普通小麦旗叶大小QTL定位与候选基因分析

doi:10.3864/j.issn.0578-1752.2026.18.001

中国农业科学 ›› 2026, Vol. 59 ›› Issue (18): 3949-3972.doi: 10.3864/j.issn.0578-1752.2026.18.001

• 作物遗传育种·种质资源·分子遗传学 • 上一篇下一篇

普通小麦旗叶大小QTL定位与候选基因分析

关攀锋¹(), 李盟欣¹, 万紫栋¹, 葛帅¹, 汪欣燕¹, 崔东洁¹, 池青¹, 孙昊¹, 马若男¹, 李兵¹, 许航博¹, 张世洁¹, 郭淑卿¹, 卢家玲¹, 焦浈¹, 朱育攀¹(), 魏文辉²()

¹ 郑州大学农业与生物制造学院/河南省离子束绿色农业生物工程重点实验室，郑州 450001
² 昭通学院农学与生命科学学院，云南昭通 657000

收稿日期:2026-02-03 接受日期:2026-04-01 出版日期:2026-09-16 发布日期:2026-09-20
通信作者:
朱育攀，E-mail：zhuyupan@zzu.edu.cn。
魏文辉，E-mail：20240001@ztu.edu.cn。
联系方式: 关攀锋，E-mail：guanpanfeng@zzu.edu.cn。
基金资助:
国家自然科学基金(32201862); 河南省科技攻关项目(242102111135)

QTL Mapping and Candidate Gene Analysis of Flag Leaf Size in Common Wheat (Triticum aestivum L.)

GUAN PanFeng¹(), LI MengXin¹, WAN ZiDong¹, GE Shuai¹, WANG XinYan¹, CUI DongJie¹, CHI Qing¹, SUN Hao¹, MA RuoNan¹, LI Bing¹, XU HangBo¹, ZHANG ShiJie¹, GUO ShuQing¹, LU JiaLing¹, JIAO Zhen¹, ZHU YuPan¹(), WEI WenHui²()

¹ School of Agriculture and Biomanufacturing, Zhengzhou University/Henan Key Laboratory of Ion-Beam Green Agriculture Bioengineering, Zhengzhou 450001
² College of Agronomy and Life Sciences, Zhaotong University, Zhaotong 657000, Yunnan

Received:2026-02-03 Accepted:2026-04-01 Published:2026-09-16 Online:2026-09-20

摘要/Abstract

摘要：

【目的】小麦旗叶大小是株型结构的关键特征，对光合作用、产量形成和胁迫响应至关重要。挖掘调控旗叶大小相关性状的主效QTL与候选基因，为小麦理想株型高产分子设计育种提供理论支撑与基因资源。【方法】以我国骨干型品种百农4199（BN4199）为母本、国外引进材料郑引麦2号（ZYM2）为父本，构建包含184个家系的重组自交系（recombinant inbred line，RIL）群体，利用小麦120K液相芯片（120K-4HWA）进行基因分型，并构建高密度遗传连锁图谱；基于复合区间作图法（composite interval mapping，CIM），结合2年2点4个大田环境下旗叶大小相关性状（包括旗叶长度、宽度、长宽比及面积）的表型数据和最佳线性无偏估计（best linear unbiased estimate，BLUE）值，进行全基因组QTL定位，获得稳定主效QTL并分析其加性效应；同时，结合多个小麦功能基因组学研究数据库，开展候选基因筛选和鉴定，以及组织表达模式分析，并根据QTL区间多态性SNP位点开发相应的竞争性等位基因特异性PCR标记（kompetitive allele-specific PCR，KASP）。【结果】构建了BN4199/ZYM2-RIL群体的高密度遗传连锁图谱，包含27个连锁群，3 666个SNP标记，覆盖小麦21条染色体，总图距为5 194.09 cM，标记间的平均距离为1.42 cM，遗传图谱与中国春物理图谱共线性较好，表明图谱具有较高的质量。共检测到130个QTL，包含31个旗叶长度QTL、34个旗叶宽度QTL、33个旗叶长宽比QTL和32个旗叶面积QTL，分布于除小麦3B、3D和4D外的所有染色体上；其中，位于1A、2A、2B、2D、4B、5A、7A和7B等染色体上的20个位点是多环境稳定QTL，且10个是本研究新鉴定到的位点。在20个稳定QTL区间内共注释到3 923个基因，GO分析显示，其主要富集于转录延伸因子复合物、UDP-糖基转移酶活性、节律过程、染色质介导的转录维持等条目。根据QTL区间基因功能注释，以及对应的水稻同源基因的克隆和功能鉴定，并结合小麦功能基因发掘交互式分析平台的QTG miner模块分析，共筛选出55个候选基因，其在小麦根、茎、叶、穗和籽粒不同发育时期呈现不同的表达模式。此外，开发了10个与新鉴定的QTL紧密连锁的KASP分子标记。【结论】构建了一张小麦120K液相芯片的高密度遗传连锁图谱，检测到20个控制旗叶大小相关性状的稳定QTL，其中10个为新位点，开发了与其紧密连锁的KASP分子标记，并筛选出55个潜在候选基因。

