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作物学报 ›› 2023, Vol. 49 ›› Issue (1): 140-152.doi: 10.3724/SP.J.1006.2023.23020

所属专题: 玉米:遗传育种·种质资源·分子遗传学

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

玉米籽粒淀粉含量全基因组关联分析和候选基因预测

王锐璞1(), 董振营1,2(), 高悦欣1, 鲍建喜1, 殷芳冰1, 李金萍2, 龙艳1,2,*(), 万向元1,2,*()   

  1. 1北京科技大学生物与农业研究中心 / 化学与生物工程学院 / 顺德研究生院 / 北京中智生物农业国际研究院, 北京 100083
    2北京首佳利华科技有限公司 / 主要作物生物育种北京市工程实验室 / 生物育种北京市国际科技合作基地, 北京 100192
  • 收稿日期:2022-02-27 接受日期:2022-05-05 出版日期:2023-01-12 网络出版日期:2022-05-24
  • 通讯作者: 龙艳,万向元
  • 作者简介:王锐璞, E-mail: 15903575520@163.com;
    董振营, E-mail: zydong@ustb.edu.cn第一联系人:**同等贡献
  • 基金资助:
    国家重点研发计划项目“农业生物种质资源挖掘与创新利用”重点专项(2021YFD1200700)

Genome-wide association study and candidate gene prediction of kernel starch content in maize

WANG Rui-Pu1(), DONG Zhen-Ying1,2(), GAO Yue-Xin1, BAO Jian-Xi1, YIN Fang-Bing1, LI Jin-Ping2, LONG Yan1,2,*(), WAN Xiang-Yuan1,2,*()   

  1. 1Zhongzhi International Institute of Agricultural Biosciences, Shunde Graduate School, School of Chemistry and Biological Engineering, Research Center of Biology and Agriculture, University of Science and Technology Beijing (USTB), Beijing 100083, China
    2Beijing Engineering Laboratory of Main Crop Bio-Tech Breeding, Beijing International Science and Technology Cooperation Base of Bio-Tech Breeding, Beijing Solidwill Sci-Tech Co. Ltd., Beijing 100192, China
  • Received:2022-02-27 Accepted:2022-05-05 Published:2023-01-12 Published online:2022-05-24
  • Contact: LONG Yan,WAN Xiang-Yuan
  • About author:First author contact:**Contributed equally to this work
  • Supported by:
    National Key Research and Development Program of China(2021YFD1200700)

摘要:

玉米是重要的粮食作物, 其籽粒重量的70%来自于淀粉。淀粉不仅是人类及其他动物的主要能量来源, 同时也是化工等行业的重要原料。本研究利用711份玉米自交系作为关联群体, 对2个环境下玉米籽粒湿基淀粉含量和干基淀粉含量进行统计分析, 结合覆盖玉米全基因组的2799个单核苷酸多态性(single nucleotide polymorphism, SNP)标记, 通过FarmCPU模型对玉米籽粒淀粉含量性状开展全基因组关联分析(genome-wide association study, GWAS), 共关联到67个显著SNP位点, 其中23个高可信度显著SNP位点可在多个环境重复检测到。3个高可信度SNP标记位点为本研究首次发现与玉米籽粒淀粉含量相关, 其余20个SNP标记位于前人已定位QTL (quantitative trait locus)置信区间或/和已报道与籽粒淀粉含量显著相关SNP标记1 Mb之内。进一步通过基因功能注释、基因本体论(gene ontology, GO)分析及籽粒胚乳基因表达分析, 在23个高可信度显著SNP位点上下游各200 kb候选区间共挖掘45个重要候选基因, 涉及淀粉生物合成与代谢、碳水化合物代谢、糖代谢、激素代谢等途径, 同时检测到2个已报道调控玉米籽粒淀粉含量的基因Ae1Pin1。通过等位变异效应分析鉴定出9个主效SNP位点及其优异等位变异。本研究为深入解析玉米籽粒淀粉含量遗传机制提供了新的信息, 为加速培育高产、优质玉米新品种提供重要基因资源。

关键词: 玉米, 籽粒淀粉含量, 全基因组关联分析, 候选基因

Abstract:

Maize is an important food crop worldwide, and about 70% of its grain weight comes from starch. Starch is not only the main energy resource for human and other animals but also an important raw material for chemical industries. In this study, an association panel including 711 maize inbred lines was used for the examination of both wet-base starch content and dry-base starch content of maize kernel from two environments combined with 2799 single nucleotide polymorphism (SNP) markers spaning the whole genome of maize, genome-wide association study (GWAS) was carried out using FarmCPU model. 67 significant SNPs were identified, of which 23 highly reliable significant SNPs (HRS-SNPs) could be repeatedly associated in different environments. Three HRS-SNPs were reported for the first time by our study, and the remaining 20 HRS-SNPs were either located within the known quantitative trait loci (QTLs) or within 1 Mb of known SNPs associated with mazie kernel starch content. Through gene function annotation, gene ontology (GO) analysis and gene expression analysis, a total of 45 important candidate genes, involving starch biosynthesis, carbohydrate metabolism, sugar metabolism, hormone metabolism, and other pathways were identified within 200 kb regions around the HRS-SNPs. And two genes Ae1 and Pin1 which had been reported to regulate the maize kernel starch content were also detected. Furthermore, elite alles of the nine major SNPs was identified by allelic variation effect analysis. Our study provides new genetic information for further disecting the genetic mechanism of maize kernel starch content and provides important gene resources for accelerating the breeding of new maize varieties with high yield and quality.

Key words: maize, kernel starch content, genome-wide association study, candidate gene

表1

玉米籽粒淀粉含量数据统计分析"

性状及环境
Trait and environment
变异范围
Range (%)
平均值
Average (%)
标准差
SD
变异系数
CV (%)
偏度
Skewness
峰度
Kurtosis
WSYC2019 61.46-71.16 66.95 1.62 2.42 -0.40 -0.03
WSPG2020 59.66-68.94 64.55 1.57 2.43 -0.17 -0.29
DSYC2019 68.14-75.97 72.59 1.21 1.67 -0.49 0.64
DSPG2020 66.40-75.22 71.19 1.27 1.78 -0.25 0.14

附图1

玉米籽粒湿基(A)和干基(B)淀粉含量频率分布 WSYC2019: 2019年崖城玉米籽粒湿基淀粉含量; WSPG2020: 2020年平谷玉米籽粒湿基淀粉含量; DSYC2019: 2019年崖城玉米籽粒干基淀粉含量; DSPG2020: 2020年平谷玉米籽粒干基淀粉含量。"

表2

玉米籽粒淀粉含量数据相关性分析"

性状及环境
Trait and environment
相关系数Correlation coefficient
WSYC2019 WSPG2020 DSYC2019 DSPG2020
WSYC2019 1
WSPG2020 0.627** 1
DSYC2019 0.804** 0.526** 1
DSPG2020 0.563** 0.844** 0.610** 1

附图2

玉米自交系系统进化树"

图1

玉米籽粒淀粉含量全基因组关联分析Manhattan图(A)和QQ图(B) WSBLUP和DSBLUP分别表示玉米籽粒湿基和干基淀粉含量最佳线性无偏预测值; Chr表示染色体。其他处理缩写同表1。"

附表1

本研究所鉴定67个玉米籽粒淀粉含量关联分析显著SNP信息"

