中国农业科学 ›› 2019, Vol. 52 ›› Issue (11): 1839-1857.doi: 10.3864/j.issn.0578-1752.2019.11.001
史大坤1,姚天茏1,刘楠楠2,邓敏3,段海洋1,王路林1,万炯1,高炯浩1,谢惠玲1,汤继华1,张雪海1()
收稿日期:
2019-01-21
接受日期:
2019-02-28
出版日期:
2019-06-01
发布日期:
2019-06-11
通讯作者:
张雪海
作者简介:
史大坤,E-mail:912320091@qq.com。|姚天茏,E-mail:yaotianlong0629@qq.com。
基金资助:
SHI DaKun1,YAO TianLong1,LIU NanNan2,DENG Min3,DUAN HaiYang1,WANG LuLin1,WAN Jiong1,GAO JiongHao1,XIE HuiLing1,TANG JiHua1,ZHANG XueHai1()
Received:
2019-01-21
Accepted:
2019-02-28
Online:
2019-06-01
Published:
2019-06-11
Contact:
XueHai ZHANG
摘要:
目的 叶绿素含量与作物产量呈正相关。通过提高叶绿素含量来提高作物产量是作物育种的方向之一。因此,利用全基因组关联分析(genome-wide association study, GWAS)解析玉米叶绿素含量的遗传基础,可为玉米高光效理想株型设计育种提供理论指导。方法 以538份玉米自交系构成的关联群体为研究对象,在5个环境下,通过对其授粉后5 d的棒三叶(穗位叶、穗上叶、穗下叶)叶绿素含量进行测定,并借助覆盖玉米全基因组的558 629个单核苷酸多态性标记(SNPs),利用3种模型(Q、K和Q+K)对叶绿素含量进行全基因组关联分析,随后选择最优模型的GWAS结果并结合eQTL(expression quantitative trait loci)分析对叶绿素含量的自然变异进行解析。结果 5个环境下,棒三叶叶绿素含量均遵从正态分布且叶绿素含量间呈正相关;方差分析表明棒三叶叶绿素含量的环境效应、基因型效应、基因型与环境互作效应均达到了极显著水平;此外,穗上叶、穗位叶和穗下叶叶绿素含量的遗传力分别为0.66、0.66和0.67。比较3种模型发现K模型对假阳性(I型错误)控制最好,在此模型下共检测到29个与棒三叶叶绿素含量显著关联的SNP(P≤3.99×10-6),涉及到18个位点,共有76个候选基因落在这18个位点内,其中85.5%(65/76)的候选基因具有eQTL,11.8%(9/76)的候选基因与对应表型显著相关(P<0.05),说明这9个基因可能是通过表达量变化来调控表型变异。在这76个基因中,60个候选基因有功能注释,功能涉及到能量代谢、物质输送代谢途径和生物合成调节等过程。此外还发现2个可以在不同环境或不同叶片共定位的位点,其中,共定位位点内的基因GRMZM2G074759编码一种与AAE3高度相似的酰基活化酶,该基因通过提高α-酮戊二酸(ALA)和草酰乙酸含量进而影响氨基酸生物合成,提高籽粒赖氨酸含量,改善玉米品质。此外,ALA的合成会促使叶绿素含量升高,进而提高作物产量,推测该基因为最可能的候选基因。结论 K模型对假阳性的控制效果最好,基于K模型,共检测到18个玉米叶绿素含量显著关联位点,发现多个参与叶绿素合成途径相关基因。
史大坤, 姚天茏, 刘楠楠, 邓敏, 段海洋, 王路林, 万炯, 高炯浩, 谢惠玲, 汤继华, 张雪海. 玉米叶绿素含量的全基因组关联分析[J]. 中国农业科学, 2019, 52(11): 1839-1857.
SHI DaKun, YAO TianLong, LIU NanNan, DENG Min, DUAN HaiYang, WANG LuLin, WAN Jiong, GAO JiongHao, XIE HuiLing, TANG JiHua, ZHANG XueHai. Genome-Wide Association Study of Chlorophyll Content in Maize[J]. Scientia Agricultura Sinica, 2019, 52(11): 1839-1857.