关键词: 小麦, 旗叶大小, QTL定位, 120K液相芯片, 候选基因

Abstract:

【Objective】Flag leaf size is a critical component of wheat plant architecture, playing a vital role in photosynthesis, yield formation, and stress response. The identification of major quantitative trait loci (QTL) and candidate genes regulating flag leaf size-related traits will provide theoretical support and genetic resources for the molecular design breeding of high-yielding wheat ideotypes.【Method】A recombinant inbred line (RIL) population comprising 184 lines was derived from a cross between the Chinese elite cultivar Bainong 4199 (BN4199, female parent) and the exotic variety Zhengyinmai 2 (ZYM2, male parent). Genotyping of the BN4199/ZYM2-RIL population was performed using the wheat 120K liquid chip (120K-4HWA) to construct a high-density genetic linkage map. Based on the composite interval mapping (CIM) method, QTL mapping was performed to identify stable major-effect loci and to analyze their additive effects using the flag leaf size-related traits, including flag leaf length (FLL), flag leaf width (FLW), flag leaf length/width ratio (FLR), and flag leaf area (FLA), and the best linear unbiased estimate (BLUE) values obtained across four environments (two locations over two years). Furthermore, candidate gene annotation and identification, as well as tissue expression pattern analysis, were conducted using several wheat functional genomics databases. In addition, corresponding kompetitive allele-specific PCR markers (KASP) were developed based on polymorphic SNP sites located within the stable QTL intervals.【Result】A high-density genetic linkage map of the BN4199/ZYM2-RIL population was constructed, comprising 27 linkage groups with 3 666 SNP markers, covering all 21 wheat chromosomes. The map spanned 5 194.09 cM with an average inter-marker distance of 1.42 cM. High collinearity with the Chinese Spring physical map indicated the robustness of this genetic map. A total of 130 QTL were detected, including: 31 for FLL, 34 for FLW, 33 for FLR, and 32 for FLA, distributed across all chromosomes except 3B, 3D, and 4D. Among these, 20 loci on chromosomes 1A, 2A, 2B, 2D, 4B, 5A, 7A, and 7B were environmentally stable QTL, with 10 being novel loci reported for the first time. A total of 3 923 genes were annotated within the 20 stable QTL intervals. Gene Ontology (GO) analysis revealed significant enrichment in transcription elongation factor complex, UDP-glycosyltransferase activity, rhythmic processes, and chromatin-mediated maintenance of transcription. Based on gene function annotation, information from the cloning and characterization of rice orthologs, and analysis via the QTG-miner module of an interactive wheat functional gene discovery platform, 55 candidate genes were prioritized within stable QTL intervals, exhibiting different expression patterns at various developmental stages of wheat root, stem, leaf, spike and grain tissues. In addition, 10 KASP markers closely linked to the novel QTL were developed.【Conclusion】A high-density genetic linkage map was constructed using the wheat 120K liquid chip, and 20 stable major QTL controlling flag leaf size-related traits were detected, 10 of which were novel loci. The KASP molecular markers closely linked to novel QTL were developed, and 55 potential candidate genes were identified in this study.

Key words: wheat, flag leaf size, QTL mapping, 120K liquid chip, candidate genes

关攀锋, 李盟欣, 万紫栋, 葛帅, 汪欣燕, 崔东洁, 池青, 孙昊, 马若男, 李兵, 许航博, 张世洁, 郭淑卿, 卢家玲, 焦浈, 朱育攀, 魏文辉. 普通小麦旗叶大小QTL定位与候选基因分析[J]. 中国农业科学, 2026, 59(18): 3949-3972.

GUAN PanFeng, LI MengXin, WAN ZiDong, GE Shuai, WANG XinYan, CUI DongJie, CHI Qing, SUN Hao, MA RuoNan, LI Bing, XU HangBo, ZHANG ShiJie, GUO ShuQing, LU JiaLing, JIAO Zhen, ZHU YuPan, WEI WenHui. QTL Mapping and Candidate Gene Analysis of Flag Leaf Size in Common Wheat (Triticum aestivum L.)[J]. Scientia Agricultura Sinica, 2026, 59(18): 3949-3972.

