作物杂志, 2026, 42(5): 15-22 doi: 10.16035/j.issn.1001-7283.2026.05.002

专题综述

多维度分析小麦粒重基因:研究现状与育种展望

许娜丽,, 刘晓宇, 汪丽平, 钟思志, 周国勤, 尹志刚, 陈宏, 陈真真, 马家运, 陈俊, 朱保磊,, 石守设,

信阳市农业科学院, 464000, 河南信阳

Multidimensional Analysis of Wheat Grain Weight Genes: Research Status and Breeding Prospects

Xu Nali,, Liu Xiaoyu, Wang Liping, Zhong Sizhi, Zhou Guoqin, Yin Zhigang, Chen Hong, Chen Zhenzhen, Ma Jiayun, Chen Jun, Zhu Baolei,, Shi Shoushe,

Xinyang Academy of Agricultural Sciences, Xinyang 464000, Henan, China

通讯作者: 朱保磊,主要从事小麦遗传育种研究,E-mail:1067354457@qq.com;石守设为共同通信作者,主要从事小麦遗传育种研究,E-mail:shiss68@163.com

收稿日期: 2025-08-24   修回日期: 2025-09-30   网络出版日期: 2026-03-16

基金资助: 国家小麦产业技术体系项目(CARS-03-01A)
河南省小麦产业技术体系项目(Z2010-01-01)
河南省科技攻关项目(252102111143)

Received: 2025-08-24   Revised: 2025-09-30   Online: 2026-03-16

作者简介 About authors

许娜丽,主要从事小麦遗传育种研究,E-mail:441259781@qq.com

摘要

小麦是世界上三大粮食作物之一,对保障粮食安全有着极其重要的作用,而粒重是影响小麦产量的三要素之一。目前,国内外学者对粒重基因的研究很多,明确粒重是多基因控制的数量性状,受多种因素的影响。本文基于小麦粒重的影响因素、生物学调控途径、遗传定位分析、粒重相关基因的克隆及其变异类型、粒重分子标记在辅助育种中的应用5个方面,系统论述小麦粒重基因的研究进展,明确了小麦粒重是由多基因通过多种途径调控完成,遗传定位发掘小麦粒重相关数量性状位点(QTL)分布情况,同时明确现有粒重基因的优异等位变异及在小麦种质资源中的利用情况,为后续小麦产量性状的改良以及培育高产品种提供理论依据。

关键词: 小麦; 粒重; 基因; 调控途径; 遗传定位; 等位变异

Abstract

Wheat is one of the three major food crops in the world and plays an extremely important role in ensuring food security, while grain weight is one of the three yield components restricting yield levels. At present, domestic and foreign scholars have conducted extensive research on grain weight genes, confirming that grain weight is a quantitative trait controlled by multiple genes and is affected by many factors. In this paper, the research progress of wheat grain weight genes is systematically reviewed based on five aspects: the influencing factors of wheat grain weight, biological regulation pathways, genetic mapping analysis, cloning and variation types of grain weight-related genes, and the application of grain weight molecular markers in assisted breeding. It clarifies that wheat grain weight is regulated by multiple genes through multiple pathways, summarizes the distribution of quantitative trait loci (QTLs) related to wheat grain weight, and reveals the favorable allelic variation of existing grain weight genes and their utilization in wheat germplasm resources, which provides a theoretical basis for the improvement of wheat yield traits and the cultivation of high-yield wheat varieties.

Keywords: Wheat; Grain weight; Gene; Regulation pathway; Genetic mapping; Allelic variation

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本文引用格式

许娜丽, 刘晓宇, 汪丽平, 钟思志, 周国勤, 尹志刚, 陈宏, 陈真真, 马家运, 陈俊, 朱保磊, 石守设. 多维度分析小麦粒重基因:研究现状与育种展望. 作物杂志, 2026, 42(5): 15-22 doi:10.16035/j.issn.1001-7283.2026.05.002

Xu Nali, Liu Xiaoyu, Wang Liping, Zhong Sizhi, Zhou Guoqin, Yin Zhigang, Chen Hong, Chen Zhenzhen, Ma Jiayun, Chen Jun, Zhu Baolei, Shi Shoushe. Multidimensional Analysis of Wheat Grain Weight Genes: Research Status and Breeding Prospects. Crops, 2026, 42(5): 15-22 doi:10.16035/j.issn.1001-7283.2026.05.002

小麦(Triticum aestivum L.)作为世界及我国最重要的粮食作物之一,为我国的粮食生产和安全提供了重要保障,也为全球提供了21%的能量和20%的蛋白质,是全球近35%~40%人口的主要粮食来源[1]。在城镇化快速推进的背景下,提高国家粮食产量是当务之急,而小麦产量的高低与其穗粒数、有效穗数和粒重相关,其中粒重是控制产量的直接因素,在产量三要素中,粒重的遗传力高且稳定,受环境的影响也相对较小[2-4]。因此,粒重研究对小麦产量的提高具有重要意义。张泽源等[5]对216个家系重组自交系(RIL)群体进行55K SNP基因型分析,表明主效数量性状位点(QTL)Qtkw. nwafu-2D.1和Qtkw.nwafu-7D利用加性互作效应可显著提高小麦的千粒重和粒宽。黄倩楠等[6]认为粒重对单株产量有直接正向影响,可以作为提高新疆冬小麦单株产量的筛选指标。梁倩等[7]利用小麦100K液相芯片,对以扬麦18和扬麦22为双亲构建的RIL群体进行分析,发现粒型与粒重间存在显著相关性。张天星等[8]利用30份小麦材料进行目标基因核苷酸多态性检测,并用323份普通小麦群体进行验证,结果发现在千粒重和籽粒大小方面,含有TaCYP78A17-Ap-HapⅡ或TaCYP78A17-Ap- HapⅢ单倍型的小麦显著低于含有TaCYP78A17- Ap-HapI单倍型的小麦,且单倍型之间的差异明显。随着对小麦粒重基因的研究逐渐深入,通过增加粒重来提高小麦产量是目前最经济有效的方式之一。本研究从小麦粒重的影响因素、生物学调控机制、遗传定位分析、粒重相关基因的克隆及其变异类型、粒重分子标记在辅助育种中的应用5个方面进行论述,并对后续应用进行展望,以期为小麦粒重性状的改良提供理论依据,同时促进小麦高产新品种的培育。

1 小麦粒重的影响因素

小麦粒重不但受自身因素的影响,还受外界因素的影响。外界因素包括光照[9]、温度[9]和水分[10]等,其中温度胁迫是指影响小麦生长发育、产量及品质形成的逆境,主要包括冬前冻害、早春冻害、倒春寒和花后高温。自身因素比较复杂,主要由粒长、粒宽、粒形(粒长与粒宽之比)、粒厚和粒体积等要素构成,是多基因控制的数量性状,受遗传因素的影响;此外,自身因素还包括籽粒灌浆速率和持续时间,灌浆速率受基因型控制,与粒重呈显著正相关,而特定地区的气候和耕作栽培制度决定小麦灌浆持续的时间[11-12],一般情况下,小麦灌浆持续时间越长,其体内积累的物质越多,产量也就越高。粒重是提高产量的直接因素,而籽粒灌浆速率是影响粒重的关键因素,通过增加籽粒灌浆物质累积量可提高小麦的产量。吴晓丽等[13]在3个生态点对10个四川小麦品种进行分析,结果表明粒重与灌浆速率呈显著正相关。范涛等[14]利用RIL群体开发了1对与籽粒粒长主效QTL连锁的KASP分子标记,QKl.sau-3D.2对株高、千粒重、粒宽、粒径比和穗粒重有正向影响。刘皓等[15]对硬粒小麦Altar84与粗山羊草CIGM86.940合成的双二倍体的114个株系进行分析,发现随着小麦粒长或粒宽的增加会导致粒重提高,说明粒长、粒宽与粒重具有较强的相关性,但粒长同粒重的相关性较粒宽偏强。宋红玲等[16]通过对76份山西主推小麦品种的籽粒性状进行分析,发现千粒重与籽粒周长、粒长、粒宽、籽粒面积极显著正相关。由此可知,小麦粒重受多因素影响,其中自身因素对小麦粒重起到显著正向作用。因此,需加强对小麦粒重构成因素的研究,进而提高小麦产量。

