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作物学报 ›› 2023, Vol. 49 ›› Issue (8): 2025-2038.doi: 10.3724/SP.J.1006.2023.22051

• 作物遗传育种·种质资源·分子遗传学 •    下一篇

水稻OsGMS2基因的鉴定及其核不育系种子繁殖体系构建

唐杰1(), 龙湍2(), 吴春瑜1, 李新鹏1, 曾翔1, 吴永忠1, 黄培劲1,*()   

  1. 1 海南波莲科技有限公司, 海南海口 570125
    2 海南大学热带作物学院, 海南海口 570208
  • 收稿日期:2022-09-18 接受日期:2023-02-10 出版日期:2023-08-12 网络出版日期:2023-03-13
  • 通讯作者: 黄培劲
  • 作者简介:唐杰, E-mail: jesontom@126.com
    龙湍, E-mail: longtuan2001@aliyun.com第一联系人：**同等贡献
  • 基金资助:
    中国水稻研究所水稻生物学国家重点实验室开放课题项目(20190203)

Identification of OsGMS2 and construction of seed production system for genic male sterile line in rice

TANG Jie1(), LONG Tuan2(), WU Chun-Yu1, LI Xin-Peng1, ZENG Xiang1, WU Yong-Zhong1, HUANG Pei-Jin1,*()   

  1. 1 Hainan Bolian Technology Co., Ltd., Haikou 570125, Hainan, China
    2 College of Tropical Crops, Hainan University, Haikou 570208, Hainan, China
  • Received:2022-09-18 Accepted:2023-02-10 Published:2023-08-12 Published online:2023-03-13
  • Contact: HUANG Pei-Jin
  • About author:First author contact:**Contributed equally to this work
  • Supported by:
    Open Project of the State Key Laboratory of Rice Biology, China Rice Research Institute(20190203)

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摘要：

雄性不育是作物杂种优势利用的基础。本研究在籼稻93-11 60Co-γ辐射诱变突变体库中鉴定了一个普通核雄性不育突变体osgms2。突变体雄性彻底败育, 雌性育性正常, 其他农艺性状与野生型一致。遗传分析表明雄性不育稳定遗传且为单基因隐性突变, 通过与明恢63构建遗传群体将突变基因精细定位于水稻4号染色体长臂的分子标记S3b和4826之间86 kb物理区间内。通过测序分析发现定位区间内一编码类成束阿拉伯半乳聚糖蛋白的基因LOC_ Os04g48490编码区第118位至126位缺失连续9个碱基, 导致氨基酸整码突变。OsGMS2与已报道的OsFLA1为同一基因, 但是突变位点不同的新等位突变。粳稻中花11背景下的基因敲除和遗传互补实验证实了该基因的功能。实时荧光定量PCR (qPCR)表达分析模式显示, 该基因在水稻各组织均有表达, 但在花期和乳熟期种子表达量较高。蛋白同源比对分析发现, 该蛋白序列保守, 可能在不同物种中发挥重要功能。通过创建OsGMS2-T保持系成功实现了普通核不育系的繁殖。osgms2突变体的发现和基因鉴定为水稻普通核雄性不育研究及杂种优势利用提供了新的材料。

关键词: 水稻, 核雄性不育, osgms2, 基因定位, OsGMS2-T保持系

Abstract:

Male sterility is the basis for application of crop heterosis. The common nuclear male sterile mutant osgms2 was isolated from a mutant library created by 60Co-γ-treated indica cultivar 93-11. The male of the mutant was completely aborted, the female was nomal, and other agronomic characters were consistent with those of the wild type. Genetic analyses indicated that the male sterility phenotype was stably controlled by a single recessive gene. The OsGMS2 gene was fine-mapped within the 86 kb physical interval between two molecular markers S3b and 4826 on the long arm of chromosome 4 with a mapping population of osgms2 and Minghui 63. Further sequencing found the gene LOC_Os04g48490, encoding a fasciclin-like arabinogalactan protein, had a 9-bp deletion at position 118 to 126 from start codon of translation, resulting in three codons mutation. The osgms2 gene was a novel allele of OsFLA1. Subsequent gene knockout and genetic complementation experiments in Zhonghua 11 background confirmed the function of the gene. Real-time quantitative PCR (qPCR) analyses showed that the gene was expressed in all tissues, with the highest level at the flowering and immature seed stages. Alignment analyses revealed that the protein sequence was conserved in different species. By creating the OsGMS2-T maintainer, seed production of the common sterile line was achieved. The identification of the allelic mutant osgms2 provides the new materials for the study on rice nuclear male sterility and application of heterosis.