图/表 13

图1

表1

表2

图2

表3

图3

表4

BN4199/ZYM2-RIL群体旗叶大小相关性状环境稳定QTL位点鉴定"

性状 Trait	QTL	染色体 Chromosome	环境 Environment	峰值标记位置 Peak marker position (cM)	LOD	表型变异率 PVE (%)	加性效应 Additive effect	遗传区间 Genetic interval (cM)	物理区间^a Physical interval (Mb)	参考文献 Reference
旗叶长度 Flag leaf length	QFll.zzu-1A.2	1A	E1	197.11	2.94	5	-0.49	192.4-210.0	576.6-584.7	[46]
		1A	E3	204.61	3.41	5	-0.56
		1A	BLUE	204.61	2.91	4	-0.35
	QFll.zzu-2B.1	2B	E1	77.01	9.02	15	0.85	74.3-80.5	60.7-72.7	[22,41,47]
		2B	E2	77.01	7.80	10	0.79
		2B	E3	77.01	9.29	15	0.98
		2B	E4	77.01	6.21	12	0.52
		2B	BLUE	77.01	11.59	16	0.75
	QFll.zzu-2D.1	2D	E2	23.51	2.80	4	0.53	20.0-27.1	18.7-23.1	[9,19,48]
	QFll.zzu-2D.1	2D	E3	23.51	2.91	4	0.58	20.0-27.1	18.7-23.1	[9,19,48]
	QFll.zzu-2D.2	2D	E1	43.91	9.69	19	0.96	31.9-46.5	26.8-35.2	[49-51]
		2D	E2	43.91	20.95	40	1.57
		2D	E3	36.91	8.08	16	0.98
		2D	BLUE	41.91	14.25	25	0.91
	QFll.zzu-7B.4	7B	E1	25.11	2.99	5	-0.47	23.9-28.1	477.2-597.6	New
		7B	E4	24.51	2.83	6	-0.40
		7B	BLUE	24.51	3.10	4	-0.37
旗叶宽度 Flag leaf width	QFlw.zzu-2D	2D	E1	43.91	3.35	7	0.04	36.9-52.1	26.8-40.4	[49-51]
	QFlw.zzu-2D	2D	E2	47.51	8.61	16	0.08	36.9-52.1	26.8-40.4	[49-51]
	QFlw.zzu-4B.1	4B	E1	37.31	5.53	10	0.05	37.0-41.3	34.4-38.7	New
	QFlw.zzu-4B.1	4B	E3	39.31	6.79	14	0.06	37.0-41.3	34.4-38.7	New
	QFlw.zzu-4B.2	4B	E1	51.81	7.53	13	0.06	47.6-54.3	47.2-97.2	New
		4B	E3	48.31	9.16	17	0.06
		4B	BLUE	51.81	3.64	7	0.04
	QFlw.zzu-4B.3	4B	E1	58.51	6.65	12	0.05	55.1-60.2	101.0-152.2	New
		4B	E3	57.81	5.68	12	0.05
		4B	BLUE	58.51	3.24	6	0.04
	QFlw.zzu-5A.1	5A	E1	136.21	3.73	7	-0.04	131.7-137.9	503.4-509.9	New
	QFlw.zzu-5A.1	5A	E3	133.61	4.10	8	-0.04	131.7-137.9	503.4-509.9	New
	QFlw.zzu-5A.3	5A	E3	189.11	3.30	7	0.04	184.2-197	579.7-611.9	[52]
	QFlw.zzu-5A.3	5A	E4	186.11	5.31	11	0.05	184.2-197	579.7-611.9	[52]
	QFlw.zzu-7A.1	7A	E1	102.21	4.35	8	-0.04	92.6-107.6	68.0-82.8	New
		7A	E2	96.81	5.72	11	-0.07
		7A	BLUE	97.81	3.47	7	-0.04
	QFlw.zzu-7A.2	7A	E1	110.61	2.98	5	-0.03	108.3-116.6	82.8-86.6	New
		7A	E2	110.31	3.69	6	-0.05
		7A	BLUE	110.61	2.66	5	-0.03
旗叶长宽比 Flag leaf length/ width ratio	QFlr.zzu-2A.3	2A	E1	131.81	3.19	6	-0.31	128.7-133.7	559.9-568.9	New
		2A	E4	130.31	3.34	6	-0.36
		2A	BLUE	130.31	5.43	10	-0.58
	QFlr.zzu-2D.1	2D	E1	7.31	2.77	5	0.30	5.6-8.5	11.7-13.4	New
		2D	E2	6.31	5.95	12	0.44
		2D	E3	6.31	5.25	10	0.55
		2D	BLUE	6.31	5.62	10	0.40
	QFlr.zzu-2D.2	2D	E1	16.21	5.55	10	0.42	11.6-18.9	14.6-18.3	[9,19,48]
		2D	E2	13.61	2.94	7	0.33
		2D	E3	16.21	7.58	14	0.65
		2D	E4	15.61	4.17	9	0.41
		2D	BLUE	15.61	6.73	13	0.45
	QFlr.zzu-7A	7A	E1	187.81	3.11	6	0.31	185.9-189.1	515.6-516.9	New
	QFlr.zzu-7A	7A	E2	187.51	6.33	14	0.47	185.9-189.1	515.6-516.9	New
旗叶面积 Flag leaf area	QFla.zzu-2B.1	2B	E1	72.01	4.82	8	1.39	67.3-78.7	50.6-69.9	[22,41,47]
		2B	E2	68.41	5.39	8	1.96
		2B	E3	76.31	5.92	11	1.69
		2B	E4	77.01	4.89	10	0.92
	QFla.zzu-2B.2	2B	E1	82.31	6.38	10	1.51	78.9-87.6	72.7-76.9	[22,41,47]
		2B	E2	85.71	6.99	10	2.17
		2B	E3	84.71	7.16	13	1.77
		2B	E4	84.71	3.40	7	0.78
		2B	BLUE	80.71	8.40	14	1.51
	QFla.zzu-2D.1	2D	E1	42.91	8.50	17	1.91	34.1-53.3	26.8-40.4	[49-51]
		2D	E2	45.91	20.56	33	3.92
		2D	E3	41.91	3.42	8	1.37
		2D	BLUE	43.91	10.02	18	1.72