2 小麦粒重的生物学调控途径

小麦粒重的分子调控机制主要围绕两大类基因展开,一类是淀粉合成代谢相关的基因;另一类是籽粒细胞发育和分化相关的基因。其中部分基因通过对籽粒发育进行正向调节进而提高小麦产量,如候选基因TaGDSL-7D[17]、果激酶基因TaFRK2- 7B1[18]和细胞分裂素氧化酶/脱氢酶TaCKX11-D[19];部分基因通过对籽粒发育进行负向调节进而增加小麦产量,如TaSG-D1-TaGAMyb信号转导模块[20]、转录因子TaDOF7.6[21]和E3泛素连接酶基因TaSDIR1-4A[22]。通过对小麦粒重的分子调控机制进行研究分析,了解影响小麦粒重的内在因素,从而能基于分子层面对粒重性状进行改良,进而提高小麦产量。

2.1 调控因子TaFT-D1的调控途径

小麦开花关键调控因子TaFT-D1可调控粒重,编码磷脂酰乙醇胺结合蛋白PEBP的TaFT-D1作为小麦籽粒重量的候选基因,其第3个外显子中插入/缺失(InDel)(G/-)的1 bp导致蛋白质长度不同,通过促进细胞增殖和淀粉合成来增加籽粒尺寸,进而促进粒重增加[23-24]。小麦粒重正调控因子TaFDL2与下游细胞周期和淀粉合成相关基因的启动子结合,激活其表达,TaFT-D1作为TaFDL2的辅助激活因子进一步增强表达,与TaFDL2相互作用形成复合体,从而正向调控小麦粒重[24]。

2.2 CAMTA2-GCN5复合体的协同作用

组蛋白乙酰转移酶(GCN5)与钙调蛋白结合转录因子CAMTA2发生物理相互作用,并调节小麦籽粒大小和重量。CAMTA2与GCN5形成复合体,直接与淀粉合成基因Sus2和SBEIc的启动子结合,并在小麦胚乳发育过程中建立H3K9ac和H3K14ac组蛋白标记,进而激活Sus2和SBEIc基因表达并促进淀粉积累[25]。

2.3 TaHDA9-TaSRK-TaPsbO调控模块

TaHDA9-TaSRK通过介导光系统II中的TaPsbO来增强小麦粒长的级联调控机制。研究[26]发现TaSRK是负向调控小麦籽粒长度的关键基因,TaHDA9可直接去乙酰化TaSRK,降低其蛋白积累,从而促进粒长和粒重增加。TaSRK可在叶绿体中与TaPsbO相互作用,磷酸化TaPsbO并降低其蛋白丰度,而TaHDA9通过去乙酰化调控TaSRK进而促进TaPsbO的蛋白积累,最终影响小麦粒长和粒重[27]。

2.4 TaSnRK1-TabHLH489调控模块

TabHLH489是植物甾醇激素油菜素内酯(BR)的负调控因子,敲除TabHLH489及其同源基因会使小麦对BR的敏感性增强,籽粒变长,千粒重增加。植物能量感受器TaSnRK1α1与磷酸化的TabHLH489相互作用,促进其降解,从而促进小麦籽粒发育早期种皮细胞的伸长,糖处理诱导TaSnRK1α1蛋白积累,同时降低TabHLH489蛋白水平,共同调控小麦籽粒发育[28]。

2.5 OsGSK3-EOG1调控模块

RNA结合蛋白EOG1是水稻和小麦中粒径和重量的负调节因子,通过促进小穗壳中的细胞增殖来调节粒度。OsGSK3与EOG1相互作用并磷酸化,从而增加EOG1的稳定性。通过基因编辑技术敲除EOG1在小麦中的同源基因TaEOG1A/B/D会显著增加籽粒的长、宽和粒重,表明EOG1在水稻和小麦中具有保守的功能[29]。

2.6 转录因子MYB44调控途径

TaMYB44通过直接抑制淀粉生物合成基因的表达和调节次级代谢途径来抑制淀粉合成,负向调控胚乳淀粉合成与粒重,而TaMYB44的敲除突变体亦可显著提高籽粒淀粉含量及粒重[30]。

3 小麦粒重的遗传定位分析

粒重是影响小麦产量的三大要素之一,主要受基因和环境作用,对其进行表观分析可能会有一定的误差,结合基因分析,可对粒重的研究更为精确。因此,国内外学者基于粒重这一性状进行遗传定位分析的研究较多。马明等[31]对F2代小麦穗部和粒型性状进行基因分型(GBS)测序与QTL定位分析,在1B和2D染色体上定位到关于粒长和粒厚的QTL,在2D、3D和6D染色体上定位到关于穗密度的QTL,并且穗密度与粒长、粒重呈极显著负相关,说明穗密度越密,粒重越小。Guo等[32]对RIL群体进行QTL检测并对主效QTL QTkw1B进行精细定位,最终将其定位在1B染色体上698.15~700.19 Mb区间内。胡西宁等[33]利用55K芯片对RIL群体进行QTL定位,共检测到132个QTL,其中2个QTL簇分别位于2D和7A染色体上,同时预测到2个粒重相关的候选基因TraesCS2D02G445500和TraesCS2D02G461700。Mangini等[34]对165份圆锥小麦进行全基因组关联分析,共检测到39个与籽粒相关性状的QTL,其中有23个QTL位于1A、1B、2B、3B、4B、5A和6B染色体上。He等[35]对RIL群体进行QTL检测,在5个环境下共检测到8个QTL,QTkw.cau-3DL是3DL染色体组上的1个新型主效QTL,对千粒重和粒长均有显著影响,分别占表型变异的16.43%和13.87%。Huang等[36]利用55K SNP芯片对198个品系组成双单倍体(DH)群体进行基因分型,共检测到41个QTL,其中位于2D染色体上的千粒重QTL QTgw-2D.3和QTgw-2D.4以及位于6A染色体上的QTgw-6A.16A,解释了4.81%~18.67%的表型变异,位于3D、5B、5D和6A染色体上的5个稳定QTL,解释了5.89%~7.08%的穗粒数表型变异,1个稳定的穗粒重QTL QGws-4A解释了6.08%~6.14%的表型变异。以上研究表明小麦粒重是个受多基因控制的复杂数量性状,在21条染色体上几乎均能检测到粒重相关基因的QTL,为研究粒重基因提供了坚实的理论基础,同时也提供了下一步思路,但对小麦粒重基因的研究远不能止步于此,应该基于先进的分子技术进行更深层次的挖掘与研究。

4 小麦粒重相关基因的克隆及其变异类型

小麦属于异源六倍体,基因组庞大、重复序列较多,给克隆和分离小麦基因带来很大困难,但是随着现代分子技术的快速发展,小麦粒重相关基因的克隆也取得较快发展。尤其在水稻和拟南芥中克隆到了许多控制籽粒大小的相关基因,基于同样的方法,在小麦中也鉴定到并克隆了一些控制籽粒性状的基因,部分小麦粒重基因及其等位变异见表1。目前,已证实有45个小麦粒重相关基因分布在除1D、3B和4B染色体之外的所有染色体上[5]。TaGW2是水稻基因OsGW2的直系同源物,编码E3泛素连接酶(RING型)并控制水稻中的粒径,而在小麦中同源克隆出3个TaGW2基因,并将其定位在小麦同源组6上,即TaGW2-6A、TaGW2-6B和TaGW2-6D[37]。在小麦中关于粒重研究较深入的基因是TaGW2-6A,而TaGW2-6B对千粒重的影响比TaGW2-6A和TaGW2-6D大。TaGW2基因有8个外显子和7个内含子,cDNA长1275 bp,编码423个氨基酸,氨基酸61~103区域是RING型结构域,该结构域发挥E3泛素连接酶活性[38]。研究[22]表明E3泛素连接酶TaSDIR1-4A是粒径的负调节因子,其启动子区存在2个核苷酸变异,即Hap-4A-1和Hap-4A-2,因乙烯反应因子TaERF3可以在Hap-4A-2中充当TaSDIR1-4A的转录阻遏因子,导致TaSDIR1-4A表达水平较低,进而使得含单倍型Hap-4A-2的小麦千粒重高于含单倍型Hap-4A-1。Ma等[39]对位于小麦染色体2A上的TaCwi-A1基因进行全长基因组DNA序列测定,发现TaCwi-A1由7个外显子和6个内含子组成,总计3676 bp,开放阅读框(ORF)为1767 bp。开发了1对互补显性标记CWI21和CWI22,CWI21在粒重较低的品种中扩增了404 bp的PCR片段,而在粒重较高的品种中,CWI22产生了402 bp的PCR片段。研究[40]发现TaGS-D1基因测序显示外显子编码序列没有多样性,但在第2个内含子中存在30个单核苷酸多态性(SNP),其中40 bp InDel和3 bp InDel分别是千粒重较高和较低的基因型。Jiang等[41]在TaSus2-2B中检测到2个SNP,可扩增出423和381 bp的片段。研究[42]发现在部分品种中TaGS2-2B基因组序列第4外显子3002 bp位有“CTT”3个碱基的缺失,导致其编码蛋白第306位苯丙氨酸缺失,分别将不含和含有“CTT”缺失的TaGS2-2B基因变异类型命名为TaGS2-2B-a和TaGS2-2B-b。Wang等[43]从面包小麦中克隆出TaGS5基因,通过测序发现在TaGS5-A1第303个外显子中有1个SNP并鉴定出2个等位基因。研究[44]发现TaGS1a基因全长3415 bp,有11个外显子和10个内含子。在内含子中检测到9个SNP和2个DNA片段的插入和缺失,产生2种单倍型Hap1和Hap2。Yue等[45]在24份小麦材料的6-SFT-A2基因中共检测到13个SNP,形成3种单倍型HapI、HapⅡ和HapⅢ。Dong等[46]研究发现TaGASR7-A1的2个主要单倍型(H1c和H1g)具有相同的ORF,但在其5′和3′近端区域存在多个差异。研究[47]发现在TaCWI-4A启动子区有2个SNP位点,1个SNP位于-1523(C/T),另1个SNP位于-1527(T/C),这2个SNP分别命名为Hap-4A-C和Hap-4A-T。

