小麦旗叶叶绿素含量的QTL定位及验证

doi:10.3724/SP.J.1006.2023.21018

作物学报 ›› 2023, Vol. 49 ›› Issue (3): 744-754.doi: 10.3724/SP.J.1006.2023.21018

• 作物遗传育种·种质资源·分子遗传学 • 上一篇下一篇

小麦旗叶叶绿素含量的QTL定位及验证

杨斌(), 乔玲(), 赵佳佳, 武棒棒, 温宏伟, 张树伟, 郑兴卫, 郑军()

山西农业大学小麦研究所 / 省部共建有机旱作农业国家重点实验室(筹), 山西临汾 041000

收稿日期:2022-02-25 接受日期:2022-06-07 出版日期:2023-03-12 网络出版日期:2022-07-07
通讯作者: 郑军
作者简介:杨斌, E-mail: sxxmsyb83@126.com
乔玲, E-mail: qiaolingsmile@163.com第一联系人：^**同等贡献
基金资助:
中央引导地方科技发展资金项目(YDZJSX2022A033);山西农业大学育种工程(YZGC013);山西省基础研究计划项目(202103021223156)

QTL mapping and validation of chlorophyll content of flag leaves in wheat (Triticum aestivum L.)

YANG Bin(), QIAO Ling(), ZHAO Jia-Jia, WU Bang-Bang, WEN Hong-Wei, ZHANG Shu-Wei, ZHENG Xing-Wei, ZHENG Jun()

Institute of Wheat Research, Shanxi Agricultural University / State Key Laboratory of Sustainable Dryland Agriculture (in preparation), Linfen 041000, Shanxi, China

Received:2022-02-25 Accepted:2022-06-07 Published:2023-03-12 Published online:2022-07-07
Contact: ZHENG Jun
About author:First author contact:^**Contributed equally to this work
Supported by:
Central Guidance on Science & Technology Development of Shanxi(YDZJSX2022A033);Shanxi Agricultural University Breeding Project(YZGC013);Basic Research Program of Shanxi Province(202103021223156)

摘要/Abstract

摘要：

旗叶是小麦主要的光合器官, 叶绿素既是旗叶最主要的光合色素, 也是品种选育中重要的表型指标, 因此挖掘和利用旗叶叶绿素含量有关的主效基因/位点, 对于培育高产稳产小麦新品种意义重大。以旗叶叶绿素含量差异较大双亲构建的双单倍体群体(DH群体)为材料, 利用小麦90K SNP芯片对5个环境旗叶叶绿素含量进行QTL分析, 共定位到20个旗叶叶绿素含量有关的遗传位点, 表型贡献率为4.10%~27.16%; 其中3个QTL (Qchl.saw-2D.1、Qchl.saw-4D.2和Qchl.saw-6A)能在多个环境条件下检测到; Qchl.saw-2D.1的遗传效应最高, 该位点与2D染色体上已报道的其他叶绿素位点不同, 初步确定是1个新的主效QTL。并进一步将Qchl.saw-2D.1紧密连锁的SNP标记开发为KASP标记, 通过在含有共同亲本金麦919的RIL群体中验证其效应, 发现在多个环境条件下具有Qchl.saw-2D.1有利等位基因的家系叶绿素含量显著或极显著高于其他家系。对Qchl.saw-2D.1、Qchl.saw-4D.2和Qchl.saw-6A所在功能区段进行基因注释, 筛选到12个与叶绿素相关的候选基因, 其中3个基因参与镁等金属离子的结合过程, 5个基因参与调控叶绿体结构组成, 4个基因参与调控光合作用过程中相关电子链的传递活性。本文研究结果为叶绿素调控的遗传机制提供了有价值的信息, 并为高光效分子标记辅助育种提供依据与参考。

关键词: 小麦, 叶绿素含量, SNP标记, QTL定位

Abstract:

Flag leaf is the main photosynthetic organ in wheat. The chlorophyll content is not only the major photosynthetic pigment in flag leaf but also an important phenotypic indicator in crop breeding. Therefore, the identification of major loci/genes related to chlorophyll content in the flag leaf play an important role in breeding wheat varieties with higher grain yields and stability. In this study, we constructed a double haploid (DH) population from a cross of two cultivars with significant difference in chlorophyll content, and the chlorophyll contents of DH lines were detected under five environments. A total of 20 QTLs associated with chlorophyll content were detected using Wheat 90K single-nucleotide polymorphism (SNP) array, with contributions to phenotypic variation explained (PVE) from 4.10% to 27.16%. Three QTLs (Qchl.saw-2D.1, Qchl.saw-4D.2, and Qchl.saw-6A) were identified under multiple environmental conditions, in which Qchl.saw-2D.1 with the strongest genetic effect was different from previous studies and identified as a novel major QTL. Furthermore, Qchl.saw-2D.1 was validated by a tightly linked kompetitive allele specific PCR (KASP) marker in a recombinant inbred line (RIL) population containing the co-parent Jinmai 919. Those lines with the favorable allele of Qchl.saw-2D.1 revealed significantly higher chlorophyll content than other lines under multiple environments. Moreover, a total of 12 candidate genes controlling chlorophyll content were identified in the three QTL regions. Based on gene annotation, three genes were involved in the binding process of metal iron, such as magnesium. Five genes were regulated the structural composition of chloroplasts, and four genes were engaged in the regulation of electron transfer activities during the photosynthetic process. In conclusion, this study will broaden the understanding of the genetic mechanism and provide a molecular basis for the marker-assisted breeding (MAS) of chlorophyll content in the flag leaf of wheat.

Key words: wheat, chlorophyll content, SNP marker, QTL mapping

杨斌, 乔玲, 赵佳佳, 武棒棒, 温宏伟, 张树伟, 郑兴卫, 郑军. 小麦旗叶叶绿素含量的QTL定位及验证[J]. 作物学报, 2023, 49(3): 744-754.

YANG Bin, QIAO Ling, ZHAO Jia-Jia, WU Bang-Bang, WEN Hong-Wei, ZHANG Shu-Wei, ZHENG Xing-Wei, ZHENG Jun. QTL mapping and validation of chlorophyll content of flag leaves in wheat (Triticum aestivum L.)[J]. Acta Agronomica Sinica, 2023, 49(3): 744-754.

图/表 7

表1

图1

表2

DH群体中检测到的叶绿素含量QTL"