表4

图4

图5

图6

表5

候选基因的鉴定及功能注释"

小麦基因ID Gene ID of wheat (IWGSC CS RefSeqv2.1)	功能描述 Functional description	水稻基因ID Gene ID of rice (IRGSP 1.0)	水稻同源基因功能分析 Functional characterization of rice orthologous genes
TraesCS1A03G1028100	尿酸降解双功能蛋白TTL Uric acid degradation bifunctional protein TTL	Os03g0390700	剑叶长度、剑叶宽度、剑叶面积、叶夹角 Flag leaf length, flag leaf width, flag leaf area, leaf angle
TraesCS1A03G1030600	痉挛蛋白 (Spastin) Spastin	Os05g0584600	叶夹角Leaf angle
TraesCS1A03G1034900	茉莉酰-L-氨基酸合成酶GH3.5 Jasmonoyl-L-amino acid synthetase GH3.5	Os05g0586200	剑叶宽度、叶夹角、开花时间 Flag leaf width, leaf angle, flowering time
TraesCS2A03G0780200	五肽重复序列蛋白(PPR) At1g09900 Pentatricopeptide repeat-containing protein At1g09900	Os02g0565400	叶发育、叶绿素 Leaf development, chlorophyll
TraesCS2A03G0793800	隐花色素-1(蓝光受体) Cryptochrome-1	Os02g0573200	赤霉素响应、叶发育 Gibberellin response, leaf development
TraesCS2A03G0801300	LRR类受体激酶SERK2 LRR receptor kinase SERK2	Os04g0457800	叶夹角、细胞分裂 Leaf angle, cell division
TraesCS2A03G0808100	含有NAC结构域蛋白100 NAC domain-containing protein 100	Os04g0460600	抽穗期调控、维管束发育、叶绿素 Heading date regulation, vascular bundle development, chlorophyll
TraesCS2A03G0810600	邻氨基苯甲酸合成酶β亚基1（叶绿体型） Anthranilate synthase beta subunit 1 (chloroplastic)	Os04g0463500	剑叶宽度、叶夹角、叶绿素合成 Flag leaf width, leaf angle, chlorophyll synthesis
TraesCS2B03G0201200	UDP-糖基转移酶79 UDP-glycosyltransferase 79	Os04g0206700	抽穗期调控、剑叶长度 Heading date regulation, flag leaf length
TraesCS2B03G0205100	类萌发蛋白8-14 Germin-like protein 8-14	Os08g0460000	叶夹角 Leaf angle
TraesCS2B03G0222600	二组分响应调节类蛋白PRR37 Two-component response regulator-like PRR37	NA	抽穗期调控 Heading date regulation
TraesCS2B03G0223300	L-抗坏血酸过氧化物酶2（胞质型） L-ascorbate peroxidase 2 (cytosolic)	Os07g0694700	剑叶长度、剑叶宽度、叶夹角 Flag leaf length, flag leaf width, leaf angle
TraesCS2D03G0084300	UDP-糖基转移酶79 UDP-glycosyltransferase 79	Os04g0206700	抽穗期调控、剑叶长度 Heading date regulation, flag leaf length
TraesCS2D03G0099000	三羟基黄酮3',4',5'-O-三甲基转移酶 Tricetin 3',4',5'-O-trimethyltransferase	Os08g0157500	剑叶长度、剑叶宽度、光合作用、维管束发育、叶绿素 Flag leaf length, flag leaf width, photosynthesis, vascular bundle development, chlorophyll
TraesCS2D03G0142400	UDP-葡萄糖基转移酶UGT13248 UDP-glucosyltransferase UGT13248	Os04g0206700	抽穗期调控、剑叶长度 Heading date regulation, flag leaf length
TraesCS2D03G0144400	类萌发蛋白8-14 Germin-like protein 8-14	Os08g0460000	叶夹角 Leaf angle
TraesCS2D03G0144600	类萌发蛋白8-14 Germin-like protein 8-14	Os08g0460000	叶夹角 Leaf angle
TraesCS2D03G0156800	二组分响应调节类蛋白PRR37 Two-component response regulator-like PRR37	Os07g0695100	抽穗期调控 Heading date regulation
