaki M, Doebley J F, Mcmullen M D, Gaut B S, Nielsen D M, Holland J B, Kresovich S, Buckler E. A unified mixed-model method for association mapping that accounts for multiple levels of relatedness. Nat Genet, 2006, 38: 203-208. doi: 10.1038/ng1702 pmid: 16380716
[22]	李天清, 王金堂, 马真胜, 雷伟, 王林, 李萌. 骨质疏松性骨折与OPG-RANK-RANKL系统基因多态性的关联分析. 中国骨质疏松杂志, 2014, 20: 247-255.
	Li T Q, Wang J T, Ma Z S, Lei W, Wang L, Li M. Correlation analysis of the relationship between the osteoporotic fracture and OPG-RANK-RANKL gene polymorphisms. Chin J Osteop, 2014, 20: 247-255 (in Chinese with English abstract).
[23]	Gabriel S B, Schaffner S F, Liu-cordero S N, Rotimi C, Adeyemo A, Cooper R, Ward R, Lander E S, Daly M J, Altshuter D, Nguyen H, Moor J M, Roy J, Blumenstiel B, Higgins J, Defelice M, Lochner A, Faggart M. The structure of haplotype blocks in the human genome. Science, 2002, 296: 2225-2229. doi: 10.1126/science.1069424 pmid: 12029063
[24]	闫雪, 史雨刚, 梁增浩, 杨斌, 李晓宇, 王曙光, 孙黛珍. 小麦旗叶形态相关性状的QTL定位. 核农学报, 2015, 29: 1253-1259. doi: 10.11869/j.issn.100-8551.2015.07.1253
	Yan X, Shi Y G, Liang Z H, Yang B, Li X Y, Wang S G, Sun D Z. QTL mapping for morphological traits of flag leaf in wheat. J Nucl Agric Sci, 2015, 29: 1253-1259 (in Chinese with English abstract). doi: 10.11869/j.issn.100-8551.2015.07.1253
[25]	宋霄君, 张敏, 李秉昌, 赵城, 刘希伟, 贾晓沛, 王琨, 蔡瑞国. 干旱胁迫对小麦营养器官物质转运和籽粒灌浆特性的影响. 中国农学通报, 2016, 32(15): 25-31. doi: 10.11924/j.issn.1000-6850.casb16010002
	Song X J, Zhang M, Li B C, Zhao C, Liu X W, Jia X P, Wang K, Cai R G. Effects of drought stress on material transport and grain-filling characteristics of wheat vegetative organs. Chin Agric Sci Bull, 2016, 32(15): 25-31 (in Chinese with English abstract).
[26]	Chen S, Liu F, Wu W X, Jiang Y, Zhan K H. A SNP-based GWAS and functional haplotype-based GWAS of flag leaf-related traits and their influence on the yield of bread wheat (Triticum aestivum L.). Theor Appl Genet, 2021, 134: 3895-3909. doi: 10.1007/s00122-021-03935-7
[27]	Khanna C R, Singh K, Shukla S, Kadam S, Singh N K. QTLs for cell membrane stability and flag leaf area under drought stress in a wheat RIL population. J Plant Biochem Biotechnol, 2020, 29: 276-286. doi: 10.1007/s13562-019-00534-y
[28]	曹英杰, 杨剑飞, 王宇. 全基因组关联分析在作物育种研究中的应用. 核农学报, 2019, 33: 1508-1518. doi: 10.11869/j.issn.100-8551.2019.08.1508
	Cao Y J, Yang J F, Wang Y. The application of GWAS in crop breeding. J Nucl Agric Sci, 2019, 33: 1508-1518 (in Chinese with English abstract). doi: 10.11869/j.issn.100-8551.2019.08.1508
[29]	Wu Q H, Chen Y X, Fu L, Zhou S H, Chen J J, Zhao X J, Zhao D, Ou-Yang S H, Wang Z Z, Li D, Wang G X, Zhang D Y, Yuan C G, Wang L X, You M S, Han J, Liu Z Y. QTL mapping of flag leaf traits in common wheat using an integrated high-density SSR and SNP genetic linkage map. Euphytica, 2016, 208: 337-351. doi: 10.1007/s10681-015-1603-0
[30]	Yang D L, Liu Y, Cheng H B, Chang L, Chen J J, Chai S X, Li M F. Genetic dissection of flag leaf morphology in wheat (Triticum aestivum L.)under diverse water regimes. BMC Genet, 2016, 17: 1-15.