表1
关联群体不同环境下玉米棒三叶叶绿素含量的描述统计分析"
环境 Environment | 性状 Traits | 变异范围 Range | 均值 Mean | 标准差 sd. | 偏度 Ske. | 峰度 Kur. |
---|---|---|---|---|---|---|
2012年鹤壁Hebi, 2012 | 穗上叶Above the uppermost ear leaf | 26.25—67.85 | 52.98 | 6.91 | -0.87 | 1.51 |
穗位叶Uppermost ear leaf | 22.25—68.60 | 52.82 | 7.17 | -0.97 | 1.69 | |
穗下叶Below the uppermost ear leaf | 18.25—66.20 | 52.92 | 7.58 | -1.08 | 1.92 | |
2017年三亚Sanya, 2017 | 穗上叶Above the uppermost ear leaf | 29.25—68.45 | 50.97 | 6.39 | -0.46 | 0.90 |
穗位叶Uppermost ear leaf | 25.83—70.45 | 51.08 | 6.37 | -0.43 | 1.16 | |
穗下叶Below the uppermost ear leaf | 25.65—69.60 | 51.03 | 6.67 | -0.48 | 0.92 | |
2018年长沙Changsha, 2018 | 穗上叶Above the uppermost ear leaf | 27.90—82.93 | 53.84 | 6.20 | -0.26 | 3.08 |
穗位叶Uppermost ear leaf | 29.47—82.95 | 53.83 | 6.10 | -0.08 | 2.98 | |
穗下叶Below the uppermost ear leaf | 22.88—95.28 | 53.93 | 7.44 | -0.09 | 4.59 | |
2018年原阳Yuanyang, 2018 | 穗上叶Above the uppermost ear leaf | 30.80—63.35 | 48.05 | 5.74 | -0.48 | 0.97 |
穗位叶Uppermost ear leaf | 30.32—61.99 | 47.88 | 5.80 | -0.39 | 0.84 | |
穗下叶Below the uppermost ear leaf | 29.73—60.64 | 47.97 | 6.13 | -0.33 | 0.52 | |
2018年永城Yongcheng, 2018 | 穗上叶Above the uppermost ear leaf | 40.92—69.59 | 54.67 | 6.61 | 0.00 | 0.00 |
穗位叶Uppermost ear leaf | 38.30—68.48 | 54.86 | 6.74 | -0.20 | 0.37 | |
穗下叶Below the uppermost ear leaf | 37.85—69.15 | 54.99 | 6.96 | -0.03 | -0.04 | |
综合Blup | 穗上叶Above the uppermost ear leaf | 43.46—58.68 | 52.51 | 3.16 | -0.57 | 1.02 |
穗位叶Uppermost ear leaf | 40.45—60.49 | 52.57 | 3.61 | -0.55 | 0.78 | |
穗下叶Below the uppermost ear leaf | 40.20—64.44 | 52.55 | 4.13 | -0.50 | 0.64 |
表2
5个环境玉米棒三叶叶绿素含量的方差分析"
性状 Trait | 变异来源 Source | 离均差平方和 Type sum of squares | 自由度 Df | 均方 Mean square | F值 F-value |
---|---|---|---|---|---|
穗上叶 Above the uppermost ear leaf | 环境E | 62052.294 | 4 | 15513.073 | 1202.047** |
基因型G | 51835.582 | 537 | 96.528 | 7.479** | |
基因型×环境 G×E | 122215.029 | 2148 | 56.897 | 4.408** | |
误差Error | 138863.674 | 10760 | 12.906 | ||
总变异Total | 374966.579 | 13449 | |||
穗位叶 Uppermost ear leaf | 环境E | 74363.802 | 4 | 18590.951 | 1466.621** |
基因型G | 56009.168 | 537 | 104.300 | 8.228** | |
基因型×环境 G×E | 126440.437 | 2148 | 58.864 | 4.643** | |
误差Error | 136394.238 | 10760 | 12.676 | ||
总变异Total | 393207.645 | 13449 | |||
穗下叶 Below the uppermost ear leaf | 环境E | 101559.963 | 4 | 25389.991 | 1822.287** |
基因型G | 59378.695 | 537 | 110.575 | 7.936** | |
基因型×环境 G×E | 135210.636 | 2148 | 62.947 | 4.518** | |