TraesCS2D03G0157900	L-抗坏血酸过氧化物酶2（胞质型） L-ascorbate peroxidase 2 (cytosolic)	Os07g0694700	剑叶长度、剑叶宽度、叶夹角 Flag leaf length, flag leaf width, leaf angle
TraesCS4B03G0116600	类SCARECROW蛋白7（SCL7） SCARECROW-like protein 7	Os03g0723000	剑叶长度、剑叶宽度、叶夹角、细胞分裂素 Flag leaf length, flag leaf width, leaf angle, cytokinin
TraesCS4B03G0117300	未鉴定糖激酶slr0537 Uncharacterized sugar kinase slr0537	Os01g0105900	剑叶长度、剑叶宽度、叶夹角、抽穗期调控、维管束发育 Flag leaf length, flag leaf width, leaf angle, heading date regulation, vascular bundle development
TraesCS4B03G0121800	7-甲基-GTP焦磷酸酶 7-methyl-GTP pyrophosphatase	Os03g0724700	剑叶宽度、赤霉素响应 Flag leaf width, gibberellin response
TraesCS4B03G0136900	类BEL1同源结构域蛋白6 BEL1-like homeodomain protein 6	Os03g0732100	叶大小、抽穗期调控、叶绿素、赤霉素响应 Leaf size, heading date regulation, chlorophyll, gibberellin response
TraesCS4B03G0139300	GRF相互作用因子1 GRF-interacting factor 1	Os03g0733600	剑叶长度、剑叶宽度、维管束发育 Flag leaf length, flag leaf width, vascular bundle development
TraesCS4B03G0155100	休眠相关蛋白同源物2 Dormancy-associated protein homolog 2	Os11g0671000	剑叶长度、抽穗期调控 Flag leaf length, heading date regulation
TraesCS4B03G0160200	胞吐复合物组分SEC3A Exocyst complex component SEC3A	Os03g0625700	叶发育、叶绿素合成 Leaf development, chlorophyll synthesis
TraesCS4B03G0254200	60S核糖体蛋白L3 60S ribosomal protein L3	Os11g0168200	抽穗期调控、叶大小 Heading date regulation, leaf size
TraesCS4B03G0255000	脱落酸胁迫成熟蛋白5（ASR5） Abscisic stress-ripening protein 5	Os11g0167800	叶发育、赤霉素响应 Leaf development, gibberellin response
TraesCS4B03G0269400	信号识别颗粒54 kDa蛋白（叶绿体型） Signal recognition particle 54 kDa protein (chloroplastic)	Os11g0153600	抽穗期调控、叶绿素、光合作用 Heading date regulation, chlorophyll, photosynthesis
TraesCS4B03G0269500	信号识别颗粒54 kDa蛋白（叶绿体型） Signal recognition particle 54 kDa protein (chloroplastic)	Os11g0153700	叶发育、叶绿素 Leaf development, chlorophyll
TraesCS5A03G0934800	光敏色素C Phytochrome C	Os03g0752100	抽穗期调控、叶夹角、叶绿素、赤霉素响应 Heading date regulation, leaf angle, chlorophyll, gibberellin response
TraesCS5A03G0949400	赤霉素调节蛋白8 Gibberellin-regulated protein 8	Os03g0760800	剑叶长度、赤霉素响应 Flag leaf length, gibberellin response
TraesCS5A03G0949800	抗菌肽 Snakin-1Snakin-1	Os03g0760800	剑叶长度、赤霉素响应 Flag leaf length, gibberellin response
TraesCS5A03G0949900	抗菌肽 Snakin-1Snakin-1	Os03g0760800	剑叶长度、赤霉素响应 Flag leaf length, gibberellin response
TraesCS5A03G0956200	Dof锌指蛋白DOF1.1 Dof zinc finger protein DOF1.1	Os03g0764900	剑叶宽度、抽穗期调控、细胞分裂素 Flag leaf width, heading date regulation, cytokinin
TraesCS5A03G0972700	富含脯氨酸类受体蛋白激酶 PERK4Proline-rich receptor-like protein kinase PERK4	Os03g0821900	叶夹角 Leaf angle
