玉米雄穗性状遗传结构与形成分子机制<sup>*</sup>

doi:10.13523/j.cb.2111003

中国生物工程杂志

2021, Vol. 41

Issue (12): 88-102 DOI: 10.13523/j.cb.2111003

玉米生物育种基础研究与关键技术专辑

玉米雄穗性状遗传结构与形成分子机制^*

王彦博^1,²,魏佳^1,²,龙艳^1,^2,³,董振营^1,^2,^**(

),万向元^1,^2,^3,^**(

)

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 100083
2 北京中智生物农业国际研究院北京 100192
3 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

Research Advances on Genetic Structure and Molecular Mechanism Underlying the Formation of Tassel Traits in Maize

WANG Yan-bo^1,²,WEI Jia^1,²,LONG Yan^1,^2,³,DONG Zhen-ying^1,^2,^**(

),WAN Xiang-yuan^1,^2,^3,^**(

)

1 Research Center of Biology and Agriculture, Shunde Graduate School, School of Chemistry and Biological Engineering, University of Science and Technology Beijing, Beijing 100083, China
2 Zhongzhi International Institute of Agricultural Biosciences, Beijing 100192, China
3 Beijing 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

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摘要：

玉米为雌雄同株异花植物,其雄穗着生于植株顶部,雌穗腋生。雄穗一方面需产生足量花粉以保证雌穗授粉结实,另一方面由于对下部叶片的遮蔽作用和自身营养需求,其生长发育会同时影响叶片光合作用效率和能量分配,因此优化雄穗结构是提高玉米产量的重要措施之一。玉米雄穗性状包括雄穗分枝数、雄穗分枝长度、雄穗主轴长度、雄穗分枝总长度、雄穗分枝角度等,均为多基因控制的数量性状。自20世纪90年代,研究者开始利用数量性状位点(quantitative trait locus,QTL)定位方法解析玉米雄穗性状遗传结构;随着玉米自交系B73等参考基因组释放,以及DNA微阵列、基因组重测序等高通量基因分型技术的日益成熟,全基因组关联分析(genome-wide association study, GWAS)成为数量性状遗传研究的主流方法,目前已鉴定出大量玉米雄穗性状遗传位点。通过总结雄穗性状遗传定位研究结果,构建一致性图谱并挖掘定位热点区间,有助于进一步了解雄穗性状遗传结构特征及指导雄穗性状候选基因克隆。此外,通过对调控雄穗发育的已知基因进行功能分类,可为解析玉米雄穗发育的遗传网络和调控通路提供理论支撑。

关键词： 雄穗; 遗传结构; 数量性状位点; QTL定位; 全基因组关联分析

Abstract:

Maize is a monoecious plant with top terminal tassels and lateral ears. The tassels need to produce sufficient pollens to fertilize ears; however, due to the shading effect on the lower leaves and their own nutritional requirements, the growth and development of tassels have a negative effect on yield through affecting leaf photosynthesis and energy distribution of the whole plant. Optimizing the tassel architecture is thus urgent for maize yield improvement. Maize tassel traits include the number of tassel branches, the length of tassel branches, the length of the principal axis of the tassel, the total length of tassel branches, and the angle of tassel branching, which are all complex quantitative traits controlled by different genetic basis. Since the 1990s, researchers have begun to analyze the genetic structure of maize tassel traits by quantitative trait locus(QTL) mapping. With the release of the reference genome of the maize inbred line B73, and the improvement of high-throughput genotyping technologies such as DNA microarrays and genome resequencing, genome-wide association study(GWAS) has been widely applied in recent years, and a large number of loci associated with the maize tassel traits have been identified. Here, the genetic loci of maize tassel traits identified from different research periods were retrieved, and a consistent physical map was built. Furthermore, the genetic mapping hotspots were isolated which will be useful for the further understanding of the genetic structure of maize tassel traits and the guidance of the cloning of tassel-trait-related genes. At the same time, the cloned genes controlling tassel traits and the corresponding functional mechanism were summarized which will be helpful for further deciphering the genetic network and regulatory pathways of tassel development in maize.

Key words: Tassel Genetic structure Quantitative trait loci QTL mapping Genome-wide association study(GWAS)

收稿日期: 2021-10-31 出版日期: 2022-01-13

ZTFLH:

Q819

基金资助: * 中央高校基本科研业务费专项资金(06500060);国家“万人计划”科技创新领军人才特殊支持经费(201608)

通讯作者: 董振营,万向元 E-mail: zydong@ustb.edu.cn;wanxiangyuan@ustb.edu.cn

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引用本文:

王彦博,魏佳,龙艳,董振营,万向元. 玉米雄穗性状遗传结构与形成分子机制^*[J]. 中国生物工程杂志, 2021, 41(12): 88-102.