[31]	连俊方, 张德强, 武炳瑾, 宋晓朋, 马文洁, 周丽敏, 冯毅, 孙道杰. 利用90K基因芯片进行小麦旗叶相关性状的QTL定位. 麦类作物学报, 2016, 36: 689-698.
	Lian J F, Zhang D Q, Wu B J, Song X P, Ma W J, Zhou L M, Feng Y, Sun D J. QTL mapping of flag leaf traits using an integrated high-density 90K genotyping chip. J Triticeae Crops, 2016, 36: 689-698 (in Chinese with English abstract).
[32]	Yan X F, Zhao L, Ren Y, Zhang N, Dong Z D, Chen F. Identification of genetic loci and a candidate gene related to flag leaf traits in common wheat by genome-wide association study and linkage mapping. Mol Breed, 2020, 6: 40-58.
[33]	严勇亮, 张恒, 张金波, 时晓磊, 耿洪伟, 肖菁, 路子峰, 倪中福, 丛花. 春小麦主要籽粒性状的全基因组关联分析. 麦类作物学报, 2022, 42: 1182-1191.
	Yan Y L, Zhang H, Zhang J B, Shi X L, Geng H W, Xiao J, Lu Z F, Ni Z F, Cong H. Genome-wide association study of grain traits in spring wheat. J Triticeae Crops, 2022, 42: 1182-1191 (in Chinese with English abstract).
[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: 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. J Plant Genet Resour, 2021, 22: 1698-1707 (in Chinese with English abstract).
[35]	赵广才. 中国小麦种植区划研究(一). 麦类作物学报, 2010, 30: 886-895.
	Zhao G C. Study on Chinese wheat planting regionalization (I). J Triticeae Crops, 2010, 30: 886-895 (in Chinese with English abstract).
[36]	茹振钢, 冯素伟, 李淦. 黄淮麦区小麦品种的高产潜力与实现途径. 中国农业科学, 2015, 48: 3388-3393. doi: 10.3864/j.issn.0578-1752.2015.17.006
	Ru Z G, Feng S W, Li G. High-yield potential and effective ways of wheat in Yellow & Huai Rivers valley facultative winter wheat region. Sci Agric Sin, 2015, 48: 3388-3393 (in Chinese with English abstract).
[37]	Zhao Z X, Zhang G Q, Zhou S M, Ren Y Q, Wang W. The improvement of salt tolerance in transgenic tobacco by overexpression of wheat F-box gene TaFBA1. Plant Sci, 2017, 259: 71-85. doi: 10.1016/j.plantsci.2017.03.010
[38]	Baute J, Polyn S, Block J D, Blomme J, Lijsebettens M V, Baute J. F-box protein FBX92 affects leaf size in Arabidopsis thaliana. Plant Cell Physiol, 2017, 58: 962-975. doi: 10.1093/pcp/pcx035 pmid: 28340173
[39]	Byeon Y, Back K. Molecular cloning of melatonin 2-hydroxylase responsible for 2-hydroxymelatonin production in rice (Oryza sativa). J Pineal Res, 2015, 58: 343-351. doi: 10.1111/jpi.12220 pmid: 25728912
[40]	Farrow S C, Facchini P J. Functional diversity of 2-oxoglutarate/ Fe(II)-dependent dioxygenases in plant metabolism. Front Plant Sci, 2014, 5: 524-539. doi: 10.3389/fpls.2014.00524 pmid: 25346740
[41]	Arnao M B, Hernández-Ruiz J. Functions of melatonin in plants: a review. J Pineal Res, 2015, 59: 133-150. doi: 10.1111/jpi.12253 pmid: 26094813
[42]	MacMillan J. Occurrence of gibberellins in vascular plants, fungi, and bacteria. J Plant Growth Regul, 2001, 20: 387-442. doi: 10.1007/s003440010038 pmid: 11986764
[43]	Park J J, Jin P, Yoon J, Yang J, Jeong H J, Ranathunge K, Schreiber L, Franke R, Lee I J, An G. Mutation in wilted dwarf and lethal 1 (WDL1) causes abnormal cuticle formation and rapid water loss in rice. Plant Mol Biol, 2010, 74: 91-103. doi: 10.1007/s11103-010-9656-x
[44]	徐文, 申浩, 郭军, 余晓丛, 李祥, 杨彦会, 马晓, 赵世杰, 宋健民. 旗叶蜡质含量不同小麦近等基因系的抗旱性. 作物学报, 2016, 42: 1700-1707.