TraesCS5A03G0981000	光系统 I 反应中心亚基Ⅲ（叶绿体型） Photosystem I reaction center subunit Ⅲ (chloroplastic)	Os03g0778100	叶绿素、光合电子传递链 Chlorophyll, photosynthetic electron transport chain
TraesCS7A03G0272100	抽穗期相关蛋白3A（Hd3a） Protein HEADING DATE 3A	Os06g0157700	抽穗期调控 Heading date regulation
TraesCS7A03G0284900	类富含脯氨酸细胞壁蛋白 Proline-rich cell wall protein-like protein	Os06g0160400	抽穗期调控、细胞分裂素 Heading date regulation, cytokinin
TraesCS7A03G0856100	推测的E3泛素蛋白连接酶 UBR7Putative E3 ubiquitin-protein ligase UBR7	Os06g0529800	叶发育、细胞分裂素 Leaf development, cytokinin
TraesCS7B03G0705500	棕榈酰-酰基载体蛋白硫酯酶（叶绿体型） Palmitoyl-acyl carrier protein thioesterase (chloroplastic)	Os06g0596300	叶夹角、叶绿素合成 Leaf angle, chlorophyll synthesis
TraesCS7B03G0707900	Argonaute蛋白 PNH1 (PINHEAD)Protein argonaute PNH1	Os06g0597400	叶发育、维管束发育 Leaf development, vascular bundle development
TraesCS7B03G0719700	组蛋白去乙酰化酶 1Histone deacetylase 1	Os06g0583400	剑叶长度、剑叶宽度、叶夹角 Flag leaf length, flag leaf width, leaf angle
TraesCS7B03G0732300	细胞色素P450 711A1 Cytochrome P450 711A1	Os06g0565100	叶片大小与形状调控 Regulation of leaf size and shape
TraesCS7B03G0737400	脱落酸受体PYL9 Abscisic acid receptor PYL9	Os06g0562200	叶绿素、维管束发育 Chlorophyll, vascular bundle development
TraesCS7B03G0745700	开花位点蛋白T（FT蛋白） Protein FLOWERING LOCUS T	Os06g0552900	抽穗期调控、卷叶叶夹角 Heading date regulation, leaf rolling and leaf angle
TraesCS7B03G0757900	血红素加氧酶1（叶绿体型） Heme oxygenase 1 (chloroplastic)	Os06g0603000	抽穗期调控、叶绿素 Heading date regulation, chlorophyll
TraesCS7B03G0771700	果糖-1,6-二磷酸醛缩酶5（胞质型）Fructose-bisphosphate aldolase 5 (cytosolic)	Os06g0608700	叶绿素、光合作用 Chlorophyll, photosynthesis
TraesCS7B03G0780400	Ras相关蛋白 RABA3Ras-related protein RABA3	Os06g0714600	叶发育、叶夹角 Leaf development, leaf angle
TraesCS7B03G0797500	基质加工肽酶（叶绿体型） Stromal processing peptidase (chloroplastic)	Os06g0625400	叶发育、叶绿素 Leaf development, chlorophyll
TraesCS7B03G0824900	含有五肽重复序列的蛋白At5g42310（叶绿体型） Pentatricopeptide repeat-containing protein At5g42310 (chloroplastic)	Os06g0710800	净光合速率、叶绿素 Net photosynthetic rate, chlorophyll
TraesCS7B03G0826400	F-box蛋白SKIP31 F-box protein SKIP31	Os06g0709100	叶大小、叶夹角 Leaf size, leaf angle
TraesCS7B03G0845600	含有CCCH锌指结构域蛋白46 Zinc finger CCCH domain-containing protein 46	Os06g0704300	叶发育、叶夹角、细胞大小 Leaf development, leaf angle, cell size
TraesCS7B03G0850100	生长素响应因子19（ARF19） Auxin response factor 19	Os06g0702600	剑叶长度、剑叶宽度、叶夹角、叶绿素合成 Flag leaf length, flag leaf width, leaf angle, chlorophyll synthesis
TraesCS7B03G0870400	富含亮氨酸重复序列蛋白2（LRR蛋白） Leucine-rich repeat protein 2	Os03g0440900	叶发育 Leaf development