关联位点
Associated SNP
染色体
Chr.
位置
Position (bp)
P
P
-value
表型变异率
PVE (%)
性状及环境
Trait and environment
SYN20034 1 13,864,423 4.83E-05 0.94 C*
PZE-101026314 1 16,041,430 1.16E-04 1.22 A
PZE-101058274 1 42,284,401 8.48E-04 2.56 D
PZE-101060842 1 44,730,532 4.27E-05 2.89 E, F
PZE-101099263 1 93,135,341 2.46E-05 0.13 B
PZE-101105385 1 109,861,210 3.43E-04 0.76 D
PZE-101107894 1 115,245,510 4.01E-07 2.86 A, C, D, F
PZE-101117618 1 144,859,179 9.63E-04 0.99 E
PZE-101182888 1 231,208,427 9.34E-05 5.05 B, C
PZE-101188800 1 237,883,989 5.74E-04 0.92 D
PZE-101224367 1 279,939,641 3.43E-04 1.09 A
PZE-101233856 1 286,821,902 2.85E-04 3.93 B, C
PZE-102000560 2 837,383 5.60E-04 0.11 B, C
SYN4735 2 9,618,618 1.02E-05 5.08 B, E
SYN19565 2 32,086,278 1.84E-04 3.23 A
PZE-102055831 2 35,216,627 4.31E-04 1.15 A
PZA02450.1 2 49,124,335 5.70E-04 2.50 F
PZE-102082146 2 69,751,058 8.97E-04 4.28 F
SYN33606 2 169,623,783 4.00E-06 0.21 F
PZE-102123716 2 176,643,239 9.36E-05 2.66 D, F
PZE-102131295 2 187,143,781 3.36E-04 1.68 B
PZA03527.1 3 5,148,837 7.57E-04 7.09 B, C
PZE-103024939 3 16,957,568 1.76E-04 1.30 A
PZE-103065358 3 105,359,736 9.15E-07 1.97 A
PZE-103112971 3 175,092,704 6.95E-05 0.19 F
PZE-103165581 3 218,640,885 2.34E-05 5.86 E, F
PZE-104044786 4 69,044,273 1.14E-04 0.47 A
PZE-104080388 4 158,252,821 3.53E-05 0.01 A
PZE-104092771 4 171,771,502 5.79E-06 3.12 E
PZE-104116111 4 196,212,782 2.20E-04 2.78 E
SYN37141 4 227,482,564 3.12E-04 1.17 E
PZE-105024985 5 12,912,185 4.13E-04 2.10 B, F
SYN27691 5 23,539,045 2.36E-05 0.13 F
PZE-105070840 5 77,119,846 4.46E-04 0.60 A
PZE-105111323 5 172,217,865 1.39E-04 2.36 B, E
PZE-105132778 5 194,317,312 1.53E-04 0.96 C, F
PZE-105158980 5 212,371,623 6.15E-07 0.11 C
SYN35254 5 218,593,404 1.48E-04 2.71 B
PZE-106041751 6 93,712,294 3.59E-06 6.34 B, D, F
PZE-106041753 6 93,713,273 5.79E-06 6.56 C, E
PZE-106049962 6 102,943,268 1.07E-04 3.99 A
PZE-106049961 6 102,943,378 1.70E-04 3.60 C
SYN4597 6 107,195,543 1.11E-04 3.71 F
PZE-106088503 6 150,161,584 5.29E-04 1.77 E
PZE-106104150 6 159,598,074 1.94E-04 0.25 B
SYN4194 6 162,814,863 2.76E-05 2.98 B, C
SYN10687 6 169,009,504 4.06E-05 0.56 E
SYN32203 7 8,799,859 2.50E-05 0.02 D
SYN34669 7 9,820,060 5.57E-05 5.43 D, E
PZE-107044349 7 91,690,556 6.53E-04 0.08 B
SYN17951 7 143,619,654 3.39E-06 5.55 D, E, F
PZE-107088998 7 148,971,333 3.54E-05 0.10 E, F
SYN34204 7 156,490,174 8.51E-04 0.96 B
PZE-108005623 8 5,939,105 8.88E-05 3.99 A, D
PZE-108038271 8 63,280,888 5.15E-04 0.31 C
PZE-108053763 8 97,541,328 1.09E-07 0.03 D, F
SYN21795 8 117,579,482 7.45E-05 5.59 B, C
PZE-108064353 8 117,743,973 1.29E-04 2.07 F
ZM012274-0351 8 120,522,562 8.98E-05 2.03 E
SYN14914 8 132,971,220 6.87E-05 0.48 D
PZE-108078728 8 138,559,450 9.03E-04 2.54 B
PZE-109003046 9 3,291,982 5.83E-04 1.73 B, C
PZA03595.2 9 96,808,133 7.94E-05 0.70 F
PZE-109054725 9 98,196,277 7.53E-04 0.97 B
PZE-109069969 9 117,528,657 1.98E-04 1.15 A
SYN12403 10 2,241,835 1.30E-06 2.31 F
PZE-110061626 10 117,865,752 9.07E-07 0.67 A, C

表3

玉米籽粒淀粉含量显著相关SNP和候选基因功能注释"