	Xu W, Shen H, Guo J, Yu X C, Li X, Yang Y H, Ma X, Zhao S J, Song J M. Drought resistance of wheat NILs with different cuticular wax contents in flag leaf. Acta Agron Sin, 2016, 42: 1700-1707 (in Chinese with English abstract). doi: 10.3724/SP.J.1006.2016.01700
[45]	凌华. 油菜GDSL脂肪酶基因BnLIP1的克隆及其原核表达. 中国农学通报, 2007, (12): 102-106.
	Ling H. Cloning of a GDSL lipase gene from Brassica napus and preliminary study of its recombinant expression in Escherichia coli. Chin Agric Sci Bull, 2007, (12): 102-106 (in Chinese with English abstract).
[46]	Zhao Y, Tian X J, Wang F, Zhang L Y, Xin M M, Hu Z R, Yao Y Y, Ni Z F, Sun Q X, Peng H R. Characterization of wheat MYB genes responsive to high temperatures. BMC Plant Biol, 2017, 17: 208-221. doi: 10.1186/s12870-017-1158-4 pmid: 29157199

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性状 Trait	环境 Environment	处理 Treatment	最小值 Min.	最大值 Max.	平均数 Mean	标准差 SD	变异系数 CV	偏度 Skewness	峰度 Kurtosis
旗叶长 FLL (cm)	E1	NI	11.27	23.87	17.51 a	2.37	0.14	0.12	‒0.32
		DS	10.76	24.08	16.54 b	2.27	0.14	0.40	0.17
	E2	NI	9.82	24.57	16.82 a	3.02	0.18	0.46	‒0.19
		DS	9.29	26.05	15.90 b	2.79	0.18	0.49	0.41
	E3	NI	14.48	24.41	19.77 a	1.53	0.08	‒0.03	0.17
		DS	13.44	23.37	18.22 b	1.55	0.09	0.20	‒0.05
	E4	NI	12.84	23.12	18.26 a	1.41	0.08	‒0.22	0.69
		DS	12.08	22.55	17.07 b	1.71	0.10	0.20	‒0.53
	BLUE	NI	14.09	25.92	18.85 a	1.97	0.10	0.43	0.19
		DS	12.61	22.74	16.93 b	1.75	0.10	0.51	0.17
旗叶宽 FLW (cm)	E1	NI	0.89	1.81	1.43 a	0.14	0.10	‒0.20	0.26
		DS	0.86	1.79	1.38 b	0.13	0.09	‒0.06	0.72
	E2	NI	0.87	1.69	1.32 a	0.13	0.10	0.17	0.04
		DS	0.93	1.65	1.30 b	0.13	0.10	0.00	‒0.07
	E3	NI	1.28	2.08	1.73 a	0.13	0.07	‒0.45	0.62
		DS	1.23	2.00	1.66 b	0.12	0.08	‒0.21	0.64
	E4	NI	1.20	2.01	1.59 a	0.11	0.07	‒0.05	0.45
		DS	1.17	2.00	1.57 b	0.13	0.08	0.28	0.74
	BLUE	NI	1.19	1.84	1.51 a	0.10	0.07	‒0.13	0.41
		DS	1.19	1.83	1.48 b	0.10	0.07	0.06	0.05
旗叶面积 FLA (cm²)	E1	NI	10.58	28.30	19.38 a	3.50	0.18	0.12	‒0.36
		DS	9.53	28.68	17.62 b	3.28	0.19	0.56	0.27
	E2	NI	8.99	29.59	17.01 a	3.99	0.23	0.71	0.28
		DS	8.30	28.89	15.99 b	3.65	0.23	0.72	0.56