表5

图7

表6

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性状 Trait	环境 Environment	亲本Parents		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
性状 Trait	环境 Environment	郑引麦2号 ZYM2	百农4199 BN4199	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
旗叶长度 Flag leaf length (cm)	E1	23.74	18.78	15.63	25.29	19.73	1.74	8.8	0.56	0.71
	E2	23.77	21.39	18.29	27.14	22.17	1.70	7.7	0.32	0.18
	E3	23.57	17.72	18.11	29.94	23.67	2.26	9.5	0.10	-0.24
	E4	21.00	12.82	13.11	20.67	16.61	1.46	8.8	0.34	0.01
	BLUE	22.34	20.04	16.02	24.64	20.28	1.82	9.0	0.21	-0.36
旗叶宽度 Flag leaf width (cm)	E1	1.83	1.71	1.29	2.16	1.67	0.12	7.2	0.57	1.82
	E2	1.95	1.78	1.62	2.26	1.88	0.13	7.1	0.41	-0.04
	E3	1.87	1.68	1.41	2.11	1.71	0.14	8.2	0.54	0.45
	E4	1.66	1.37	1.12	1.88	1.47	0.15	9.9	0.02	-0.17
	BLUE	1.98	1.81	1.33	2.12	1.70	0.14	8.3	0.34	0.10
旗叶长宽比 Flag leaf length/width ratio	E1	13.02	11.13	9.22	15.36	11.96	1.02	8.5	0.10	0.65
	E2	12.24	12.10	9.23	14.10	11.87	0.88	7.4	-0.13	-0.12
	E3	12.72	10.56	10.39	19.87	14.08	1.59	11.3	0.21	0.74
	E4	12.85	9.49	7.63	16.90	11.62	1.43	12.3	0.48	0.86
	BLUE	10.60	9.69	8.98	16.18	12.18	1.22	10.0	0.18	0.41
旗叶面积 Flag leaf area (cm²)	E1	36.23	26.81	16.75	37.72	27.52	3.75	13.6	0.54	0.34
	E2	39.43	32.10	25.77	48.20	35.13	4.52	12.9	0.50	0.02
	E3	37.07	25.54	24.19	47.55	33.63	4.60	13.7	0.35	-0.04
	E4	29.45	15.07	14.03	27.99	20.59	3.08	15.0	0.35	-0.42
	BLUE	38.61	31.27	20.06	41.10	28.54	3.96	13.9	0.44	0.08

性状Trait	方差来源Source of variance	自由度df	平方和SS	均方MS	F值F value	遗传力H² (%)
旗叶长度 Flag leaf length	基因型效应Genotype (G)	183	6706.14	36.65	12.02**	86
	环境效应Environment (E)	3	15041.74	5013.91	1645.15**
	基因型×环境互作效应G×E	530	2930.23	5.53	1.81**
	区组/环境内区组效应Block/Env	8	540.94	67.62	22.19**
	误差Error	1363	4154.01	3.05
旗叶宽度 Flag leaf width	基因型效应Genotype (G)	183	36.53	0.20	10.67**	85
	环境效应Environment (E)	3	41.13	13.71	732.68**
	基因型×环境互作效应G×E	530	17.32	0.03	1.75**
	区组/环境内区组效应Block/Env	8	2.05	0.26	13.67**
	误差Error	1363	25.51	0.02
旗叶长宽比 Flag leaf length/ width ratio	基因型效应Genotype (G)	183	3007.74	16.44	12.26**	88
	环境效应Environment (E)	3	2141.34	713.78	532.45**
	基因型×环境互作效应G×E	530	1108.24	2.09	1.56**
	区组/环境内区组效应Block/Env	8	330.69	41.34	30.83**
	误差Error	1363	1827.19	1.34
旗叶面积 Flag leaf area	基因型效应Genotype (G)	183	33263.02	181.77	11.29**	83
	环境效应Environment (E)	3	66355.80	22118.60	1373.42**
	基因型×环境互作效应G×E	530	17523.87	33.06	2.05**
	区组/环境内区组效应Block/Env	8	1722.99	215.37	13.37**
	误差Error	1363	21950.84	16.10

染色体 Chromosome	连锁群 Linkage group	差异SNP标记数量 No. of differential SNP markers	BIN标记数量 No. of BIN markers	图谱位点数量 No. of map loci	遗传长度 Genetic length (cM)	平均距离 Mean distance (cM)
1A	LG1A	2161	536	219	244.05	1.11
1B	LG1B	2891	679	256	285.06	1.11
1D	LG1D	540	377	148	292.10	1.97
2A	LG2A	3072	886	258	230.24	0.89
2B	LG2B	3841	1013	432	416.97	0.97
2D	LG2D.1	1245	118	80	172.85	2.16
2D	LG2D.2	1245	49	19	92.02	4.84
3A	LG3A.1	2203	205	85	68.61	0.81
3A	LG3A.2	2203	188	94	196.61	2.09
3B	LG3B	5123	743	241	310.09	1.29
3D	LG3D	360	243	102	318.94	3.13
4A	LG4A	2248	257	131	173.27	1.32
4B	LG4B	2522	447	177	217.84	1.23
4D	LG4D	109	26	22	57.88	2.63
5A	LG5A	2081	286	130	225.18	1.73
5B	LG5B	2667	573	263	351.75	1.34
5D	LG5D.1	438	27	15	57.42	3.83
5D	LG5D.2	438	69	27	27.42	1.02
6A	LG6A	2692	579	191	239.86	1.26
6B	LG6B	2493	428	210	238.27	1.13
6D	LG6D.1	386	57	20	27.29	1.36
6D	LG6D.2	386	175	95	229.66	2.42
7A	LG7A.1	3765	443	160	247.75	1.55
7A	LG7A.2	3765	88	43	51.30	1.19
7B	LG7B	2925	333	142	158.95	1.12
7D	LG7D.1	902	122	83	224.16	2.70
7D	LG7D.2	902	78	23	38.59	1.68
亚基因组Subgenome
A		18222	3468	1311	1676.86	1.28
B		22462	4216	1721	1978.92	1.15
D		3980	1341	634	1538.31	2.43
同源群Homoeologous group
1		5592	1592	623	821.21	1.32
2		8158	2066	789	912.09	1.16
3		7686	1379	522	894.24	1.71
4		4879	730	330	448.99	1.36
5		5186	955	435	661.76	1.52
6		5571	1239	516	735.07	1.42
7		7592	1064	451	720.74	1.60
总数Sum		44664	9025	3666	5194.09	1.42