关联位点
Associated SNP
染色体
Chr.
位置
Position (bp)
P
P
-value
表型变异率
PVE (%)
性状及环境
Trait and environment
候选基因
Candidate gene ID
基因名
Gene name
功能注释
Gene annotation
GO术语
GO term ID
PZE-101060842 1 44,730,532 4.27E-05 2.89 E, F* Zm00001d028735 Imd1 Isopropylmalate dehydrogenase1
Zm00001d028737 Glucuronoxylan 4-O-methyltransferase 2 GO:0016051
GO:0034637
GO:0033692
PZE-101107894 1 115,245,510 4.01E-07 2.86 A, C, D, F Zm00001d030236 Gibberellin-regulated protein 12
PZE-101182888 1 231,208,427 9.34E-05 5.05 B, C Zm00001d032577 CESA4 CESA4 (CELLULOSE SYNTHASE A4) GO:0016051
GO:0034637
GO:0033692
Zm00001d032578 Dof13 C2C2-Dof-transcription factor 13
PZE-101233856 1 286,821,902 2.85E-04 3.93 B, C Zm00001d034212 Hb8 Homeobox-transcription factor 8
PZE-102000560 2 837,383 5.60E-04 0.11 B, C Zm00001d001785 C2 calcium/lipid-binding plant phosphoribosyltransferase family protein
Zm00001d001791 Glucan endo-1,3-beta-D-glucosidase
SYN4735 2 9,618,618 1.02E-05 5.08 B, E Zm00001d002285 NAC22 NAC-transcription factor 22
Zm00001d002288 bHLH148 bHLH-transcription factor 148
Zm00001d002292 Gpm461b NAD(P)-binding Rossmann-fold superfamily protein GO:0016051
GO:0034637
GO:0033692
PZE-102123716 2 176,643,239 9.36E-05 2.66 D, F Zm00001d005502 Protein SMG7
Zm00001d005506
PZA03527.1 3 5,148,837 7.57E-04 7.09 B, C Zm00001d039469 Traf21 TNF receptor-associated factor 21
Zm00001d039475 Mybr22 MYB-related-transcription factor 22
PZE-103165581 3 218,640,885 2.34E-05 5.86 E, F Zm00001d044081 Hb46 Homeobox-transcription factor 46
Zm00001d044083 Pin10 PIN-formed protein10
Zm00001d044086 Nucleic acid-binding OB-fold-like protein
PZE-105024985 5 12,912,185 4.13E-04 2.10 B, F Zm00001d013500 C2H2-type domain-containing protein
Zm00001d013507 Heat shock 70 kDa protein 6 chloroplastic
PZE-105111323 5 172,217,865 1.39E-04 2.36 B, E Zm00001d016674 HSF6 HSF-transcription factor 6
Zm00001d016675 CTR1 Constitutive triple response1
Zm00001d016684 Ae1 Amylose extender1 GO:0016051
GO:0034637
GO:0033692
PZE-105132778 5 194,317,312 1.53E-04 0.96 C, F Zm00001d017391 Axial regulator YABBY 1
PZE-106041751 6 93,712,294 3.59E-06 6.34 B, D, F Zm00001d036583 BEACH domain-containing protein C2
PZE-106041753 6 93,713,273 5.79E-06 6.56 C, E Zm00001d036583 BEACH domain-containing protein C2
SYN4194 6 162,814,863 2.76E-05 2.98 B, C Zm00001d038693 Dof10 C2C2-Dof-transcription factor 10
Zm00001d038699 O-methyltransferase ZRP4
SYN34669 7 9,820,060 5.57E-05 5.43 D, E Zm00001d018936 Ribose-5-phosphate isomerase GO:0016051
Zm00001d018937 MMS19 nucleotide excision repair protein
Zm00001d018941 HSF4 HSF-transcription factor 4
SYN17951 7 143,619,654 3.39E-06 5.55 D, E, F Zm00001d021129 Alpha-(14)-fucosyltransferase GO:0016051
GO:0034637
GO:0033692
Zm00001d021130
Zm00001d021135 Pebp24 Phosphatidylethanolamine-binding protein24
Zm00001d021136 PTI3 Like tyrosine-protein kinase 3
PZE-107088998 7 148,971,333 3.54E-05 0.10 E, F Zm00001d021325 L-Aspartase-like family protein
PZE-108005623 8 5,939,105 8.88E-05 3.99 A, D Zm00001d008338 Trps14 Trehalose-6-phosphate synthase14 GO:0016051
GO:0034637
Zm00001d008345 Importin subunit alpha
PZE-108053763 8 97,541,328 1.09E-07 0.03 D, F Zm00001d010056 Delta-1-pyrroline-5-carboxylate synthase
Zm00001d010059 KH domain-containing protein
SYN21795 8 117,579,482 7.45E-05 5.59 B, C Zm00001d010487 UDP-glycosyltransferase 88A1
Zm00001d010490 CW-type Zinc Finger
PZE-109003046 9 3,291,982 5.83E-04 1.73 B, C Zm00001d044812 Pin1 PIN-formed protein1
Zm00001d044813 Gpm441 Putative fructokinase-6 chloroplastic GO:0016051
GO:0034637
GO:0033692
PZE-110061626 10 117,865,752 9.07E-07 0.67 A, C Zm00001d025409 Ereb21 Putative AP2/EREBP transcription factor superfamily protein
Zm00001d025412 Nicotinate phosphoribosyltransferase

附图3

连锁不平衡衰减分析 横坐标表示同一染色体上单核苷酸位点(SNP)之间的物理距离, 纵坐标表示连锁不平衡参数r2值。"

图2

玉米籽粒淀粉含量主效SNP位点等位变异效应分析 *和**分别表示P < 0.05和P < 0.01水平显著。n为样本量。其他处理缩写同表1。"

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