玉米茎秆抗倒伏遗传的研究进展

doi:10.3864/j.issn.0578-1752.2021.11.002

摘要/Abstract

摘要：

茎秆倒伏严重影响玉米产量、品质和机械化收获,是当前玉米生产和育种亟待解决的主要问题之一。加强对玉米茎秆抗倒伏性的研究,对提高品种抗倒伏能力具有重要意义。本文综述了玉米茎秆倒伏的主要影响因素及其遗传特征。茎秆倒伏与茎秆自身的强度密切相关。茎秆强度越高,抗倒伏性越强。茎秆强度受茎秆所处的发育阶段、茎秆内部结构和外部形态,及其细胞壁成分等影响。处于分生组织的茎秆细胞分裂旺盛,较易折断,而进入生殖生长后,茎秆表皮、厚壁组织增厚,维管束发育成熟,对茎秆的支撑作用增强。茎秆细胞壁的主要成分——纤维素、半纤维素、木质素、可溶性糖、无机物等均可提升茎秆强度。目前,研究者借助高通量表型平台,利用玉米连锁群体和自交系群体,采用各种定位方法,鉴定到一系列影响茎秆形态、强度、细胞壁成分的相关QTL和候选基因。研究表明,基于单倍型的QTL定位方法比基于单个SNP的定位效果好。一致性QTL分析将不同遗传群体的研究整合到一起,能够提高QTL结果的通用性。茎秆强度的遗传基础复杂,受微效多基因控制,位点间具有加性效应。茎秆成分QTL中的候选基因涉及细胞壁代谢、转录因子、蛋白激酶等。MAIZEWALL是玉米细胞壁相关基因的重要数据库。目前该数据库包含1 156个玉米细胞壁生物学相关的候选基因,为该领域的深入研究提供强大的资源。已鉴定到一系列影响玉米茎秆细胞壁成分、茎秆形态和强度的基因,其功能涉及纤维素合成路径,如纤维素合成酶类、Cobra类、糖基转移酶和核糖转运蛋白类;苯丙烷路径基因,如控制bm1—bm5的相关基因;植物激素类,如赤霉素、生长素、油菜素甾醇相关基因;转录因子如NAC、MYB;miRNA（ZmmiR528）以及F-box基因（stiff1）等。今后应积极探索不同发育时期玉米茎秆倒伏的力学机制;广泛发展自然群体或育种群体进行遗传分析;采取多种定位策略,提高抗倒伏相关基因鉴定的功效;针对优良等位基因,开发各类分子标记,加强抗倒伏分子标记辅助选择。本文将为玉米茎秆抗倒伏遗传机制解析及抗倒伏玉米品种的分子育种提供参考。

关键词: 玉米, 倒伏, 茎秆, 细胞壁, 遗传研究

Abstract:

Maize stalk lodging has a great adverse effect on yield, quality and mechanized harvesting, and is one of the main problems to be solved urgently in current maize production and breeding. Strengthening the research on the lodging resistance of maize stalk will have great significance for improving the lodging resistance of maize. In this paper, we summarize the main factors affecting maize stalk lodging resistance, and their genetic mechanisms. The stalk lodging resistance is closely related to the stalk strength. The greater the stalk strength, the stronger the lodging resistance. The stalk strength is affected by the developmental stage, the internal and external structures of the stalk, and the components of the stalk cell wall. The meristem zone has vigorously dividing cells and is easily broken. After entering the reproductive growth, the rind and sclerenchyma tissue of the stalk are thickened, the vascular bundles are mature, and thus the stalk strength is enhanced. The main components of the stalk cell wall, including cellulose, hemicellulose, lignin, soluble sugars, inorganic substances, can improve the strength of the stalk. To date, based on the high-throughput phenotyping platforms, various maize linkage and natural populations, and mapping methods, researchers have identified a series of QTLs and candidate genes that affect stalk morphology, strength, and cell wall components. The studies have shown that the haplotype-based mapping method is better than SNP-based mapping method. Meta-QTL analysis integrates the mapping results of different genetic populations and can improve the versatility of QTLs. The genetic basis of stalk strength is complex, which is determined by polygenes with minor effect and additive effect. Candidate genes in the QTLs involve cell wall metabolism, transcription factors, protein kinases, and so on. MAIZEWALL is an important database of genes related to maize cell wall. So far, the database contains 1 156 candidate genes related to maize cell wall biology, which provides a powerful resource for research in this field. A series of genes affecting cell wall components, stalk morphology and stalk strength in maize have been identified. Their functions of these genes are related to cellulose synthesis pathways, such as genes of cellulose synthase, Cobra, glycosyltransferase and ribose transport; phenylpropane pathway genes, such as genes regulating bm1-bm5; plant hormones genes, such as genes related to gibberellin, auxin and brassinosteroid; transcription factors such as NAC, MYB; miRNA (ZmmiR528) and F-box genes (stiff1). In the future research, it is needed to explore the mechanical mechanism of stalk lodging at different developmental stages. Develop diverse natural populations and breeding materials for genetic analysis. Employ a various of mapping strategies to improve the efficiency of identification of the QTL and genes related to lodging resistance. Design various molecular markers based on the favorable alleles to improve the molecular marker assisted selection for lodging resistance. These efforts will promote the research of the genetic mechanism of stalk lodging resistance, and provide a reference for the molecular breeding of new varieties with strong lodging resistance.

Key words: maize, lodging, stalk, cell wall, genetic mechanism

王夏青,宋伟,张如养,陈怡凝,孙轩,赵久然. 玉米茎秆抗倒伏遗传的研究进展[J]. 中国农业科学, 2021, 54(11): 2261-2272.