误差Error | 149919.461 | 10760 | 13.933 | ||
总变异Total | 446068.755 | 13449 |
表3
玉米棒三叶叶绿素含量候选基因及功能注释"
位点a Loci | 染色 Chr. | 环境 Enviro. | 性状 Trait | 峰值SNP Peak SNP | 物理位置b Posi. (bp) | P值c P value | 贡献率d R2(%) | 候选基因e Candidate gene | 功能注释 Annotation |
---|---|---|---|---|---|---|---|---|---|
1 | 1 | 2018年长沙 Changsha, 2018 | 穗位叶 Uppermost ear leaf | Chr.1.S_33879839 | 33879839 | 1.05E-06 | 5.89 | GRMZM2G110023 | ATP结合亚基CLPT1叶绿体 ATP-dependent Clp protease ATP-binding subunit CLPT1 chloroplastic |
GRMZM2G110004 | NA | ||||||||
GRMZM2G035552 | ACR8 | ||||||||
GRMZM2G035461 | 自噬蛋白9Autophagy protein 9 | ||||||||
2 | 1 | 2017年三亚 Sanya, 2017 | 穗位叶 Uppermost ear leaf | Chr.1.S_249417295 | 249417295 | 2.65E-06 | 5.43 | GRMZM2G031771 | NA |
GRMZM2G031788 | NA | ||||||||
GRMZM2G058057 | 同源线粒体导入受体亚基 Mitochondrial import receptor subunit TOM5 homolog TOM5 | ||||||||
GRMZM2G058227 | 甘油磷酰二元酸酯磷酸二酯酶Glycerophosphoryl diester phosphodiesterase | ||||||||
GRMZM2G058244 | UDP -葡萄糖6脱氢酶UDP-glucose 6-dehydrogenase | ||||||||
GRMZM2G172512 | 未知Unknown | ||||||||
GRMZM2G172584 | 未知Unknown | ||||||||
2 | 1 | 2017年三亚 Sanya, 2017 | 穗位叶 Uppermost ear leaf | Chr.1.S_249417300 | 249417300 | 2.65E-06 | 5.43 | GRMZM2G031771 | NA |
GRMZM2G031788 | NA | ||||||||
GRMZM2G058057 | TOM5同源线粒体导入受体亚基 Mitochondrial import receptor subunit TOM5 homolog | ||||||||
GRMZM2G058227 | 甘油磷酰二元酸酯磷酸二酯酶Glycerophosphoryl diester phosphodiesterase | ||||||||
GRMZM2G058244 | UDP -葡萄糖6脱氢酶UDP-glucose 6-dehydrogenase | ||||||||
GRMZM2G172512 | 未知Unknown | ||||||||
GRMZM2G172584 | 未知Unknown | ||||||||
2 | 1 | 2017年三亚 Sanya, 2017 | 穗位叶 Uppermost ear leaf | Chr.1.S_249417301 | 249417301 | 2.65E-06 | 5.43 | GRMZM2G031771 | NA |
GRMZM2G031788 | NA | ||||||||
GRMZM2G058057 | TOM5同源线粒体导入受体亚基 Mitochondrial import receptor subunit TOM5 homolog | ||||||||
GRMZM2G058227 | 甘油磷酰二元酸酯磷酸二酯酶Glycerophosphoryl diester phosphodiesterase | ||||||||
GRMZM2G058244 | UDP -葡萄糖6脱氢酶UDP-glucose 6-dehydrogenase | ||||||||
位点a Loci | 染色 Chr. | 环境 Enviro. | 性状 Trait | 峰值SNP Peak SNP | 物理位置b Posi. (bp) | P值c P value | 贡献率d R2(%) | 候选基因e Candidate gene | 功能注释 Annotation |
GRMZM2G172512 | 未知Unknown | ||||||||
GRMZM2G172584 | 未知Unknown | ||||||||
3 | 3 | 2012年鹤壁 Hebi, 2012 | 穗下叶Below the uppermost ear leaf | Chr.3.S_167437401 | 167437401 | 2.88E-07 | 6.52 | GRMZM2G701895 | NA |
GRMZM2G044055 | 甲酸精蛋白1Formin-like protein 1 | ||||||||
GRMZM2G044092 | 葡萄糖苷酶42Beta-glucosidase 42 | ||||||||