WANG Yan-bo,WEI Jia,LONG Yan,DONG Zhen-ying,WAN Xiang-yuan. Research Advances on Genetic Structure and Molecular Mechanism Underlying the Formation of Tassel Traits in Maize. China Biotechnology, 2021, 41(12): 88-102.

链接本文:

https://manu60.magtech.com.cn/biotech/CN/10.13523/j.cb.2111003 或 https://manu60.magtech.com.cn/biotech/CN/Y2021/V41/I12/88

图1 玉米花序发育过程示意图

表1 玉米雄穗性状QTLs、QTNs总结

图2 雄穗性状相关QTLs、QTNs在玉米基因组上的分布

图3 雄穗相关性状QTL、QTN热点区间在玉米基因组的分布

生物学功能	基因名称	基因号	编码蛋白	染色体位置/bp	参考文献
干细胞增殖和分化	CT2	Zm00001d027886	异三聚体GTP结合蛋白	Chr1: 16721214~16732176	[46]
	ZmWUS1	Zm00001d001948	同源结构域转录因子	Chr2: 3415296~3419504	[47]
	FEA3	Zm00001d040130	亮氨酸重复类受体蛋白	Chr3: 28709631~28715222	[48]
	FEA2	Zm00001d051012	亮氨酸重复类受体蛋白	Chr4: 136764371~136769212	[49-50]
	TD1	Zm00001d014793	亮氨酸重复类受体蛋白激酶	Chr5: 63455339~63461620	[51]
	FEA4	Zm00001d037317	bZIP转录因子	Chr6: 120722612~120728273	[52]
	ZmWUS2	Zm00001d026537	同源结构域转录因子	Chr10: 147854036~147858373	[47]
分生组织起始	BIF2	Zm00001d031068	丝苏氨酸激酶	Chr1: 175806351~175810932	[52-54]
	BA2	Zm00001d003897	bHLH 转录因子	Chr2: 65741213~65756966	[55]
	RA2	Zm00001d039694	LOB转录因子	Chr3: 12156780~12160565	[56]
	BA1	Zm00001d042989	bHLH 转录因子	Chr3: 186013129~186016764	[57]
	SPI1	Zm00001d044069	黄素单氧酶	Chr3: 218285795~218290961	[58]
	BIF4	Zm00001d037691	Aux/IAA蛋白	Chr6: 134087331~134094170	[59]
	RA1	Zm00001d020430	Cys2-His2 锌脂转录因子	Chr7: 113570910~113574437	[60]
	RA3	Zm00001d022193	海藻糖-6磷酸磷酸酶	Chr7: 172483459~172490694	[61]
	VT2	Zm00001d008700	色氨酸转氨酶	Chr8: 17393223~17400365	[62]
	BIF1	Zm00001d008749	Aux/IAA蛋白	Chr8: 18950258~18955333	[63]
侧生分生组织起始	UB2	Zm00001d031451	SBP-box转录因子	Chr1: 190381366~190388089	[64]
	CG1	GRMZM2G022489	miR156	Chr3: 6926778~6929901	[65]
	LG2	Zm00001d042777	bZIP蛋白	Chr3: 179386227~179397947	[66]
	UB3	Zm00001d052890	SBP-box转录因子	Chr4: 203609847~203617018	[67]
	BD1	Zm00001d022488	bHLH 转录因子	Chr7: 178604458~178608405	[68]
分生组织确定性	DCL1	Zm00001d027412	miRNA生物合成酶	Chr1: 4722956~4738332	[69]
分生组织确定性	TS4	GRMZM5G803935	SBP-box转录因子	Chr3: 144916511~144920220	[70]
性别决定	TS2	Zm00001d028806	氧化还原酶	Chr1: 46953827~46958171	[71]
	TS1	Zm00001d003533	脂氧合酶蛋白	Chr2: 47103687~47110872	[72]
	SILKY3	Zm00001d004130	核酸蛋白	Chr2: 85360621~85369941	[73]
	NA1	Zm00001d042843	类固醇还原酶	Chr3: 181819965~181824489	[74]
营养阶段向生殖阶段转变	ZFL2	Zm00001d002449	FLORICAULA/LEAFY同源转录因子	Chr2: 12912591~12918568	[75]
营养阶段向生殖阶段转变	ZFL1	Zm00001d026231	FLORICAULA/LEAFY同源转录因子	Chr10: 141560362~141566267	[75]
花期转变	ZMM4	Zm00001d034045	MADS转录因子	Chr1: 282134674~282167480	[76]
分生组织活性	REL2	Zm00001d024523	转录共抑制因子	Chr10: 75992328~76004412	[77]
分生组织起始和小穗分生组织命运	IDS1	Zm00001d034629	AP2转录因子	Chr1: 298421359~298428550	[78]
分生组织起始和小穗分生组织命运	SID1	Zm00001d019230	AP2转录因子	Chr7: 23052961~23067089	[78]
苞片生长发育	TSH1	Zm00001d039113	GATA 锌脂蛋白	Chr6: 170246513~170250985	[79]
苞片生长发育	TSH4	Zm00001d020941	SBP-box转录因子	Chr7: 137272100~137278639	[80]
分生组织活性	RTE	Zm00001d030656	硼转运蛋白	Chr1: 151219729~151227321	[81]
生物学功能	基因名称	基因号	编码蛋白	染色体位置/bp	参考文献
硼转运蛋白	TLS1	Zm00001d032461	硼通道蛋白	Chr1: 227447816~227454579	[82]
雄穗分枝角度	BAD1	Zm00001d005737	TCP转录因子	Chr2: 185420859~185424689	[83]
雄穗分枝数量	QDtbn1	Zm00001d053358	F-box蛋白	Chr4: 227482692~227483909	[84]
细胞分裂素合成相关	KN1	Zm00001d033859	KNOX转录因子	Chr1: 276071835~276082742	[85]