表1   小麦粒重相关基因的等位变异及其序列

Table1  Allelic variation and sequence of grain weight related genes in wheat

基因
Gene
等位变异
Allelic variation
标记类型
Marker type
序列(5′-3′)
Sequence (5′-3′)
染色体位置
Chromosomal position
参考文献
Reference
TaCwi-A1
TaCwi-A1a
STS
F:GGTGATGAGTTCATGGTTAAT
R:AGAAGCCCAACATTAAATCAAC
2AL
[48]
TaCwi-A1b
F:GTGGTGATGAGTTCATGGTTAAG
R:AGAAGCCCAACATTAAATCAAC
TaGW2-6A
Hap-6A-G
CAPS
F:AGAAAGGGCTGGTGCTATGGA
R:TAACGCTTGATAAACATAGGTAAT
6AS
[48]
Hap-6A-A
F:GTTACCTCTGGTTTGGGTGTCGTG
R:ACCTCTCGAAAATCTTCCCAATTA
TaGW2-6B
Hap-6B-1
KASP
F:GTTGGTGTCATTTGTAAAGCCCA
R:CCTGGGGTGCAAAACTGTTCGGA
6B
[49]
其他
F:GTTGGTGTCATTTGTAAAGCCCC
R:CCTGGGGTGCAAAACTGTTCGGA
TaGS-D1TaGS-D1aSTSF:AACTTAGGGAGCGAAAACAA
R:CACCAAGACTGGAGATGAAA
7DS[50]
TaGS-D1b
TaSus2-2B
TaSus2-2Ba
-
F:TAAGCGATGAATTATGGC
R:GGTGTCCTTGAGCTTCTGG
2B
[51]
TaSus2-2Bb
F:CTATAGTATGAGCTGGATCAATGGC
R:GGTGTCCTTGAGCTTCTGA
TaGS2-2BTaGS2-2B-aInDelF:GAGAGGGAGCCTCTGTCCTT
R:CATCGTGGTCCTGCAGCT
2B[42]
TaGS2-2B-b
TaGS5-A1
TaGS5-A1a
CAPS
F:GTGCAATCTTGGACAAACATCAG
R:AGTGCTTTGTCAACAACAGATGC
3AS
[52]
TaGS5-A1b
F:GTGCAATCTTGGACAAACATCAT
R:AGTGCTTTGTCAACAACAGATGC
TaGS1aHap1CAPSF:CAGAACCATTTACCTCTGTGAAGTGCAGTTC
R:CAGTTACTACTTCCTCCGGTTCTAAATACTTGTC
6D[44]
Hap2
TaGS5-3ATaGS5-3A-TCAPSF:CCCTCCAACTTCACATG
R:TCATCGGTGTGTAGGAAG
3A[53]
TaGS5-3A-G
6-SFT-A2HapICAPSF:CTCTCTAGACATAATCAAAAGGGA
R:TTCTTTGATCCAATGTAGCTTCA
4A[45]
HapII
HapIII
TaGASR7-A1
H1c
KASP
F:ATAACTGCTCACCCCCCACC
R:CGGGAAGGACGGGATGTATA
7A
[46]
H1g
F:CACGGTAGAGGAGCCGGTTC
R:CGGGAAGGACGGGATGTATA
TaCWI-4AHap-4A-CSNPF:AGGGCGTCCGACCAAAGTG
R:GCGACCTAGCGTGTATCAAGGAG
4A[47]
Hap-4A-T
TaTGW6-4A
TaTGW6-4Aa
KASP
F:ACGGGCCCGAGAGCGTCGCCTTC
R:TGTAGGGGCCTCGGCCCTGAGCGT
4A
[47]
TaTGW6-4Ab
F:AGTGCACGGGCCCGAGAGCGTCG
R:TGTAGGGGCCTCGGCCCTGAGCGT
TaGW8-7BTaGW8-B1aSTSF:CGCTCATCCATTCCTTCATCG
R:GCTATATGGGTTGGTGTCGC
7BS[54]
TaGW8-B1b
TaFlo2-A1TaFlo2-A1aInDelF:ACCCCTCCTCCGTTATCGTC
R:CCTCCTTCTTCTTGCGGTCG
2AL[55]
TaFlo2-A1b

新窗口打开| 下载CSV


5 小麦粒重分子标记在辅助育种中的应用

粒重对于提高小麦产量有极其重要的作用,国内外学者利用小麦粒重分子标记对小麦品种(系)进行鉴定分析,能有效准确鉴定出小麦粒重相关基因及变异类型,为后续研究提供了一定的理论依据,以期培育出高产小麦品种。曹俊梅等[56]利用KASP标记明确了14个粒重相关基因在253份小麦种质资源中的分布情况,并筛选出对育种辅助选择效应高的相关粒重基因。董海滨等[57]利用4个粒重基因分子标记鉴定了其在395份河南省小麦品种(系)中等位基因的分布情况。张维军等[58]利用4个粒重基因对209份小麦种质资源进行检测,明确不同粒重基因单倍型在种质资源中的分布情况,其中TaGW2-6B、TaGASR和TaCWI-4A能够区分粒重大小,有益于小麦粒重性状选择。李娜等[59]利用青海省育成的64份小麦品种探究粒重基因TaGW2- 6A优异等位变异,结果表明单倍型Hap-6A-G是优异等位变异,可用于小麦育种研究。时佳等[60]分析3个粒重基因TaGW2-6A、TaCwi-A1和TaSus2-2B在219份新疆小麦中的等位变异情况及分布规律,为后续小麦产量改良提供理论依据。仝靖洋等[61]分析TaGW2-6A、TaCwi-A1和TaSus2-2B在262份小麦微核心种质材料中的等位变异情况,结果表明,单倍型TaCwi-A1和Sus2-2B-H是优异等位变异。张芳等[55]利用粒重相关基因TaGS-D1和TaFlo2-A1分析其对298份国内外冬小麦品种产量的影响情况,结果表明携带TaGS-D1a和TaFlo2-A1b等位变异的小麦其千粒重分别高于携带TaGS-D1b和TaFlo2- A1a的小麦,证明其是优异等位变异。杨雪敏等[62]鉴定粒重相关基因TaCwi-A1、TaSus2-2B和TaGW2-6A在252份贵州小麦品种中的等位变异情况,结果表明其等位变异较少,不利于种质资源育种工作的开展。