GRMZM2G118362 | 含五肽复合物的线粒体蛋白 Pentatricopeptide repeat-containing protein mitochondrial | ||||||||
4 | 3 | 综合Blup | 穗上叶Above the uppermost ear leaf | PZE-103136749 | 192318356 | 6.48E-07 | 5.10 | GRMZM2G038953 | 54 kD蛋白3信号识别粒子Signal recognition particle 54 kD protein 3 |
5 | 5 | 综合Blup | 穗上叶Above the uppermost ear leaf | PZE-105049606 | 41552842 | 1.46E-06 | 4.80 | GRMZM2G007185 | β13-半乳糖基转移酶4 Probable beta-13-galactosyltransferase 4 |
5 | 5 | 综合Blup | 穗上叶Above the uppermost ear leaf | PZE-105049607 | 41552873 | 2.58E-06 | 4.53 | GRMZM2G007185 | β13-半乳糖基转移酶4 Probable beta-13-galactosyltransferase 4 |
6 | 5 | 2012年鹤壁 Hebi, 2012 | 穗位叶 Uppermost ear leaf | Chr.5.S_174139937 | 174139937 | 2.23E-06 | 5.36 | GRMZM5G836353 | 单氧酶1 Monooxygenase 1 |
GRMZM2G339523 | 单氧酶1 Monooxygenase 1 | ||||||||
GRMZM2G087169 | 单氧酶1 Monooxygenase 1 | ||||||||
GRMZM2G087068 | 类Cmc1细胞色素c氧化酶生物发生蛋白 Cytochrome c oxidase biogenesis protein Cmc1-like | ||||||||
GRMZM2G388099 | 未知Unknown | ||||||||
GRMZM2G087040 | 乙烯反应转录因子ERF021Ethylene-responsive transcription factor ERF021 | ||||||||
GRMZM2G086946 | TPX2(靶向蛋白为Xklp2)蛋白家族 TPX2 (targeting protein for Xklp2) protein family | ||||||||
7 | 5 | 综合Blup | 穗下叶Below the uppermost ear leaf | Chr.5.S_216586758 | 216586758 | 2.75E-06 | 5.03 | GRMZM2G014116 | 26S蛋白酶体非ATP酶调节亚基 26S proteasome non-ATPase regulatory subunit 12 homolog B |
GRMZM2G014793 | O-岩藻糖转移酶家族蛋白O-fucosyltransferase family protein | ||||||||
GRMZM2G015090 | 植物细胞内ras -group相关LRR蛋白 Plant intracellular Ras-group-related LRR protein 9 | ||||||||
GRMZM2G145061 | 脯氨酰4羟化酶亚基α2 Prolyl 4-hydroxylase alpha-2 subunit | ||||||||
GRMZM2G145107 | 蔗糖转运2 Sucrose transporter2 | ||||||||
GRMZM2G145128 | 含有重复PPR的线粒体蛋白Pentatricopeptide repeat-containing protein | ||||||||
位点a Loci | 染色 Chr. | 环境 Enviro. | 性状 Trait | 峰值SNP Peak SNP | 物理位置b Posi. (bp) | P值c P value | 贡献率d R2(%) | 候选基因e Candidate gene | 功能注释 Annotation |
GRMZM2G145133 | 未知Unknown | ||||||||
GRMZM2G445944 | 转录因子TCP21Transcription factor TCP21 | ||||||||
GRMZM2G445958 | 复制蛋白a32 kD亚基A Replication protein A 32 kD subunit A | ||||||||
GRMZM2G145473 | 未知Unknown | ||||||||
8 | 7 | 2012年鹤壁 Hebi, 2012 | 穗下叶Below the uppermost ear leaf | Chr.7.S_9204823 | 9204823 | 4.39E-07 | 6.37 | GRMZM2G704349 | Pumilio同源3 Pumilio homolog 3 |
GRMZM2G046776 | Pumilio同源3 Pumilio homolog 3 | ||||||||
GRMZM2G046750 | 脂质转运蛋白Lipid-transfer protein 种子贮藏2S白蛋白超家族蛋白Seed storage 2S albumin superfamily protein | ||||||||
GRMZM2G046529 | 叶绿体中蛋白质脂肪酸输出1 Protein FATTY ACID EXPORT 1 chloroplastic | ||||||||