WANG XiaQing,SONG Wei,ZHANG RuYang,CHEN YiNing,SUN Xuan,ZHAO JiuRan. Genetic Research Advances on Maize Stalk Lodging Resistance[J]. Scientia Agricultura Sinica, 2021, 54(11): 2261-2272.

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https://www.chinaagrisci.com/CN/Y2021/V54/I11/2261

图/表 3

图1

表1

玉米茎秆抗倒伏相关性状遗传定位统计"

序号 Order	性状 Trait	材料 Material	定位方法 Mapping method	主要结果 Main result	文献 Reference
1	茎秆弯曲强度 Stalk bending strength	216个RIL家系(B73×Ce03005) 216 RILs (B73×Ce03005)	复合区间作图 CIM	微效多基因遗传特征 Polygenic with minor effect inheritance	[10]
2	茎皮穿刺强度 Rind penetrometer strength	4692个NAM家系,及 196个IBM的RIL家系 4692 NAM, 196 IBM RILs	连锁分析、关联分析 Linkage analysis, GWAS	鉴定到与苯丙烷和纤维素合成相关的位点 QTLs were related to the synthesis of phenylpropane and cellulose	[11]
3	茎皮穿刺强度 Rind penetrometer strength	RIL家系(H127R× Chang7- 2)、(B73×By804) RILs (H127R× Chang7-2), (B73×By804)	复合区间作图 CIM	候选基因与细胞壁组分相关 Candidate genes were related to cell wall components	[12]
4	茎粗、茎秆弯曲强度、茎皮穿刺强度 Stalk diameter, stalk bending strength, rind penetrometer strength	257个自交系 257 inbred lines	多位点关联分析 Multi-locus association analysis	茎秆强度的改良可通过多个优良基因聚合实现 The improvement of stalk strength can be achieved through the accumulation of multiple favorable alleles	[13]
5	茎秆弯曲强度、茎皮穿刺强度 Stalk bending strength, rind penetrometer strength	189个RIL家系 (B73×Ki11) 189 RILs (B73×Ki11)	复合区间作图、关联分析 CIM, GWAS	鉴定到一个控制茎秆强度的基因stiff1 stiff1 dominates stalk strength	[30]
6	茎秆柔韧度 Stalk flexibility	313个F_2:3家系(J724×J724A1) 313 F_2:3(J724×J724A1)	混合群体分离分析 BSA	定位到1个控制茎秆柔韧度的QTL位点 One QTL was identified to control stalk flexibility	[14]
7	纤维素、半纤维、木质素 Cellulose, hemicellulose, lignin	368个自交系 368 inbred lines	关联分析 GWAS	候选基因涉及细胞壁代谢、转录因子、蛋白激酶 Candidate genes involve cell wall metabolism, transcription factors, protein kinases	[31]
8	酸性洗涤纤维、中性洗涤纤维 Acid detergent fiber, neutral detergent fiber	368个自交系 368 inbred lines	关联分析 GWAS	鉴定了ZmC3H2,提出56个候选基因 ZmC3H2 and 56 candidate genes were identified	[32]
9	6个细胞壁成分 6 cell wall components	188个RIL家系 (B73 ×By804) 188 RILs (B73×By804)	完备区间作图 ICIM	一半以上的QTL表型变异解释率超过10% More than half of the QTLs explained more than 10% phenotypic variation	[33]
10	木质素、葡萄糖和木糖 Lignin, glucose and xylose	263个IBM家系,以及 282个自交系 263 IBM, 282 inbred lines	连锁分析、关联分析 Linkage analysis, GWAS	鉴定到11个与木质素和含糖量有关的QTL 11 QTLs were related to lignin and sugar content	[34]
11	木质素及其单体含量 Lignin and its monomer content	242个RIL家系(F838×F286) 242 RILs (F838×F286)	复合区间作图 CIM	定位了80个QTL,包含7个热点区 80 QTLs were mapped, including 7 hot spots	[35]
12	细胞壁成分 Cell wall components	11个群体 11 populations	一致性QTL分析 Meta-QTL analysis	鉴定到与细胞壁组成、秸秆消化率相关的QTL QTLs related to cell wall composition and straw digestibility were identified	[36]
13	糖分含量 Stalk sugar content	202个RIL家系(YXD053×Y6-1) 202 RILs (YXD053×Y6-1)	复合区间作图 CIM	QTL之间有较强的上位性 QTLs with strong epistasis effect	[37]
14	株高与穗位高比例 Ratio of ear height to plant height	183个热带玉米自交系 183 tropical maize inbred lines	单倍型关联分析 Haplotype GWAS	单倍型关联分析更适用于倒伏性状的定位 Haplotype GWAS was more efficient for the mapping of lodging-related traits	[38]
15	茎粗 Stalk diameter	17个群体 17 populations	一致性QTL分析 Meta-QTL analysis	20个茎粗的Meta-QTLs 20 Meta-QTLs were related to stalk diameter	[39]
16	玉米最上节茎秆的维管束数目 Vascular bundle number at the uppermost internode of maize stalk	866个BC₂S₃,HIF材料 866 BC₂S₃, HIF	多QTL模型 Multiple QTL mapping	维管束数目受大量微效的QTL控制 Vascular bundle number was dominated by many small-effect QTLs	[40]
17	茎皮厚度、维管束数目、密度、茎粗 Rind thickness, vascular bundle number and density, stalk diameter	942个玉米自交系 942 inbred lines	关联分析 GWAS	鉴定到3个控制维管束密度的QTL位点 Three loci were associated with vascular bundle density	[41]
18	30个维管束性状 30 vascular traits	480个玉米自交系 480 inbred lines	多位点关联分析 Multi-locus association analysis	鉴定到84个维管束表型候选基因 84 candidate genes were related to vascular bundle phenotype	[42]

表1

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