6 展望

小麦作为第三大粮食作物为国家粮食安全提供了保障,有效解决了粮食可能短缺的问题。如何增进粮食产量是目前需要解决的一大难题,已知小麦粒重是影响产量的关键因素之一。当前对小麦粒重形成的生物学基础、生理基础及与环境因子的关系研究较为深入,随着生物信息学和基因组学的发展,对粒重相关基因克隆的研究逐步深入[63-64]。粒重基因TaCwi-A1、TaSus2-2B、TaGS-D1a和TaGW2-6A等应用较为广泛,而且其等位变异较易区分,能够较好地应用于小麦育种改良[55,65-66]。此外,基因编辑技术凭借其精准、高效、操作简单及可定向修饰的优势,成为突破传统育种瓶颈的关键手段,为小麦粒重改良提供了全新路径[67]。其中,基因编辑技术(以CRISPR-Cas9为代表)可助力突破小麦多倍体基因组克隆难点,针对不同基因组上的同源粒重基因,设计特异性sgRNA,可精准定位并验证不同基因组拷贝的功能。同时,对初步定位的候选基因进行敲除或激活,能快速验证其与粒重的关联,大幅缩短基因克隆与功能验证的周期。例如Zhang等[68]通过CRISPR/Cas9技术对TaGW2-6A进行编辑,创制了单突变体、双突变体和三突变体材料。基因编辑还能激活野生近缘种中沉默的隐性高粒重基因,如敲除其抑制因子,将野生资源的遗传潜力转化为可用育种材料;此外,对推广品种中的粒重基因进行定向编辑以提升千粒重,可通过精准靶点设计避免影响邻近的抗病、抗逆基因,解决传统育种的性状连锁问题。

目前对粒重基因的研究多是基因克隆与种质资源的鉴定,因此发掘粒重基因的遗传机制使其进入到生产实践中尤为重要,为了能让粒重基因更好地应用于小麦育种选择中,要做到以下几点:一、将禾本科植物(水稻、玉米等)及模式植物(拟南芥)中已发现的功能明确的粒重基因(如水稻的GW2和GW8基因)作为小麦粒重基因的研究依据,依据基因的同源性和共线性,同时依托高通量测序、基因编辑和蛋白互作分析等高科技技术平台,利用转录组测序、芯片技术和比较基因组学等方法对小麦基因组进行精细定位,克隆具有保守功能的同源基因。针对小麦多倍体基因组的复杂性,需重点挖掘非同源染色体上的特异性基因片段,开发可精准鉴定小麦粒重大小的分子标记,为后续基因筛选与应用奠定基础。二、针对已克隆的小麦粒重基因,借助亚细胞定位、生物信息学分析(如基因序列比对、蛋白结构预测)与转录组分析(如差异表达基因筛选、代谢通路富集分析)等技术手段分析基因在小麦籽粒形成、灌浆和成熟期等关键生育期的表达模式。同时,深入探究基因在激素代谢(如赤霉素和生长素调控)与信号传导(如蔗糖信号和胁迫响应信号通路)中的作用,明确基因在籽粒生长发育和光合产物转运分配等过程的分子调控机制。三、利用已开发的粒重基因分子标记,对小麦种质资源(包括地方品种、野生近缘种和育种中间材料)进行鉴定分析,明确优异粒重基因在不同生态区和不同遗传背景下的分布规律,筛选出携带优异基因的核心种质资源。在此基础上,通过杂交育种和回交育种等传统手段,将优异粒重基因与抗病、抗逆及品质等优良性状基因进行聚合,培育兼顾千粒重提升与综合农艺、品质性状优良的小麦新品种。同时,结合分子标记辅助选择技术,可缩短育种周期,提高优异基因的传递效率,加速育种成果向生产实践转化。四、优化基因编辑转化方法(如采用农杆菌介导转化或基因枪转化),提高编辑植株的获得率,加强编辑后代的田间鉴定与分子检测,确保编辑位点的稳定性,避免脱靶效应对其他农艺性状产生不利影响。结合单细胞测序与合成生物学,可精准解析粒重基因在籽粒发育不同细胞中的表达规律,为基因编辑提供更精细的靶点设计,推动小麦粒重改良从“定向修饰”向“精准调控”升级。

小麦产量由有效穗数、穗粒数、千粒重三大要素共同决定,三者的协调提升是实现小麦增产的关键。粒重基因的挖掘与应用可直接突破千粒重的遗传限制,同时通过调控粒重与其他产量要素的协同关系,进一步提升小麦整体产量潜力。在未来研究中,需持续深化粒重基因的分子调控机制研究,结合基因组学和合成生物学等前沿手段,开发更高效的基因编辑与育种技术,以“高产、优质、广适”为目标培育小麦品种,为保障全球粮食安全提供重要的品种与技术支撑。

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【目的】小麦是世界总产量第二的粮食作物,而粒重是影响小麦产量的重要因素。以和尚头(HST)和陇春23(LC23)衍生的216个家系重组自交系(recombinant inbred lines,RIL)群体为材料,基于55K SNP基因型数据,针对小麦粒重相关性状进行QTL定位,开发和验证粒长主效QTL的共分离标记,为分子标记辅助选择育种提供参考。【方法】利用小麦55K SNP芯片对亲本和RIL群体进行基因分型,构建高密度遗传连锁图谱,并与中国春参考基因组IWGSC RefSeq v1.0进行相关性分析。基于完备区间作图法对多环境粒重相关性状进行QTL定位;通过对主效QTL进行方差分析,判断不同QTL间的加性互作效应,并分析其对粒重相关性状的影响。同时,根据粒长主效QTL的共分离SNP位点开发相应的竞争性等位基因特异性PCR标记(kompetitive allele specific PCR,KASP),并在242份国内外小麦种质构成的自然群体中进行验证。【结果】构建了和尚头/陇春23 RIL群体的高密度遗传图谱,全长4 543 cM,共包含22个连锁群,覆盖小麦21条染色体,平均遗传距离为1.7 cM。遗传图谱与物理图谱具有显著相关性,Pearson相关系数为0.77—0.99(P<0.001)。共检测到51个粒重相关QTL,其中,有4个为3个及以上环境稳定表达的主效QTL,分布在2D、5A、6B和7D染色体。根据物理区间和功能标记分析主效QTL Qtkw.nwafu-2D.1和Qtkw.nwafu-7D分别为光周期基因Ppd-D1和开花基因FT-D1,方差分析表明,二者具有显著的互作效应;Qtkw.nwafu-2D.1和Qtkw.nwafu-7D优异等位基因的聚合显著提高了小麦的千粒重和粒宽。此外,根据粒长主效位点Qgl.nwafu-5A的共分离SNP开发了相应的KASP分子标记AX-111067709,该标记在242份小麦组成的自然群体中与粒长和粒重性状显著相关,在不同环境下能增加粒长3.33%—4.59%(P<0.001)和粒重5.70%—10.35%(P<0.05)。【结论】和尚头(HST)和陇春23(LC23)的粒重相关性状由多个遗传位点控制,其中,Qtkw.nwafu-2D.1和Qtkw.nwafu-7D通过加性互作效应可显著提高小麦的千粒重和粒宽。Qgl.nwafu-5A与粒重和粒长具有显著相关性,其共分离分子标记AX-11106770可应用于分子标记辅助选择育种。

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新疆冬小麦种质资源主要农艺性状遗传多样性分析

新疆农业科学, 2023, 60(5):1050-1058.

DOI:10.6048/j.issn.1001-4330.2023.05.002      [本文引用: 1]

【目的】研究新疆冬小麦种质资源主要农艺性状遗传多样性。【方法】分析232份冬小麦种质资源的株高、有效分蘖数、千粒重等9个主要农艺性状,进行遗传多样性、相关性和聚类综合评价。【结果】新疆232份小麦品种遗传多样性指数较高,9个农艺性状的多样性指数介于1.68~2.10,变异类型丰富,变异系数幅度为10.3%~43.6%,各农艺性状变异系数顺序为单株产量>有效分蘖数>主穗粒重>主穗长>主穗粒数>最多粒数>株高>小穗数>千粒重;主要农艺性状与单株产量的相关程度为:有效分蘖数(0.838)>主穗粒重(0.564)>主穗粒数(0.559)>最多粒数(0.301)>千粒重(0.269)>主穗长(0.224)>株高(-0.196)>小穗数(-0.102),有效分蘖数、主穗粒重、主穗粒数、千粒重对单株产量的直接通径系数和相关系数均达显著水平;在欧式距离为16 的位置将232个品种划分为5个类群,分析各类群的9个农艺性状特征发现,类群间存在较大差异。【结论】新疆冬小麦种质资源主要农艺性状变异较大,且遗传多样性丰富。有效分蘖数、主穗粒重、主穗粒数、千粒重对单株产量的影响是直接且正向的,可以作为提高单株产量的筛选指标,且有效分蘖数这一性状应优先选择。

梁倩, 刘颖, 石如晴, 等.