GRMZM2G004301 | Ⅳ型肌醇多磷酸酶Type Ⅳ inositol polyphosphate | ||||||||
9 | 7 | 2017年三亚 Sanya, 2017 | 穗下叶Below the uppermost ear leaf | Chr.7.S_145681493 | 145681493 | 1.72E-06 | 5.54 | GRMZM2G168913 | RNA聚合酶Ⅱ的中介体亚单位8 Mediator of RNA polymerase Ⅱtranscription subunit 8 |
GRMZM2G168858 | NADPH-细胞色素P450还原酶NADPH--cytochrome P450 reductase 2 | ||||||||
GRMZM2G357688 | 锌指蛋白2 Zinc finger protein 2 | ||||||||
GRMZM2G436511 | BTB/POZ蛋白域 BTB/POZ domain-containing protein | ||||||||
GRMZM2G135651 | 环核苷酸封闭通道Cyclic nucleotide-gated ion channel 14 | ||||||||
9 | 7 | 2017年三亚 Sanya, 2017 | 穗下叶Below the uppermost ear leaf | Chr.7.S_145682189 | 145682189 | 5.85E-07 | 6.07 | GRMZM2G168913 | RNA聚合酶Ⅱ的中介体亚单位8 Mediator of RNA polymeraseⅡ transcription subunit 8 |
GRMZM2G168858 | NADPH-细胞色素P450还原酶2 NADPH--cytochrome P450 reductase 2 | ||||||||
GRMZM2G357688 | 锌指蛋白2Zinc finger protein 2 | ||||||||
GRMZM2G436511 | BTB/POZ蛋白域BTB/POZ domain-containing protein | ||||||||
GRMZM2G135651 | 环核苷酸封闭通道Cyclic nucleotide-gated ion channel 14 | ||||||||
10 | 8 | 2012年鹤壁 Hebi, 2012 | 穗上叶Above the uppermost ear leaf | Chr.8.S_1454124 | 1454124 | 1.32E-06 | 5.55 | GRMZM2G378906 | 外被体蛋白β亚基Coatomer subunit beta |
GRMZM2G110368 | 未知Unknown | ||||||||
11 | 8 | 综合Blup | 穗位叶 Uppermost ear leaf | Chr.8.S_1454124 | 1454124 | 3.95E-06 | 4.70 | GRMZM2G378906 | 外被体蛋白β亚基Coatomer subunit beta |
GRMZM2G110368 | 未知Unknown | ||||||||
位点a Loci | 染色 Chr. | 环境 Enviro. | 性状 Trait | 峰值SNP Peak SNP | 物理位置b Posi. (bp) | P值c P value | 贡献率d R2(%) | 候选基因e Candidate gene | 功能注释 Annotation |
12 | 9 | 综合Blup | 穗上叶Above the uppermost ear leaf | Chr.9.S_9638369 | 9638369 | 1.32E-06 | 4.86 | GRMZM2G159402 | 假定锌指蛋白Putative zinc finger protein |
GRMZM2G033130 | DUF1296结构域蛋白家族 Putative DUF1296 domain containing family protein | ||||||||
12 | 9 | 综合Blup | 穗上叶Above the uppermost ear leaf | Chr.9.S_9639904 | 9639904 | 1.32E-06 | 4.86 | GRMZM2G159402 | 假定锌指蛋白Putative zinc finger protein |
GRMZM2G033130 | DUF1296结构域蛋白家族Putative DUF1296 domain containing family protein | ||||||||
12 | 9 | 综合Blup | 穗上叶Above the uppermost ear leaf | chr9.S_9642472 | 9642472 | 1.32E-06 | 4.86 | GRMZM2G159402 | 假定锌指蛋白Putative zinc finger protein |
GRMZM2G033130 | DUF1296结构域蛋白家族Putative DUF1296 domain containing family protein | ||||||||
12 | 9 | 综合Blup | 穗上叶Above the uppermost ear leaf | Chr.9.S_9643507 | 9643507 | 1.32E-06 | 4.86 | GRMZM2G159402 | 假定锌指蛋白Putative zinc finger protein |