	E3	NI	15.62	35.55	26.41 a	2.98	0.11	0.12	0.71
		DS	12.76	31.29	23.35 b	2.92	0.13	0.04	0.44
	E4	NI	14.96	30.80	22.36 a	2.59	0.12	0.13	0.26
		DS	11.91	32.90	20.67 b	3.65	0.18	0.48	0.31
	BLUE	NI	14.75	29.77	21.29 a	2.56	0.12	0.40	0.21
		DS	13.90	27.45	19.40 b	2.64	0.14	0.58	0.14

性状 Trait	变异来源 Source of variance	平方和 SS	均方 MS	F值 F-value	P值 P-value	遗传力 h²
FLL	基因型 Genotype (G)	16,768.48	56.08	9.26	1×10^‒5	0.77
	环境 Environment (E)	19,735.89	6578.63	1085.89	1×10^‒5
	基因型×环境 G×E	13,444.44	15.01	2.48	1×10^‒5
FLW	基因型 Genotype (G)	71.96	0.24	10.52	1×10^‒5	0.85
	环境 Environment (E)	198.12	66.04	2886.16	1×10^‒5
	基因型×环境 G×E	35.80	0.04	1.75	1×10^‒5
FLA	基因型 Genotype (G)	42,619.36	142.54	7.11	1×10^‒5	0.75
	环境 Environment (E)	138,568.38	46,189.46	2302.74	1×10^‒5
	基因型×环境 G×E	37,193.72	41.51	2.07	1×10^‒5

性状 Trait	标记名称 Marker name	单倍型 Haplotype	等位基因 Allele	频率 Frequency (%)	平均值 Mean
性状 Trait	标记名称 Marker name	单倍型 Haplotype	等位基因 Allele	频率 Frequency (%)	NI (cm)	DS (cm)
FLL	Kukri_c1406_275	FLL-Hap1	AT	77.78	18.89 a	16.93 a
		FLL-Hap2	AC	18.89	18.94 a	17.16 a
		FLL-Hap3	GC	3.33	17.43 b	15.29 b
FLA	wsnp_bq170165A_Ta_1_1	FLA-Hap1	GA	48.19	22.21 a	20.35 a
		FLA-Hap3	AA	21.01	21.27 b	19.45 b
		FLA-Hap2	GG	30.80	19.80 c	17.73 c

染色体 Chr.	位点 Marker name	处理Treatment	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
2B	IACX6324	DS	FLL	744.68	TraesCS2B01G548800	2-酮戊二酸(2OG)和Fe(II)依赖性加氧酶超家族蛋白 2-oxoglutarate (2OG) and Fe(II)-dependent oxygenase superfamily protein
3A	RAC875_c10194_673	NI	FLL, FLA	686.74	TraesCS3A01G446100	F-box家族蛋白 F-box family protein
5B	Tdurum_contig29319_256	NI	FLW	557.70	TraesCS5B01G379500	GDSL酯酶/脂肪酶 GDSL esterase/lipase
6A	RFL_Contig2206_1694	NI/DS	FLL, FLW	603.18	TraesCS6A01G385500	Myb转录因子家族蛋白 Myb family transcription factor family protein
7A	wsnp_bq170165A_Ta_1_1	NI/DS	FLA	554.49	TraesCS7A01G379000	F-box家族蛋白 F-box family protein
7B	BS00067530_51	NI/DS	FLL	648.13	TraesCS7B01G382400	F-box家族蛋白 F-box family protein

Genome-wide association analysis of morphological traits of flag leaf in wheat

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