中国农业科学 ›› 2021, Vol. 54 ›› Issue (12): 2487-2498.doi: 10.3864/j.issn.0578-1752.2021.12.001

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

水稻粒宽突变体gw87的基因定位及候选基因分析

张翔宇(),郭佳,王三,陈聪萍,孙昌辉,邓晓建,王平荣()   

  1. 四川农业大学水稻研究所,成都611130
  • 收稿日期:2020-11-18 接受日期:2021-01-15 出版日期:2021-06-16 发布日期:2021-06-24
  • 通讯作者: 王平荣
  • 作者简介:张翔宇,E-mail:964670742@qq.com
  • 基金资助:
    国家自然科学基金(31971869);国家重点研发计划(2017YFD0100201)

Gene Mapping and Candidate Gene Analysis of Grain Width Mutant gw87 in Rice

ZHANG XiangYu(),GUO Jia,WANG San,CHEN CongPing,SUN ChangHui,DENG XiaoJian,WANG PingRong()   

  1. Rice Research Institute, Sichuan Agricultural University, Chengdu 611130
  • Received:2020-11-18 Accepted:2021-01-15 Online:2021-06-16 Published:2021-06-24
  • Contact: PingRong WANG

摘要:

【目的】对水稻粒宽突变体gw87grain width 87)进行表型鉴定、遗传分析、基因定位及候选基因分析,为探明该基因调控水稻籽粒大小的分子机制及应用潜力奠定基础。【方法】利用甲基磺酸乙酯诱变籼稻恢复系材料676R,获得一份籽粒宽度和千粒重显著增加的突变体gw87。对该突变体进行表型观察、农艺性状调查及外源油菜素内酯(BL)敏感性、叶绿素含量、光合参数测定,了解其表型特征及生理特性。调查gw87与676R杂交F1的表型和F2群体的分离情况,分析其遗传行为;选取该群体中的突变植株进行高通量测序,并利用gw87与粳稻品种日本晴杂交的F2代作为定位群体,通过MutMap分析和分子标记定位,遴选候选基因并进行DNA和cDNA测序验证。利用qRT-PCR分析gw87和676R中BR合成途径基因OsDWARF4D11D2的表达差异。【结果】与野生型亲本676R相比,gw87突变体的株高降低,节间缩短,其中,倒一节间长度缩短最大,并呈现出扭曲的形态;叶片长度减少,宽度增加;单株有效穗数、主穗穗长和结实率显著降低,但籽粒宽度和千粒重显著增大。BL敏感性试验显示,gw87突变体幼苗对外源BL的敏感性降低。光合色素和光合参数测定表明,gw87光合色素含量增加,光合速率也有所增加。遗传分析表明gw87的突变性状是由1对隐性核基因控制。MutMap分析显示gw87突变基因位于第5染色体中部,在该染色体区域仅有1个碱基突变引起编码氨基酸变化;分子标记连锁分析表明该突变基因位于InDel标记X2和X3之间约101 kb的染色体区域;综合这两方面分析结果,最后遴选出gw87候选基因是编码一个含有AP2/EREBP DNA绑定结构域的转录因子基因LOC_Os05g32270。对该候选基因进行DNA和cDNA测序验证,发现gw87突变体中该基因DNA的第1 041位的碱基由G突变为A,导致与该位点相邻的76 bp内含子序列被剪接为外显子,引起阅读框移码,蛋白翻译提前终止。qRT-PCR分析显示gw87突变体中BR合成途径基因的表达量显著上调,表明gw87突变体中BR信号减弱。【结论】gw87smos1shbrla1ngr5的新等位突变体,但与这些突变体表型不同,gw87籽粒的宽度和千粒重显著增加,可能是LOC_Os05g32270的突变位点不同,导致其编码蛋白的功能活性不同所造成。

关键词: 水稻(Oryza sativa L.), 籽粒, 千粒重, 油菜素内酯, AP2类转录因子

Abstract:

【Objective】Phenotypic identification, genetic analysis, gene mapping and candidate gene analysis of grain width mutant gw87 (grain width87) were performed in rice, so as to lay a good foundation for further understanding molecular mechanism and utilization potentiality of this gene regulating rice grain size.【Method】Through ethyl methanesulfonate mutagenesis, we isolated gw87 mutant showing significant increase of grain width and 1000-grain weight from indica restorer line material 676R. To identify phenotypic and physiological characteristics of the mutant, we performed observation of phenotypes, investigation of agronomic traits, sensitivity analysis of exogenous brassinolide (BL), determination of chlorophyll contents and photosynthetic parameters by using gw87 and its wild type. To analyze its genetic behavior, we investigated F1 phenotypes and F2 segregation of the crossing combination between gw87 and its wild-type parent 676R. To find candidate gene of gw87, we conducted high-throughput sequencing and MutMap analysis of 30 mutant plants from the F2 population of gw87×676R, and molecular mapping using the F2 population crossing gw87 and japonica cultivar Nipponbare. Meanwhile, we confirmed the candidate gene by sequencing its DNA and cDNA sequences. In addition, we detected expression levels of brassinolide synthesis pathway genes OsDWARF4, D11 and D2 using qRT-PCR analysis.【Result】Compared with the wild-type parent 676R, the gw87 mutant showed significant reduction of productive panicle number per plant, main panicle length and seed setting rate, but significant increase of grain width and 1000-grain weight. Meanwhile, the mutant displayed decreased plant height and shortened internodes, in which the first internode was distorted and its length is shortened the most. Besides, leaf length of gw87 was decreased but the width was increased. Sensitivity assays of exogenous BL suggested that the sensitivity of gw87 seedlings to exogenous BL was reduced. Determination of photosynthetic pigments and photosynthetic parameters showed that chlorophyll content and net photosynthetic rate were decreased. Genetic analysis revealed that the mutant phenotype of gw87 was governed by a single recessive nuclear gene. High-throughput sequencing and MutMap analysis suggested that the gw87 locus was located in the middle of chromosome 5, and only one base mutation causing amino acid change of the encoded protein occurred in this chromosome region. Molecular marker linkage analysis demonstrated that the gw87 gene was mapped to a 101 kb genomic region between InDel markers X2 and X3. The MutMap and molecular mapping revealed that the candidate gene was LOC_Os05g32270 that encodes a transcription factor containing the AP2/EREBP DNA-binding domain. Subsequently, we confirmed the candidate gene by sequencing its DNA and cDNA. In the gw87 mutant, a single nucleotide G-to-A substitution occurred at position 1041 in its DNA sequence, and the 76-bp intron sequence adjacent to this mutated nucleotide was spliced into exon in its cDNA, which caused shift of the reading frame and then premature termination of the protein translation. In addition, qRT-PCR analysis showed that the expression of BR synthetic genes were significantly up-regulated in gw87, indicating that BR signal was weakened in the mutant.【Conclusion】gw87 is a new allelic mutant of smos1, shb, rla1, and ngr5. However, unlike these mutants, gw87 exhibited significant increase of grain width and 1000-grain weight relative to its wild type. The reason for the differentiation could be that different mutation sites in the LOC_Os05g32270 gene, resulting in different functional activities of the encoded protein.

Key words: rice (Oryza sativa L.), grain, 1000-grain weight, brassinosteroid, AP2-type transcription factor