小麦粒重和粒型相关QTL定位及遗传效应分析

江苏农业科学, 2025, 53(8):122-130.

[本文引用: 1]

张天星, 李梦, 吴林楠, 等.

小麦籽粒大小相关基因TaCYP78A17的功能标记开发

作物学报, 2024, 50(12):3025-3034.

DOI:10.3724/SP.J.1006.2024.41009      [本文引用: 1]

籽粒大小是影响小麦产量的重要因素之一。为了开发小麦籽粒大小相关的功能标记, 本研究克隆得到一个潜在籽粒大小相关基因TaCYP78A17, 并对其进行了系统进化、表达模式、等位变异分析和功能标记开发。结果表明,TaCYP78A17基因是小麦细胞色素P450 CYP78A家族的一员, 且在小麦幼穗和籽粒中高表达; 在30份普通小麦品种中发现TaCYP78A17-Ap存在6个SNP和2个InDel; 根据InDel 4和InDel 8开发功能标记InDel-A17, 可将30份普通小麦品种分为TaCYP78A17-Ap-HapI, TaCYP78A17-Ap-HapII和TaCYP78A17-Ap-HapIII三种单倍型; 利用该功能标记在323份普通小麦品种中进行验证, 发现该功能标记可以有效区分上述3种单倍型; 表型调查发现具有TaCYP78A17-Ap-HapI单倍型的小麦其千粒重和籽粒大小显著高于具有TaCYP78A17-Ap-HapII或TaCYP78A17- Ap-HapIII单倍型的小麦。研究结果有望应用于小麦分子标记辅助选择育种。

Li X L, Lv X, Wang X H, et al.

Effects of abiotic stress on anthocyanin accumulation and grain weight in purple wheat

Crop and Pasture Science, 2018, 69(12):1208-1214.

DOI:10.1071/CP18341      URL     [本文引用: 2]

This study explored the effects of abiotic stress on anthocyanin accumulation and grain weight of purple wheat (Triticum aestivum L. cv. Guizi 1). Plants were treated with five abiotic stress factors at different developmental periods of the grain, then mature grains were analysed for anthocyanin content and 1000-grain weight (TGW). Stresses included temperature (cold and heat), light (shading irradiance and continuous irradiance), drought (20 and 40 mm PEG 6000), salinity (100 and 200 mm NaCl), and wounding (four types). Of all treatments, 25% tended to increase both anthocyanin content and TGW, although only 5% reached a significant level, 27.5% decreased both, 40% increased anthocyanins but decreased TGW, and 7.5% increased TGW but decreased anthocyanins. Heat stress tended to increase anthocyanin content and decrease TGW, whereas cold stress increased anthocyanins only at 15 days post-anthesis (DPA) and decreased both traits at other times. Shading irradiance decreased anthocyanins but had little effect on TGW. Continuous irradiance increased anthocyanins and TGW at 35 DPA but decreased both at 10 DPA. The lower levels of drought (20 mm PEG 6000) and salinity (100 mm NaCl) stress tended to increase anthocyanins and decrease TGW; however, the effects of 40 mm PEG 6000 and 200 mm NaCl were different, and were period-dependent. The four types of wounds increased anthocyanins and TGW at 30 DPA, but their individual effects were different in the other four periods. The results quantified the effects of five abiotic stress factors on anthocyanin accumulation and grain weight, and provide the foundation for further study of defence-responsive molecular mechanisms of abiotic stress in purple wheat.

倪胜利, 何瑞, 刘媛, 等.

不同水分条件下小麦粒重QTL定位及其元分析

甘肃农业大学学报, 2021, 56(3):45-54.

[本文引用: 1]

苗永杰, 阎俊, 赵德辉, 等.

黄淮麦区小麦主栽品种粒重与籽粒灌浆特性的关系

作物学报, 2018, 44(2):260-267.

[本文引用: 1]

王丽娜, 韩玉林, 邹少奎, 等.

不同小麦品种粒重与籽粒灌浆特性探究

山东农业科学, 2019, 51(10):40-44.

[本文引用: 1]

吴晓丽, 汤永禄, 李朝苏, 等.

四川盆地小麦籽粒的灌浆特性

作物学报, 2014, 40(2):337-345.

[本文引用: 1]

范涛, 李治, 蒋庆, 等.

小麦单位面积穗数和粒长主效QTL紧密连锁KASP标记的开发及其效应评价

中国农业科学, 2021, 54(14):2941-2951.

DOI:10.3864/j.issn.0578-1752.2021.14.002      [本文引用: 1]

【目的】小麦单位面积穗数和籽粒粒长是小麦产量相关的重要农艺性状,对其进行遗传改良有利于提高小麦的产量。通过对前期QTL定位鉴定到的提高单位面积穗数的主效QTL位点QSn.sau-2D.2和提高籽粒粒长的主效QTL位点QKl.sau-3D.2开发相应的KASP分子标记,并在川农18和T1208构建的RILs群体中进行验证及评价,为更好地利用这两个QTL以及分子标记辅助育种奠定基础。【方法】利用前期在川农18和T1208构建的高代自交群体中鉴定到的控制小麦单位面积穗数主效QTL位点QSn.sau-2D.2和控制籽粒粒长主效QTL位点QKl.sau-3D.2,结合在这两个QTL区间内的55K SNP分子标记序列,开发设计KASP分子标记,并在亲本间筛选具有多态性的KASP分子标记。将筛选到的KASP分子标记在川农18×T1208的RILs群体中分别进行基因分型和鉴定相应表型性状的高低,并分析这两个主效QTL对于其他农艺性状的影响。【结果】KASP-AX-111151907和KASP-AX-109962767在亲本中具有多态性,KASP-AX-111151907和KASP-AX-109962767在群体中的验证表明这两个分子标记分别与QSn.sau-2D.2和QKl.sau-3D.2连锁。KASP-AX-111151907和KASP-AX-10996276能将群体材料的基因型分为2类,按照表型划分,在3年试验中,KASP-AX-111151907对多穗材料的平均选择率均达到72.58%,对少穗材料的平均选择率达到71.68%;KASP-AX-10996276对长粒材料的平均选择率达到69.86%,对短粒基因型的平均选择率可达61.52%,表明这两个标记的可靠性。基于KASP分子标记的基因分型结果表明,这两个QTL对于株高、千粒重、粒长、粒宽、粒径比、单位面积穗数、穗粒重均具有显著性影响。在川农17×川农11的RILs群体中进行验证也表明这两个分子标记对相应性状的选择具有一定的作用。【结论】针对单位面积穗数主效QTL位点QSn.sau-2D.2和籽粒粒长主效QTL位点QKl.sau-3D.2分别开发了1对与之连锁的KASP分子标记,可用于相应性状的选择,与KASP标记连锁的QTL分别能显著提高单位面积穗数和籽粒粒长。QSn.sau-2D.2对株高、千粒重、粒长、粒宽、粒径比、穗粒重是负向影响,QKl.sau-3D.2对株高、千粒重、粒宽、粒径比和穗粒重是正向影响,但对单位面积穗数是负向影响,这两个QTL及开发的KASP标记可应用于小麦高产育种中。

刘皓, 王华忠, 王立晖, 等.

小麦粒长粒宽和粒重的相关性分析

农产品加工, 2015, 6(12):57-58.

[本文引用: 1]

宋红玲, 杨进文, 李宁, 等.

山西省主推小麦品种籽粒形态性状分析

中国农学通报, 2022, 38(13):1-6.

DOI:10.11924/j.issn.1000-6850.casb2022-0037      [本文引用: 1]

旨在研究小麦籽粒相关形态性状与粒重的关系,筛选籽粒形态性状优良品种,为小麦籽粒形态相关性状的遗传改良提供参考。以76个山西省主推小麦品种为试材,对小麦7个籽粒相关性状进行测量,运用主成分分析和聚类分析等方法对其进行分析和评价。供试材料的千粒重与籽粒面积、粒宽、籽粒周长、粒长存在极显著正相关,与籽粒长宽比负相关;依据综合评价值(D值)将参试小麦品种聚为3类,其中第Ⅰ-1类综合评价值最高,包括编号60、72的品种,平均D值达0.896,籽粒性状综合表现最好,尤其是千粒重、籽粒面积和籽粒周长。千粒重、籽粒面积和籽粒周长可作为小麦粒部性状考察的重要指标。

He F, Du D J, Chen Z Y, et al.