Key words: rice, genic male sterile, osgms2, gene mapping, OsGMS2-T maintainer line

表1

初定位和精细定位引物"

引物名称
Primer name		 上游引物
Forward primer (5°-3°)		 下游引物
Reverse primer (5°-3°)
RM17332 CGGTACATCACGGTATCAAATCG TAAATGCTGGAGCGATGCTAACC
RM280 GTGCTCTCCATGTCGGATTATGC CAAGGCAACAAGATTGGTTAGTGG
RM17351 ATAAAGGAGGAGGGCCTCAGATGG CACGGTTTGGAGGTTGGAAGC
RM17352 GCTTGGCATCTGCTTCTGTTGTTGG CTCGCTGCTGATCGAGGTGTCG
RM303 ATCGATGTAGGTAGAGGGACACC CAGATCTAGTCGACATGGTTGG
4826 ACACCATCTCTCTTCTTTTTCTAT ATATGGGTAGGTTTGGATATTCG
S4b GTGTGTGTGAGTAAAATCCTAGTGCA
AAAAAGTGTGTGTGAGTAAAATCCTAGAGCC		 ATTTGTACTCCTATGTTTAGAATAGC
S3b ACAAATATATAGCAAAATCGGTGACC
AAAAAACAAATATATAGCAAAATCGGTTACG		 GTGGTTTTGTGGATGTTTTGTAACT
S2 AAGTATTTGTAATGCACTATGTAAAGGT
AAAAAAAGTATTTGTAATGCACTATGTAATGGC		 TTAAGAGCACACACTTCCAATAATATGT
S1 CTGGGCGCGGTGCGGCGGGCGAGGC
AAAAACTGGGCGCGGTGCGGCGGGCGTGGT		 CCGCCTCAGCGCCACCGCCAAGCTGA
S8 AAGTTGTGTTTAGCACTATGTTATTACG
AAAAAAAGTTGTGTTTAGCACTATGTTATGACA		 TTTAGCATAATAACTACTATTCATCATT
S10 GCAGGAGACACTTGGTGCCGCCTCTC
AAAAAGCAGGAGACACTTGGTGCCGCCACTT		 GCAGATTATTTTCGGTGGGTCCCGTCTC

表2

基因克隆相关引物"

引物名称
Primer name		 上游引物
Forward primer (5°-3°)		 下游引物
Reverse primer (5°-3°)
LOC_Os04g48490_1 AAACAGAAAGCCCCAATG TGCCGCAGTACGCGCCCAAG
LOC_Os04g48490_2 TTGTCCATGCCGGTGTCCAT GGTCACGGCACAAACTCA
InD48490 GCTCCGGCTGTTGATCT GCCTGCTCTTCCTCCTG
GAPDH-RT GAATGGCTTTCCGTGTT CAAGGTCCTCCTCAACG
T1-sg GCGGTCGGTGGCGGCCATGG
SP CCCGACATAGATGCAATAACTTC GCGCGGTGTCATCTATGTTACT
B1 AAACCCACGCCCAGAAA GCCAGGAGGAAGAGCAG
3148 HB CgcgtttcgaaatttTGATTTCTTCATCGCACT GtcgcgatcgcatgcACAACATGGTGCAACAGTG

图1

野生型93-11和突变体osgms2表型鉴定 A~F: 93-11和突变体osgms2整株(A)、穗(B)、小花(C)、去壳后颖花(D)、单个花药(E)和花粉碘染(F)。"