序号 No.	QTL	标记名称 Marker name	引物标签 Primer tag	引物序列 Primer sequences (5′-3′)
1	QFlr.zzu-2A.3	SNP7433_2A	primer_X	GAAGGTGACCAAGTTCATGCTGCAGGAAGAGCATGTAGCGCT
			primer_Y	GAAGGTCGGAGTCAACGGATTGCAGGAAGAGCATGTAGCGCA
			primer_C	ACGGGCGTTCATCCGGGCCA
2	QFlr.zzu-2D.1	SNP12582_2D	primer_X	GAAGGTGACCAAGTTCATGCTGCGTGCTGGTGGTCTGCTC
			primer_Y	GAAGGTCGGAGTCAACGGATTCGCGTGCTGGTGGTCTGCTT
			primer_C	AGCAGACCACCAGCACGCGC
3	QFlw.zzu-4B.1	SNP23868_4B	primer_X	GAAGGTGACCAAGTTCATGCTGCATGCGGTCAACATGCTTTTGATA
			primer_Y	GAAGGTCGGAGTCAACGGATTGCATGCGGTCAACATGCTTTTGATT
			primer_C	TGTTTATAGCGACCATCAATCCTTGAAG
4	QFlw.zzu-4B.2	SNP24149_4B	primer_X	GAAGGTGACCAAGTTCATGCTTGTTTTCCGGTACTGGAGACAGA
			primer_Y	GAAGGTCGGAGTCAACGGATTGTTTTCCGGTACTGGAGACAGG
			primer_C	CGCCGAGGCGTACTACGGTG
5	QFlw.zzu-4B.3	SNP24289_4B	primer_X	GAAGGTGACCAAGTTCATGCTCAAGCACTTCGTCACGGTGC
			primer_Y	GAAGGTCGGAGTCAACGGATTCCAAGCACTTCGTCACGGTGT
			primer_C	CAGTGTTGTTCCTGAGCTTGAACTC
6	QFlw.zzu-5A.1	SNP27278_5A	primer_X	GAAGGTGACCAAGTTCATGCTTGATTGGCCTGAACCAGAATCAC
			primer_Y	GAAGGTCGGAGTCAACGGATTCTTGATTGGCCTGAACCAGAATCAT
			primer_C	TATTGCCGGGAGGGATGGAACC
7	QFlw.zzu-7A.1	SNP37923_7A	primer_X	GAAGGTGACCAAGTTCATGCTTCCCTCCCCGTTGGCAATAC
			primer_Y	GAAGGTCGGAGTCAACGGATTCTCCCTCCCCGTTGGCAATAT
			primer_C	TTGCCCTCCTGACCCGATGGTT
8	QFlw.zzu-7A.2	SNP37968_7A	primer_X	GAAGGTGACCAAGTTCATGCTGAATAACACCACCATCTGACCCTTA
			primer_Y	GAAGGTCGGAGTCAACGGATTGAATAACACCACCATCTGACCCTTT
			primer_C	GGTTGGAGGAAGTTCTATGAAGACAAC
9	QFlr.zzu-7A	SNP39426_7A	primer_X	GAAGGTGACCAAGTTCATGCTGTTCCTGAGCTTGAACTCCGTC
			primer_Y	GAAGGTCGGAGTCAACGGATTTGTTCCTGAGCTTGAACTCCGTT
			primer_C	TCACCAAGCACTTCGTCGCGGT
10	QFll.zzu-7B.4	SNP42098_7B	primer_X	GAAGGTGACCAAGTTCATGCTATATGAATGCGCATGAAGCCCAC
			primer_Y	GAAGGTCGGAGTCAACGGATTCATATGAATGCGCATGAAGCCCAA
			primer_C	GGTGATCTTCGATGACACCAACTATG

普通小麦旗叶大小QTL定位与候选基因分析

QTL Mapping and Candidate Gene Analysis of Flag Leaf Size in Common Wheat (Triticum aestivum L.)

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