QTL	环境 Environment	染色体 Chr.	LOD值 LOD value	表型贡献率 PVE (%)	加性效应 Additive	左标记 Left marker	右标记 Right marker	遗传位置 Genetic interval (cM)	物理位置 Physical interval (Mb)
Qchl.saw-2A.1	E4	2A	2.55	4.71	‒0.64	Excalibur_c18630_283	Kukri_c13187_556	0.00-2.50	18.30-32.53
Qchl.saw-2A.2	E2	2A	3.33	7.10	‒0.69	Kukri_rep_c106992_239	BS00041816_51	46.70-48.30	611.30-611.32
Qchl.saw-2A.3	E2	2A	4.98	10.33	‒0.84	TA005399-1698	Ra_c7093_381	52.60-54.60	685.96-695.51
Qchl.saw-2A.4	E2	2A	3.04	7.75	‒0.73	Kukri_c47259_416	RAC875_c37510_329	62.30-65.30	717.19-727.17
Qchl.saw-2B.1	E3	2B	2.95	6.97	0.59	BS00076596_51	BS00077463_51	13.30-18.20	8.34-13.76
Qchl.saw-2B.2	E2	2B	4.01	8.27	0.74	BS00105036_51	GENE-1343_556	23.60-30.20	18.18-24.10
Qchl.saw-2B.3	E4	2B	2.74	4.60	‒0.65	Kukri_c43403_346	BS00068550_51	59.90-65.50	76.82-159.60
	E5	2B	4.34	7.85	‒0.66	Kukri_c43403_346	BS00068550_51	58.50-64.30	76.82-159.60
Qchl.saw-2D.1	E3	2D	3.69	8.21	‒0.66	BS00012148_51	Excalibur_c18324_390	0.00-1.60	3.77-7.97
	E4	2D	6.84	14.03	‒1.11	BS00012148_51	Excalibur_c18324_390	1.10-1.60	3.77-7.97
	E5	2D	12.85	26.07	‒1.19	BS00012148_51	Excalibur_c18324_390	1.50-1.80	3.77-7.97
	BLUP	2D	9.84	23.15	‒0.78	BS00012148_51	Excalibur_c18324_390	1.40-2.10	3.77-7.97
Qchl.saw-2D.2	E1	2D	2.66	7.00	‒1.12	BobWhite_c35859_105	GENE-1086_99	2.10-3.30	9.93-12.33
	E5	2D	8.81	19.64	‒1.22	BobWhite_c35859_105	GENE-1086_99	2.20-3.20	9.93-12.33
Qchl.saw-2D.3	E5	2D	13.60	27.16	‒1.23	BobWhite_rep_c64429_660	D_contig04712_355	5.30-6.70	20.63-22.14
	BLUP	2D	9.09	22.18	‒0.76	BobWhite_rep_c64429_660	D_contig04712_355	5.60-6.80	20.63-22.14
Qchl.saw-2D.4	E2	2D	3.62	9.14	‒0.78	Ex_c66554_69	D_contig73920_552	18.50-25.10	41.92-61.81
Qchl.saw-3A	E3	3A	3.50	7.50	‒0.62	BS00023222_51	Excalibur_c91430_125	123.60-129.00	714.59-724.06
	E5	3A	3.12	5.30	‒0.54	BS00023222_51	Excalibur_c91430_125	123.50-129.30	714.59-724.06
Qchl.saw-3B	E4	3B	4.15	9.30	0.91	Kukri_rep_c113136_131	IACX6214	21.40-35.00	12.03-18.56
Qchl.saw-4A	E2	4A	2.79	6.51	‒0.78	Excalibur_rep_c114266_234	GENE-2399_666	15.00-15.70	612.06-618.46
Qchl.saw-4B	E5	4B	3.37	5.11	‒0.53	Kukri_c12661_326	CAP11_c1143_339	43.10-49.50	28.73-50.40
Qchl.saw-4D.1	E4	4D	2.96	5.77	0.69	Excalibur_c32965_551	Excalibur_c55561_127	22.50-24.50	43.40-119.91
	BLUP	4D	2.73	4.77	0.36	Excalibur_c32965_551	Excalibur_c55561_127	13.60-24.00	43.40-119.91
Qchl.saw-4D.2	E4	4D	4.25	7.98	0.82	D_contig16493_653	CAP7_c7991_177	25.80-31.60	386.44-451.30
	E5	4D	3.65	6.33	0.58	D_contig16493_653	CAP7_c7991_177	25.30-31.20	386.44-451.30
	BLUP	4D	3.37	5.92	0.40	D_contig16493_653	CAP7_c7991_177	25.00-31.80	386.44-451.30
Qchl.saw-6A	E2	6A	3.93	8.14	0.74	Excalibur_c1708_1975	Tdurum_contig95236_98	59.90-62.70	29.56-32.88
	E4	6A	5.12	9.74	0.90	Excalibur_c1708_1975	Tdurum_contig95236_98	60.70-63.30	29.56-32.88
	BLUP	6A	2.60	4.10	0.34	Excalibur_c1708_1975	Tdurum_contig95236_98	60.70-64.20	29.56-32.88
Qchl.saw-6B.1	E3	6B	2.91	6.08	‒0.56	RAC875_c50436_101	wsnp_RFL_Contig1075_104560	79.60-100.10	686.19-694.40
Qchl.saw-6B.2	E1	6B	2.86	5.27	‒0.74	BS00062776_51	tplb0046e21_618	116.20-116.90	706.34-708.03

表2

图2

表3

图3

表4

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环境 Environment	亲本 Parents		DH群体 DH Population
环境 Environment	晋春7号 Jinchun 7	金麦919 Jinmai 919	平均值 Mean	标准差 SD	最大值 Max.	最小值 Min.	峰度 Kurtosis	偏度 Skewness	遗传力H² Heritability
E1	57.07^*	54.96	54.78	3.19	62.77	41.33	1.572	-0.336	0.58
E2	46.17	44.43	48.42	2.55	53.60	39.80	1.397	-0.856
E3	58.59	57.79	56.90	2.23	62.38	50.86	-0.185	-0.091
E4	55.92^*	52.76	53.76	2.85	61.58	48.10	-0.392	-0.271
E5	53.59^*	50.87	50.56	2.31	56.14	44.92	-0.056	-0.027
BLUP	54.57	52.38	52.88	1.62	57.23	47.64	0.258	-0.376