表2 玉米雄花序发育相关基因总结

染色体	QTLs共定位区间/Mb	QTNs共定位区间/Mb	重叠区间/Mb	花序发育相关基因
1	15.446~25.306¹ 33.147~35.164 43.914~47.670² 69.750~78.359 159.020~159.029 196.666~197.169 206.926~208.963 285.747~287.742 290.528~292.719 295.779~295.781 297.925~297.927	10.058~10.754 193.872~194.362 249.337~250.115		CT2¹ TS2²
2	3.380~4.863^3,4 13.851~19.120 26.728~28.948 30.533~30.966 32.090~32.227 189.209~189.291 224.131~224.195	2.707~3.563³ 4.073~5.949⁴ 11.325~14.995 23.168~25.037 48.245~49.011 185.577~186.715 212.155~216.148 241.047~242.030	3.380~4.073³ 13.851~14.995	ZmWUS1³、LG1⁴
3	147.771~147.966 170.148~192.544^5,6,7	19.166~20.083 53.391~53.964 152.498~153.470 174.370~175.283 179.523~184.667⁶ 186.752~187.900 188.380~189.319 214.197~214.926 215.696~217.604 218.524~220.242	174.370~175.283 179.523~184.667⁶ 186.752~187.900 188.380~189.319	LG2⁵ NA1⁶ BA1⁷
4	36.909~38.554 144.880~160.501 175.576~175.640 181.244~181.286 216.731~224.943	155.912~156.842 170.332~171.161 172.606~173.980 175.739~176.981 178.465~185.036 185.702~186.583 189.294~190.050	155.912~156.842 181.244~181.286
5	9.944~10.018 155.423~159.990 188.176~200.181	1.128~3.404 10.351~11.126 201.218~201.669 204.942~207.519 209.887~210.713 211.628~212.586 219.599~220.017
6	133.981~134.017 145.192~146.467 152.460~154.392 155.916~157.948	166.532~168.826
染色体	QTLs共定位区间/Mb	QTNs共定位区间/Mb	重叠区间/Mb	花序发育相关基因
7	28.786~39.161 39.469~39.629 75.813~76.357 83.489~85.541 111.723~112.716 113.233~113.290 113.760~113.769 119.717~120.786 124.148~127.859 172.473~173.548¹⁰ 176.727~179.050	22.844~23.779⁸ 111.381~112.591 136.145...138.378⁹ 152.803~153.336 171.930~174.337¹⁰ 178.044~178.752	111.723~112.591 172.473~173.548 178.044~178.752	SID1⁸ TSH4⁹ RA3¹⁰
8	15.980~15.989 136.686~137.739 173.517~173.711 175.355~175.398 175.627~176.060	17.301~21.477^11,12 169.183~170.471 177.137~179.089		VT2¹¹ BIF1¹²
9	51.718~68.851 93.176~102.973	136.739~137.501 154.942~155.936
10	87.396~88.064 93.611~115.710	14.451~14.796 141.562~142.409¹³		ZFL1¹³

表S1 玉米雄穗分枝数的QTLs、QTNs热点区间总结

表S2 玉米雄穗主轴长QTLs、QTNs热点区间

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