GRMZM2G033130 | DUF1296结构域蛋白家族Putative DUF1296 domain containing family protein | ||||||||
12 | 9 | 综合Blup | 穗上叶Above the uppermost ear leaf | Chr.9.S_9644832 | 9644832 | 1.32E-06 | 4.86 | GRMZM2G159402 | 假定锌指蛋白Putative zinc finger protein |
GRMZM2G033130 | DUF1296结构域蛋白家族Putative DUF1296 domain containing family protein | ||||||||
13 | 9 | 2017年三亚 Sanya, 2017 | 穗位叶 Uppermost ear leaf | Chr.9.S_151327393 | 151327393 | 1.72E-06 | 6.01 | GRMZM2G178787 | 叶绿体蛋白激酶Protein kinase 2B chloroplastic |
GRMZM2G178826 | 脱氢奎尼酸合酶3-dehydroquinate synthase | ||||||||
GRMZM2G479529 | 未知Unknown | ||||||||
GRMZM2G178847 | 未知Unknown | ||||||||
GRMZM2G178859 | 原血红素Ⅸ法氏转移酶Protoheme Ⅸ farnesyltransferase | ||||||||
GRMZM2G178880 | 甘露聚糖合酶7 Probable mannan synthase 7 | ||||||||
GRMZM2G178916 | 含有重复PPR的线粒体蛋白 Pentatricopeptide repeat-containing protein mitochondrial | ||||||||
GRMZM2G479581 | 未知Unknown | ||||||||
GRMZM2G178945 | O-岩藻糖转移酶家族蛋白O-fucosyltransferase family protein | ||||||||
GRMZM2G178960 | 核酮糖磷酸3-表异构酶Ribulose-phosphate 3-epimerase | ||||||||
GRMZM2G057950 | 核糖体蛋白L32含有蛋白质的异构体1%3B核糖体蛋白L32,含有蛋白质的异构体 2Ribosomal protein L32 containing protein isoform 1%3B, Ribosomal protein L32 containing protein isoform 2 | ||||||||
位点a Loci | 染色 Chr. | 环境 Enviro. | 性状 Trait | 峰值SNP Peak SNP | 物理位置b Posi. (bp) | P值c P value | 贡献率d R2(%) | 候选基因e Candidate gene | 功能注释 Annotation |
GRMZM2G058037 | 类HVA22蛋白Ⅰ HVA22-like proteinⅠ | ||||||||
GRMZM2G361256 | ATP结合盒(ABC)蛋白家族3 ABC transporter C family member 3 | ||||||||
14 | 9 | 2017年三亚 Sanya, 2017 | 穗位叶 Uppermost ear leaf | Chr.9.S_153737236 | 153737236 | 7.91E-07 | 6.59 | GRMZM2G382537 | 未知Unknown |
GRMZM2G382534 | 羟化酶5 Hydroxylase5 | ||||||||
GRMZM2G060564 | PLAC8 家族蛋白PLAC8 family protein | ||||||||
GRMZM2G060507 | 假定类结转录因子家族蛋白Putative knotted-like transcription factor family protein | ||||||||
GRMZM2G064005 | Protein FIZZY-RELATED 1 FIZZY-RELATED蛋白 | ||||||||
GRMZM2G063931 | 假定泛素连接酶酶家族,E2泛素连接酶 Putative ubiquitin-conjugating enzyme family%3B, ubiquitin-conjugating enzyme E2 | ||||||||
15 | 10 | 2012年鹤壁 Hebi, 2012 | 穗位叶 Uppermost ear leaf | Chr.10.S_45404195 | 45404195 | 2.81E-06 | 4.76 | NO | - |
16 | 10 | 2012年鹤壁 Hebi, 2012 | 穗位叶 Uppermost ear leaf | Chr.10.S_46780569 | 46780569 | 2.76E-06 | 4.75 | GRMZM2G135800 | NA |
GRMZM2G371316 | 未知Unknown | ||||||||
GRMZM2G371345 | 黄酮醇3-O-葡糖基转移酶Flavonol 3-O-glucosyltransferase | ||||||||
16 | 10 | 2012年鹤壁 Hebi, 2012 | 穗位叶 Uppermost ear leaf | Chr.10.S_46780652 | 46780652 | 2.76E-06 | 4.75 | GRMZM2G135800 | NA |