A natural variation in the promoter of TaGDSL-7D contributes to grain weight and yield in wheat

Plant Biotechnology Journal, 2025, 23(9):1-13.

DOI:10.1111/pbi.v23.1      URL     [本文引用: 1]

Yun L T, Chen T, Chen M, et al.

The wheat fructokinase TaFRK2-7B1 positively regulates grain size by modulating fructose metabolism

International Journal of Biological Macromolecules, 2025, 318:144939.

DOI:10.1016/j.ijbiomac.2025.144939      URL     [本文引用: 1]

Qian Q, Hu W G, Bi Z Z, et al.

The cytokinin oxidase/ dehydrogenase TaCKX11-D positively regulates grain size in wheat

Journal of Experimental Botany, 2025, 76(22):6758-6775.

DOI:10.1093/jxb/eraf196      URL     [本文引用: 1]

Grain size is one of the critical determinants of yield in wheat (Triticum aestivum) and is affected by cytokinin signaling. Cytokinin oxidase/dehydrogenase (CKX) regulates cytokinin homeostasis by catalysing its degradation. In this study, we determined that TaCKX11-D positively regulates wheat grain size. Overexpression of TaCKX11-D significantly increased grain length, width, thickness, and weight in wheat and Arabidopsis, whilst silencing TaCKX11-D in wheat decreased grain size and weight. Cytological analysis revealed that TaCKX11-D affected wheat grain size by regulating the cell size in the outer pericarp. Overexpression of TaCKX11-D was also found to reduce the endogenous cytokinin content in wheat, demonstrating that it promotes cell expansion and increases grain size via regulation of cytokinin homeostasis. Furthermore, we found that TaCKX11-D directly interacts with and is phosphorylated by TaMPK3 and TaMPK6. Our findings provide potentially valuable insights for breeding improved wheat varieties.

Zhou Y, Cui G X, Wei S J, et al.

TaSG-D1-TaGAMyb signaling module regulates seed weight in wheat (Triticum aestivum L.)

The Plant Journal, 2025, 123(2):e70377.

DOI:10.1111/tpj.v123.2      URL     [本文引用: 1]

Zhang P P, Gao S, Jin J H, et al.

Suppression of the transcription factor TaDOF7.6 enhances photosynthesis and energy metabolism to boost wheat yield

Journal of Agricultural and Food Chemistry, 2025, 73(26):16660-16671.

DOI:10.1021/acs.jafc.5c03339      URL     [本文引用: 1]

Wang J Y, Wang R T, Mao X G, et al.

RING finger ubiquitin E3 ligase gene TaSDIR1-4A contributes to grain size in common wheat

Journal of Experimental Botany, 2020, 71(18):5377-5388.

DOI:10.1093/jxb/eraa271      URL     [本文引用: 2]

Salt and drought-induced RING finger1 (SDIR1) is a RING-type E3 ubiquitin ligase that plays a key role in ABA-mediated responses to salinity and drought stress via the ubiquitination pathway in some plant species. However, its function in wheat (Triticum aestivum) is unknown. Here, we isolated a SDIR1 member in wheat, TaSDIR1-4A, and characterized its E3 ubiquitin ligase activity. DNA polymorphism assays showed the presence of two nucleotide variation sites in the promoter region of TaSDIR1-4A, leading to the detection of the haplotypes Hap-4A-1 and Hap-4A-2 in wheat populations. Association analysis showed that TaSDIR1-4A haplotypes were associated with 1000-grain weight (TGW) across a variety of different environments, including well-watered and heat-stress conditions. Genotypes with Hap-4A-2 had higher TGW than those with Hap-4A-1. Phenotypes in both gene-silenced wheat and transgenic Arabidopsis showed that TaSDIR1-4A was a negative regulator of grain size. Gene expression assays indicated that TaSDIR1-4A was most highly expressed in flag leaves, and expression was higher in Hap-4A-1 accessions than in Hap-4A-2 accessions. The difference might be attributable to the fact that TaERF3 (ethylene response factor) can act as a transcriptional repressor of TaSDIR1-4A in Hap-4A-2 but not in Hap-4A-1. Examination of modern wheat varieties shows that the favorable haplotype has been positively selected in breeding programs in China. The functional marker for TaSDIR1-4A developed in this study should be helpful for future wheat breeding.

Liu H, Zhang X T, Xu Y F, et al.

Identification and validation of quantitative trait loci for kernel traits in common wheat (Triticum aestivum L.)

BMC Plant Biology, 2020, 20(1):529.

DOI:10.1186/s12870-020-02661-4      [本文引用: 1]

Kernel weight and morphology are important traits affecting cereal yields and quality. Dissecting the genetic basis of thousand kernel weight (TKW) and its related traits is an effective method to improve wheat yield.

Zhang Y H, Liu H X, Wang Y J, et al.

TaFT-D1 positively regulates grain weight by acting as a coactivator of TaFDL2 in wheat

Plant Biotechnology Journal, 2025, 23(6):2207-2223.

DOI:10.1111/pbi.v23.6      URL     [本文引用: 2]

Zhang R J, An K X, Gao Y J, et al.

The transcription factor CAMTA2 interacts with the histone acetyltransferase GCN5 and regulates grain weight in wheat

The Plant Cell, 2024, 36(12):4895-4913.

DOI:10.1093/plcell/koae261      URL     [本文引用: 1]

Grain weight and size are major traits targeted in breeding to improve wheat (Triticum aestivum L.) yield. Here, we find that the histone acetyltransferase GENERAL CONTROL NONDEREPRESSIBLE 5 (GCN5) physically interacts with the calmodulin-binding transcription factor CAMTA2 and regulates wheat grain size and weight. gcn5 mutant grains were smaller and contained less starch. GCN5 promoted the expression of the starch biosynthesis genes SUCROSE SYNTHASE 2 (Sus2) and STARCH-BRANCHING ENZYME Ic (SBEIc) by regulating H3K9ac and H3K14ac levels in their promoters. Moreover, immunoprecipitation coupled to mass spectrometry (IP–MS) revealed that CAMTA2 physically interacts with GCN5. The CAMTA2–GCN5 complex activated Sus2 and SBEIc by directly binding to their promoters and depositing H3K9ac and H3K14ac marks during wheat endosperm development. camta2 knockout mutants exhibited similar phenotypes to gcn5 mutants, including smaller grains that contained less starch. In gcn5 mutants, transcripts of high-molecular-weight (HMW) Glutenin (Glu) genes were downregulated, leading to reduced HMW glutenin protein levels, gluten content, and sodium dodecyl sulfate (SDS) sedimentation volume. However, the association of GCN5 with Glu genes was independent of CAMTA2, since GCN5 enrichment on Glu promoters was unchanged in camta2 knockouts. Finally, we identified a CAMTA2-AH3 elite allele that corresponded with enhanced grain size and weight, serving as a candidate gene for breeding wheat varieties with improved grain weight.

陈代英. 小麦籽粒大小调控基因TaSRK的克隆及功能研究. 郑州: 河南农业大学, 2023.

[本文引用: 1]

Zhang N, Li S G, Sun Z P, et al.

A lectin receptor-like kinase TaSRK that is deacetylated by TaHDA9 regulates wheat grain length by mediating the photosystem II protein TaPsbO

Science Bulletin, 2025, 70(19):3120-3124.

DOI:10.1016/j.scib.2025.04.028      PMID:40328611      [本文引用: 1]

Lyu J Y, Wang D Z, Sun N, et al.

The TaSnRK1-TabHLH489 module integrates brassinosteroid and sugar signalling to regulate the grain length in bread wheat

Plant Biotechnology Journal, 2024, 22(7):1989-2006.

DOI:10.1111/pbi.v22.7      URL     [本文引用: 1]

Yan L, Jiao B Y, Duan P G, et al.

Control of grain size and weight by the RNA-binding protein EOG1 in rice and wheat

Cell Reports, 2024, 43(11):114856.

DOI:10.1016/j.celrep.2024.114856      URL     [本文引用: 1]

Liu Y C, Wang M M, Wang Y J, et al.