染色体 Chromosome	QTL	左标记 Left marker	右标记 Right marker	物理位置 Physical position (Mb)	表型贡献率 PVE (%)	参考文献 Reference
2D	qHChlb2D	Xcfd53	—	23.03	14.31	[16]
2D	QXcfd0116-2D	Xcfd0116	wPt-2644	205.62	—	[41]
2D	Qchc.iiwbr-2D	Xbarc228	—	570.40	16.55	[42]
2D	Qlchc7a.iiwbr-2D	Xwmc18	—	130.83	31.36	[42]
2D	QChl-2D	Xwmc0018	wPt-0298	130.83	6.46	[40]
2D	qCHN-2Da	Xbcd611	Xcdo1379	—	4.73	[43]
2D	qCHN-2Db	Xfba074	Xfba062	—	5.49	[43]
2D	QXcfd53-2D	Xcfd53	—	23.03	—	[38]
2D	QXbarc168.1-2D	Xbarc168.1	—	44.74	—	[38]
2D	QXgwm539-2D	Xgwm539	—	513.10	—	[38]
4D	qChlH-4D	Xgwm192	Xwmc331	412.72	19.90	[39]
6A	QXgwm617	Xgwm617	—	—	11.00	[22]
6A	QXgwm334	Xgwm334	—	9.25	23.00	[22]
6A	QChl-6A.1	Xgwm570	Xpsp3071	447.12	12.80	[18]
6A	Qspad.acs-6A.1	Xwmc201	Xwmc684	461.84	—	[24]
6A	Qspad.acs-6A.2	Xbarc171	Xgwm427	354.592	—	[24]
6A	Qspad.acs-6A.3	Xgwm169	Xwmc580	595.378	—	[24]
6A	Qspad.acs-6A.4	Xbarc113	Xwmc621	495.11	—	[24]
6A	QChl-6A	aca/caa-5	wPt-7599	32.67	7.44	[40]
6A	qCHO-6A	Xmwg2053	Xmwg573	—	7.72	[43]
6A	QChl.ksu-6A	Xgwm427	Xgwm169	595.38	16.10	[2]
6A	QChlL.igdb‐6A	Xgwm570	—	579.13	12.40	[44]
6A	QHtscc.ksu-6A	Xbarc113	CTGTGC93	—	17.57	[5]

QTL名称	染色体	标记区间	基因	水稻同源基因	物理位置	基因注释
QTL name	Chr.	Marker interval	Gene	Rice homologous gene	Position (bp)	Gene annotation
Qchl.saw-2D.1	2D	BS00012148_51-Excalibur_c18324_390	TraesCS2D02G007500	—	4038523	镁离子结合 Magnesium ion binding
Qchl.saw-2D.1	2D	BS00012148_51-Excalibur_c18324_390	TraesCS2D02G008700	—	4685051	镁离子结合 Magnesium ion binding
Qchl.saw-2D.1	2D	BS00012148_51-Excalibur_c18324_390	TraesCS2D02G010200	Os04g0118700	5230187	叶绿体RNA加工 Chloroplast RNA processing
Qchl.saw-2D.1	2D	BS00012148_51-Excalibur_c18324_390	TraesCS2D02G014800	—	7148652	叶绿体组织 Chloroplast organization
Qchl.saw-4D.2	4D	D_contig16493_653-CAP7_c7991_177	TraesCS4D02G273000	—	442898720	光系统I中的光合电子传递 Photosynthetic electron transport in photosystem I
Qchl.saw-4D.2	4D	D_contig16493_653-CAP7_c7991_177	TraesCS4D02G248100	Os03g0212700	416933144	金属离子结合 Metal ion binding
Qchl.saw-4D.2	4D	D_contig16493_653-CAP7_c7991_177	TraesCS4D02G270900	—	441653004	电子传递活性 Electron transfer activity
Qchl.saw-4D.2	4D	D_contig16493_653-CAP7_c7991_177	TraesCS4D02G263300	Os03g0196800	434273089	叶绿体基质 Chloroplast stroma
Qchl.saw-4D.2	4D	D_contig16493_653-CAP7_c7991_177	TraesCS4D02G277400	Os03g0163300 (OsGR1)	449725264	电子传递活性 Electron transfer activity
Qchl.saw-4D.2	4D	D_contig16493_653-CAP7_c7991_177	TraesCS4D02G267100	Os03g0192400	437810691	电子传递链 Electron transport chain
Qchl.saw-4D.2	4D	D_contig16493_653-CAP7_c7991_177	TraesCS4D02G230000	Os03g0240500	389572209	叶绿体外膜 Chloroplast outer membrane
Qchl.saw-4D.2	4D	D_contig16493_653-CAP7_c7991_177	TraesCS4D02G275300	Os03g0161200	446137350	叶绿体 Chloroplast

小麦旗叶叶绿素含量的QTL定位及验证

QTL mapping and validation of chlorophyll content of flag leaves in wheat (Triticum aestivum L.)

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