GRMZM2G371316 | 未知Unknown | ||||||||
GRMZM2G371345 | 黄酮醇3-O-葡糖基转移酶Flavonol 3-O-glucosyltransferase | ||||||||
17 | 10 | 2018年长沙 Changsha, 2018 | 穗位叶 Uppermost ear leaf | Chr.10.S_148643242 | 148643242 | 3.04E-06 | 5.30 | GRMZM2G074787 | DNA错配修复蛋白MSH3 DNA mismatch repair protein MSH3 |
GRMZM2G074773 | 核转录因子Y亚基C-4 Nuclear transcription factor Y subunit C-4 | ||||||||
GRMZM2G074759 | 酰基活化酶3 Acyl activating enzyme3 | ||||||||
GRMZM2G074754 | C2钙/脂结合植物磷脂酰转移酶家族蛋白 C2 calcium/lipid-binding plant phosphoribosyltransferase family protein | ||||||||
GRMZM2G074718 | 表达蛋白Expressed protein | ||||||||
GRMZM2G074107 | DNA结合蛋白DNA binding protein | ||||||||
位点a Loci | 染色 Chr. | 环境 Enviro. | 性状 Trait | 峰值SNP Peak SNP | 物理位置b Posi. (bp) | P值c P value | 贡献率d R2(%) | 候选基因e Candidate gene | 功能注释 Annotation |
17 | 10 | 2018年长沙 Changsha, 2018 | 穗位叶 Uppermost ear leaf | Chr.10.S_148643247 | 148643247 | 3.87E-06 | 5.27 | GRMZM2G074787 | DNA错配修复蛋白MSH3 DNA mismatch repair protein MSH3 |
GRMZM2G074773 | 核转录因子Y亚基C-4 Nuclear transcription factor Y subunit C-4 | ||||||||
GRMZM2G074759 | 酰基活化酶3 Acyl activating enzyme3 | ||||||||
GRMZM2G074754 | C2钙/脂结合植物磷脂酰转移酶家族蛋白 C2 calcium/lipid-binding plant phosphoribosyltransferase family protein | ||||||||
GRMZM2G074718 | 表达蛋白Expressed protein | ||||||||
GRMZM2G074107 | DNA结合蛋白DNA binding protein | ||||||||
17 | 10 | 2018年长沙 Changsha, 2018 | 穗位叶 Uppermost ear leaf | Chr.10.S_148643249 | 148643249 | 1.78E-07 | 6.71 | GRMZM2G074787 | DNA错配修复蛋白MSH3 DNA mismatch repair protein MSH3 |
GRMZM2G074773 | 核转录因子Y亚基C-4 Nuclear transcription factor Y subunit C-4 | ||||||||
GRMZM2G074759 | 酰基活化酶3 Acyl activating enzyme3 | ||||||||
GRMZM2G074754 | C2钙/脂结合植物磷脂酰转移酶家族蛋白 C2 calcium/lipid-binding plant phosphoribosyltransferase family protein | ||||||||
GRMZM2G074718 | 表达蛋白Expressed protein | ||||||||
GRMZM2G074107 | DNA结合蛋白DNA binding protein | ||||||||
18 | 10 | 2018年长沙 Changsha, 2018 | 穗上叶Above the uppermost ear leaf | Chr.10.S_148643249 | 148643249 | 1.58E-06 | 5.80 | GRMZM2G074787 | DNA错配修复蛋白MSH3 DNA mismatch repair protein MSH3 |
GRMZM2G074773 | 核转录因子Y亚基C-4 Nuclear transcription factor Y subunit C-4 | ||||||||
GRMZM2G074759 | 酰基活化酶3 Acyl activating enzyme3 | ||||||||
GRMZM2G074754 | C2钙/脂结合植物磷脂酰转移酶家族蛋白 C2 calcium/lipid-binding plant phosphoribosyltransferase family protein | ||||||||
GRMZM2G074718 | 表达蛋白Expressed protein | ||||||||
GRMZM2G074107 | DNA结合蛋白DNA binding protein |
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