The transcription factor MYB44 suppresses starch synthesis to negatively regulate grain weight and yield in wheat and rice

Molecular Plant, 2025, 18(7):1193-1209.

DOI:10.1016/j.molp.2025.06.007      URL     [本文引用: 1]

马明, 王掌军, 李清峰, 等.

小麦穗部和粒型性状相关分析及QTL定位

现代农业科技, 2025(2):170-174.

[本文引用: 1]

Guo H R, Liu W, Yan G L, et al.

Conditional QTL analysis and fine mapping for thousand-kernel weight in common wheat

Plants, 2025, 14(12):1848.

DOI:10.3390/plants14121848      URL     [本文引用: 1]

胡西宁, 许晴, 郭祖良, 等.

小麦粒重相关性状的QTL定位

中国农业科技导报(中英文), 2026, 28(5):39-49.

[本文引用: 1]

Mangini G, Nigro D, Curci P L, et al.

Genome-wide association study identifies QTL and candidate genes for grain size and weight in a Triticum turgidum collection

The Plant Genome, 2025, 18(1):e20562.

DOI:10.1002/tpg2.v18.1      URL     [本文引用: 1]

He X, Liu J L, Ren X M, et al.

Mapping and validation of QTkw.cau-3DL, a major QTL controlling thousand-kernel weight in wheat

Theoretical and Applied Genetics, 2025, 138(2):46.

DOI:10.1007/s00122-025-04824-z      [本文引用: 1]

Huang F, Li X S, Du X Y, et al.

SNP-based identification of QTLs for thousand-grain weight and related traits in wheat 8762/Keyi 5214 DH lines

Journal of Integrative Agriculture, 2023, 22(10):2949-2960.

DOI:10.1016/j.jia.2023.03.004      [本文引用: 1]

<p class="15"> As important yield-related traits, thousand-grain weight (TGW), grain number per spike (GNS) and grain weight per spike (GWS) are crucial components of wheat production. &nbsp;To dissect their underlying genetic basis, a double haploid (DH) population comprised of 198 lines derived from 8762/Keyi 5214 was constructed. &nbsp;We then used genechip to genotype the DH population and integrated the yield-related traits TGW, GNS and GWS for QTL mapping. &nbsp;Finally, we obtained a total of 18&nbsp;942 polymorphic SNP markers and identified 41 crucial QTLs for these traits. &nbsp;Three stable QTLs for TGW were identified on chromosomes 2D (<i>QTgw-2D</i>.<i>3</i>&nbsp;and <i>QTgw-2D</i>.<i>4</i>) and 6A (<i>QTgw-6A</i>.<i>1</i>), with additive alleles all from the parent 8762, explaining 4.81–18.67% of the phenotypic variations. &nbsp;Five stable QTLs for GNS on chromosomes 3D, 5B, 5D and 6A were identified. &nbsp;<i>QGns-5D</i>.<i>1</i>&nbsp;was from parent 8762, while the other four QTLs were from parent Keyi 5214, explaining 5.89–7.08% of the GNS phenotypic variations. &nbsp;In addition, a stable GWS genetic locus <i>QGws-4A</i>.<i>3</i>&nbsp;was detected from the parent 8762, which explained 6.08–6.14% of the phenotypic variations. &nbsp;To utilize the identified QTLs, we developed STARP markers for four important QTLs, <i>Tgw2D</i>.<i>3-2</i>, <i>Tgw2D</i>.<i>4-1</i>, <i>Tgw6A</i>.<i>1</i>&nbsp;and <i>Gns3D</i>.<i>1</i>. &nbsp;Our results provide important basic resources and references for the identification and cloning of genes related to TGW, GNS and GWS in wheat.</p>

Jaiswal V, Gahlaut V, Mathur S, et al.

Identification of novel SNP in promoter sequence of TaGW2-6A associated with grain weight and other agronomic traits in wheat (Triticum aestivum L.)

PLoS One, 2017, 10(6):e129400.

[本文引用: 1]

冯烨宏, 王倩茜, 吴旭, 等.

TaGW2基因在超高产小麦‘烟农1212’中的结构和功能分析

分子植物育种. (2025-02-08) [2025-08-24]. https://link-cnki-net-s.webvpn.hnagri.org.cn/urlid/46.1068.S.20250208.1459.008.

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Ma D Y, Yan J, He Z H, et al.

Characterization of a cell wall invertase gene TaCwi-A1 on common wheat chromosome 2A and development of functional markers

Molecular Breeding, 2012, 29(1):43-52.

DOI:10.1007/s11032-010-9524-z      URL     [本文引用: 1]

Zhang Y J, Liu J D, Xia X C, et al.

TaGS-D1, an ortholog of rice OsGS3, is associated with grain weight and grain length in common wheat

Molecular Breeding, 2014, 34(3):1097-1107.

DOI:10.1007/s11032-014-0102-7      URL     [本文引用: 1]

Jiang Q Y, Hou J, Hao C Y, et al.

The wheat (T. aestivum) sucrose synthase 2 gene (TaSus2) active in endosperm development is associated with yield traits

Functional & Integrative Genomics, 2011, 11(1):49-61.

[本文引用: 1]

王沙沙, 裴星旭, 黄超, 等.

小麦TaGS2基因等位变异与粒重之间的关系分析

植物遗传资源学报, 2022, 23(5):1438-1445.

[本文引用: 2]

Wang S S, Zhang X F, Chen F, et al.

A Single-nucleotide polymorphism of TaGS5 gene revealed its association with kernel weight in Chinese bread wheat

Frontiers in Plant Science, 2015, 6:1166.

[本文引用: 1]

Guo Y, Sun J J, Zhang G Z, et al.

Haplotype, molecular marker and phenotype effects associated with mineral nutrient and grain size traits of TaGS1a in wheat

Field Crops Research, 2013, 154:119-125.

DOI:10.1016/j.fcr.2013.07.012      URL     [本文引用: 2]

Yue A Q, Li A, Mao X G, et al.

Identification and development of a functional marker from 6-SFT-A2 associated with grain weight in wheat

Molecular Breeding, 2015, 35(2):63.

DOI:10.1007/s11032-015-0266-9      URL     [本文引用: 2]

Dong L L, Wang F M, Liu T, et al.

Natural variation of TaGASR7-A1 affects grain length in common wheat under multiple cultivation conditions

Molecular Breeding, 2014, 34 (3):937-947.

DOI:10.1007/s11032-014-0087-2      URL     [本文引用: 2]

Jiang Y M, Jiang Q Y, Hao C Y, et al.

A yield-associated gene TaCWI, in wheat: its function, selection and evolution in global breeding revealed by haplotype analysis

Theoretical and Applied Genetics, 2015, 128(1):131-143.

DOI:10.1007/s00122-014-2417-5      URL     [本文引用: 3]

韩利明, 杨芳萍, 夏先春, 等.

株高、粒重及抗病相关基因在不同国家小麦品种中的分布

麦类作物学报, 2011, 31(5):824-831.

[本文引用: 2]

Qin L, Hao C Y, Hou J, et al.

Homologous haplotypes, expression, genetic effects and geographic distribution of the wheat yield gene TaGW2

BMC Plant Biology, 2014, 14(1):107.

DOI:10.1186/1471-2229-14-107      [本文引用: 1]

陈杰, 冉午玲, 陈建辉, 等.

黄淮麦区(南片)小麦粒重基因TaGS-D1和TaCwi-A1等位变异检测及效应分析

麦类作物学报, 2023, 43(4):409-416.

[本文引用: 1]

陈杰, 徐永贵, 冉午玲, 等.

小麦粒重基因在河南小麦品种中的分布及效应分析

分子植物育种. (2023-06-16)[2025-08-24]. https://link-cnki-net-s.webvpn.hnagri.org.cn/urlid/46.1068.S.20230616.1112.012.

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李静雯, 董亚超, 肖永贵, 等.

黄淮麦区部分小麦品种粒重基因TaGS5-A1的等位变异分布及其效应分析

麦类作物学报, 2021, 41(3):272-280.

[本文引用: 1]

Ma L, Li T, Hao C Y, et al.

TaGS5-3A, a grain size gene selected during wheat improvement for larger kernel and yield

Plant Biotechnology Journal, 2016, 14(5):1269-1280.