表3

突变体osgms2的遗传分析"

杂交组合
Cross combination		 异交结实率
Out-crossing seed-setting rate		 F2 χ23:1 χ20.05
可育株Fertile plants 不育株Sterile plants
osgms2/93-11 71.27%±14.67% 57 13 0.85 3.841
osgms2/ZH11 62.94%±14.34% 80 16 0.57 3.841

图2

osgms2的精细定位 A: 突变基因初定位在4号染色体RM303和S10之间387.8 kb物理区间; B: 精细定位在标记S3b和4826之间86 kb物理区间; C: 精细定位区间内的11个预测基因, 其中标红色ORF11为候选基因; D: 候选基因LOC_Os04g48490的结构及其在osgms2突变体中的突变位点(突变体碱基AACAGCTAC缺失导致天冬酰胺、丝氨酸和酪氨酸的缺失)。"

表4

目标区域预测基因"

序号
Serial number		 基因
Gene name		 功能注释
Function annotation
ORF1 LOC_Os04g48390 SPX-Major Facility Superfamily (MFS) protein
ORF2 LOC_Os04g48400 HOTHEAD precursor
ORF3 LOC_Os04g48410 Copper chaperone for superoxide dismutase
ORF4 LOC_Os04g48416 OsSub45-putative subtilisin homologue
ORF5 LOC_Os04g48440 The expressed protein
ORF6 LOC_Os04g48450 The expressed protein
ORF7 LOC_Os04g48460 Cytochrome P450
ORF8 LOC_Os04g48470 The expressed protein
ORF9 LOC_Os04g48480 The exostosin family domain containing protein
ORF10 LOC_Os04g48484 The expressed protein
ORF11 LOC_Os04g48490 The fasciclin-like arabinogalactan protein

图3

突变体的共分离验证 osgms2扩增出140 bp的产物; MH63扩增出149 bp的产物; F2可育表型植株扩增出149 bp或同时扩增出149 bp和140 bp的产物; 雄性不育表型的植株扩增出140 bp的产物。"

图4

突变基因的敲除及遗传互补 A: 敲除载体靶序列及纯合突变株cr-osgms2-3测序结果, 黑色加粗下画线碱基为PAM区, 红色标注为AT碱基缺失; B: osgms2基因组互补载体构建图谱; C~F: ZH11和敲除纯合株cr-osgms2-3的整株(C)、花(D)、花药(E)、花粉碘染(F); G~J: ZH11和遗传互补株Com-osgms2的整株(G)、花(H)、花药(I)、花粉碘染(J)。标尺: (C) 10 cm; (D)~(E) 2 mm; (F) 0.1 mm; (G) 10 cm; (H)~(I) 2 mm; (J) 0.1 mm。"

图5

OsGMS2氨基酸序列比对"

图6

OsGMS2在水稻不同组织部位相对表达量"

图7

OsGMS2不育系繁殖 A: OsGMS2-T载体; B: OsGMS2不育系繁殖流程; C: 左、野生型ZH11 (基因型Ms/Ms), 中、OsGMS2-T保持系(基因型ms/Ms-T), 右、osgms2不育系(基因型ms/ms)花粉1%碘-碘化钾染色; D: 明场(BF)、荧光(FRFP)以及合成(Merged)的OsGMS2-T保持系种子。标尺: (C) 0.1 mm; (D) 5 mm。"

图8

T1代OsGMS2-T保持系和osgms2不育系表型鉴定 A: OsGMS2-T保持系和osgms2不育系整株; B: OsGMS2-T保持系和osgms2不育系穗; C: 荧光视野OsGMS2-T保持系种子; D: OsGMS2-T保持系花粉1%碘-碘化钾染色; E: osgms2不育系花粉1%碘-碘化钾染色。标尺: (A) 10 cm; (B) 5 cm; (C) 1 cm; (D)~(E) 0.1 mm。"

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