DOI:10.1111/pbi.12492      PMID:26480952      [本文引用: 1]

Grain size is a dominant component of grain weight in cereals. Earlier studies have shown that OsGS5 plays a major role in regulating both grain size and weight in rice via promotion of cell division. In this study, we isolated TaGS5 homoeologues in wheat and mapped them on chromosomes 3A, 3B and 3D. Temporal and spatial expression analysis showed that TaGS5 homoeologues were preferentially expressed in young spikes and developing grains. Two alleles of TaGS5-3A, TaGS5-3A-T and TaGS5-3A-G were identified in wheat accessions, and a functional marker was developed to discriminate them. Association analysis revealed that TaGS5-3A-T was significantly correlated with larger grain size and higher thousand kernel weight. Biochemical assays showed that TaGS5-3A-T possesses a higher enzymatic activity than TaGS5-3A-G. Transgenic rice lines overexpressing TaGS5-3A-T also exhibited larger grain size and higher thousand kernel weight than TaGS5-3A-G lines, and the transcript levels of cell cycle-related genes in TaGS5-3A-T lines were higher than those in TaGS5-3A-G lines. Furthermore, systematic evolution analysis in diploid, tetraploid and hexaploid wheat showed that TaGS5-3A underwent strong artificial selection during wheat polyploidization events and the frequency changes of two alleles demonstrated that TaGS5-3A-T was favoured in global modern wheat cultivars. These results suggest that TaGS5-3A is a positive regulator of grain size and its favoured allele TaGS5-3A-T exhibits a larger potential application in wheat high-yield breeding. © 2015 Society for Experimental Biology, Association of Applied Biologists and John Wiley & Sons Ltd.

张福彦, 程仲杰, 陈晓杰, 等.

黄淮麦区小麦粒重基因等位变异的分子鉴定及育种应用

作物学报, 2021, 47(11):2091-2098.

DOI:10.3724/SP.J.1006.2021.01083      [本文引用: 1]

采用特异性引物PCR扩增方法对183份黄淮海麦区的小麦品种(系)的粒重基因TaCwi-A1、TaGw8-B1和TaGS-D1的等位变异进行分子鉴定, 并结合2016&#x02014;2017和2017&#x02014;2018年度的千粒重表型数据, 分析不同等位变异类型对小麦粒重的影响, 从而找出优势基因型组合。结果表明, 不同年份间参试品种(系)的千粒重差异达到极显著水平(P&lt;0.01); TaCwi-A1位点上发现TaCwi-A1a和TaCwi-A1b两种等位变异, 其分布频率分别为66.7%和33.3%; TaGw8-B1位点上TaGw8-B1a等位变异分布频率较高, 为94.5%, 而TaGw8-B1b等位变异分布频率极低, 仅为5.5%; TaGS-D1位点上发现TaGS-D1a和TaGS-D1b两种等位变异, 其分布频率分别为79.8%和20.2%。不同等位变异组合的品种千粒重存在显著差异(P&lt;0.05), 其中具有3个高千粒重等位变异组合TaCwi-A1a/TaGS-D1a/TaGw8-B1a品种的平均千粒重最高, 与具有TaCwi-A1b/TaGS-D1a/TaGw8-B1a品种的平均千粒重差异不显著, 但是显著高于其他组合(P&lt;0.05)。TaCwi-A1a/TaGS-D1a/TaGw8-B1b基因型组合小麦品种(系)的平均千粒重最低。TaCwi-A1、TaGw8-B1和TaGS-D1位点上的不同等位变异均会导致小麦千粒重的显著变化, 其中TaGw8-B1和TaGS-D1位点上的等位变异对小麦粒重的影响更为重要。在参试材料中没有发现具有3个低千粒重等位变异组合TaCwi-A1b/TaGS-D1b/TaGw8-B1b的品种, 在7种不同等位变异组合中, 具有3个高千粒重等位变异组合TaCwi-A1a/TaGS-D1a/TaGw8-B1a品种的平均千粒重最高, 是优势基因型组合。

张芳, 任毅, 严勇亮, 等.

小麦TaGS-D1与TaFlo2-A1等位变异对粒重的影响

麦类作物学报, 2020, 40(5):533-546.

[本文引用: 3]

曹俊梅, 周安定, 刘娜, 等.

新疆冬小麦种质资源粒重基因的分布与育种适用性分析

麦类作物学报, 2025, 45(1):37-44.

[本文引用: 1]

董海滨, 任伟, 韩留鹏, 等.

河南省小麦品种(系)粒重相关基因等位变异的鉴定与分析

麦类作物学报, 2024, 44(8):996-1002.

[本文引用: 1]

张维军, 赵俊杰, 何进尚, 等.

宁夏小麦种质资源粒重基因KASP标记检测及验证

植物遗传资源学报, 2021, 23(2):493-507.

[本文引用: 1]

李娜, 曹东, 陈文杰, 等.

TaGW2-6A等位变异对青海省小麦育成品种千粒重的影响

麦类作物学报, 2018, 38(1):16-21.

[本文引用: 1]

时佳, 白璐, 任毅, 等.

新疆小麦TaGW2-6A、TaCwi-A1、TaSus2-2B等位变异对粒重的影响及应用

分子植物育种, 2018, 16(3):848-858.

[本文引用: 1]

仝靖洋, 李少鹏, 刘胜杰, 等.

小麦粒重基因等位变异的高通量分子检测及组合分析

麦类作物学报, 2018, 38(11):1300-1308.

[本文引用: 1]

杨雪敏, 李鲁华, 任明见, 等.

贵州小麦品种(系)粒重相关基因TaCwi-A1、TaSus2-2B和TaGW2-6A等位变异类型鉴定

南方农业学报, 2020, 51(9):2071-2081.

[本文引用: 1]

潘庆民, 于振文, 王月福, 等.

小麦粒重研究进展

山东农业大学学报, 1999, 30(1):92-97.

[本文引用: 1]

吕广德, 孙宪印, 亓晓蕾, 等.

普通小麦粒重相关基因克隆的研究进展

麦类作物学报, 2017, 37(10):1301-1308.

[本文引用: 1]

陈杰, 赵君瑶, 宋全昊, 等.

黄淮南片麦区小麦粒重基因TaSus2-2B和TaGw8-B1等位变异的分子检测

分子植物育种, 2024, 22(18):5944-5953.

[本文引用: 1]

王志伟, 王志龙, 乔祥梅, 等.

云南小麦品种(系)株高和粒重相关功能基因的KASP标记检测

种子, 2020, 39(3):1-6.

[本文引用: 1]

王超. 小麦粒重候选基因挖掘与小麦TaGW2突变体材料创制. 杨凌: 西北农林科技大学, 2023.

[本文引用: 1]

Zhang Y, Li D, Zhang D B, et al.

Analysis of the functions of TaGW2 homoeologs in wheat grain weight and protein content traits

The Plant Journal, 2018, 94(5):857-866.

DOI:10.1111/tpj.13903      PMID:29570880      [本文引用: 1]

GW2 is emerging as a key genetic determinant of grain weight in cereal crops; it has three homoeologs (TaGW2-A1, -B1 and -D1) in hexaploid common wheat (Triticum aestivum L.). Here, by analyzing the gene editing mutants that lack one (B1 or D1), two (B1 and D1) or all three (A1, B1 and D1) homoeologs of TaGW2, several insights are gained into the functions of TaGW2-B1 and -D1 in common wheat grain traits. First, both TaGW2-B1 and -D1 affect thousand-grain weight (TGW) by influencing grain width and length, but the effect conferred by TaGW2-B1 is stronger than that of TaGW2-D1. Second, there exists functional interaction between TaGW2 homoeologs because the TGW increase shown by a double mutant (lacking B1 and D1) was substantially larger than that of their single mutants. Third, both TaGW2-B1 and -D1 modulate cell number and length in the outer pericarp of developing grains, with TaGW2-B1 being more potent. Finally, TaGW2 homoeologs also affect grain protein content as this parameter was generally increased in the mutants, especially in the lines lacking two or three homoeologs. Consistent with this finding, two wheat end-use quality-related parameters, flour protein content and gluten strength, were considerably elevated in the mutants. Collectively, our data shed light on functional difference between and additive interaction of TaGW2 homoeologs in the genetic control of grain weight and protein content traits in common wheat, which may accelerate further research on this important gene and its application in wheat improvement.© 2018 The Authors The Plant Journal © 2018 John Wiley & Sons Ltd.

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