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作物学报 ›› 2022, Vol. 48 ›› Issue (7): 1569-1582.doi: 10.3724/SP.J.1006.2022.12044

• 作物遗传育种·种质资源·分子遗传学 •    下一篇

水稻典败型隐性核雄性不育突变体ap90的鉴定与基因定位

陈驰1,2(), 陈代波2(), 孙志豪2, 彭泽群2, 贺登美2, 张迎信2, 程海涛1, 于萍2, 马兆慧1, 宋建3, 曹立勇2, 程式华2, 孙廉平2,*(), 占小登2,*(), 吕文彦1,*()   

  1. 1沈阳农业大学农学院, 辽宁沈阳 110866
    2中国水稻研究所/国家水稻改良中心/浙江省超级稻研究重点实验室/水稻生物学国家重点实验室, 浙江杭州 311401
    3浙江省农业科学院作物与核技术利用研究所, 浙江杭州 310021
  • 收稿日期:2021-06-29 接受日期:2021-10-19 出版日期:2022-07-12 网络出版日期:2021-11-03
  • 通讯作者: 孙廉平,占小登,吕文彦
  • 作者简介:陈驰, E-mail: 595954943@qq.com
    陈代波, E-mail: chendaibo@caas.cn第一联系人：

    **同等贡献

  • 基金资助:
    浙江省自然科学基金项目(LY17C130003);浙江省自然科学基金项目(LY21C130003);水稻生物学国家重点实验室开放课题(20200105);国家自然科学基金项目(31801440);中国农业科学院创新工程项目(CAAS-ASTIP-2013-CNRRI);辽宁省科技重大专项(2019JH1/10200001)

Characterization and genetic mapping of a classic-abortive-type recessive genic-male-sterile mutant ap90 in rice (Oryza sativa L.)

CHEN Chi1,2(), CHEN Dai-Bo2(), SUN Zhi-Hao2, PENG Ze-Qun2, Adil Abbas2, HE Deng-Mei2, ZHANG Ying-Xin2, CHENG Hai-Tao1, YU Ping2, MA Zhao-Hui1, SONG Jian3, CAO Li-Yong2, CHENG Shi-Hua2, SUN Lian-Ping2,*(), ZHAN Xiao-Deng2,*(), LYU Wen-Yan1,*()   

  1. 1College of Agronomy, Shenyang Agricultural University, Shenyang 110866, Liaoning, China
    2China National Rice Research Institute / National Center for Rice Improvement / Key Laboratory for Zhejiang Super Rice Research / State Key Laboratory of Rice Biology, Hangzhou 311401, Zhejiang, China
    3Institute of Crop and Nuclear Technology Utilization, Zhejiang Academy of Agricultural Sciences, Hangzhou 310021, Zhejiang, China
  • Received:2021-06-29 Accepted:2021-10-19 Published:2022-07-12 Published online:2021-11-03
  • Contact: SUN Lian-Ping,ZHAN Xiao-Deng,LYU Wen-Yan
  • About author:First author contact:

    **Contributed equally to this work

  • Supported by:
    Natural Science Foundation of Zhejiang Province of China(LY17C130003);Natural Science Foundation of Zhejiang Province of China(LY21C130003);Open Project of State Key Laboratory of Rice Biology(20200105);National Natural Science Foundation of China(31801440);Chinese Academy of Agricultural Sciences Innovation Project(CAAS-ASTIP-2013-CNRRI);Liaoning Provincial Major Science and Technology Projects(2019JH1/10200001)

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摘要：

通过甲基磺酸乙酯(Ethyl Methyl Sulfone, EMS)诱变籼稻恢复系中恢8015 (Zhonghui 8015, wild-type, WT)筛选获得了一个性状稳定的典败型雄性不育突变体材料abortive pollen 90 (ap90)。突变体ap90与野生型中恢8015在株高、株型、分蘖数、抽穗期等农艺性状上无显著差异, 但花药瘦小、呈淡乳黄色、花粉完全典败。花药发育不同阶段的半薄切片观察结果显示, ap90突变体内的花药壁细胞发育异常, 主要表现为绒毡层细胞降解异常, 小孢子细胞在有丝分裂阶段无法形成正常的花粉壁结构、无法完成淀粉充实过程, 最终小孢子细胞退化成细线状、花药亦无法开裂。花药和花粉外壁扫描电镜观察结果显示, 突变体ap90的花药外壁皱缩, 角质层排列更紧密; 花药内壁乌氏体形状不规则、排列紧密而混乱; 花粉细胞呈干瘪状, 花粉外壁表面的孢粉素形态异常、呈不规则排列。遗传分析表明, ap90突变性状受1对隐性核基因控制, 基因的初步定位将该突变位点定位于水稻7号染色体长臂RM21421和RM21435之间491.73 kb区间内; 进一步的Mut-Map测序分析证实该区间内LOC_Os07g22850的第2个外显子区存在1个37 bp的缺失和1处单碱基替换, 使得其编码序列发生移码、转录和翻译提前终止, 导致出现ap90中的花粉完全典败和小穗不育表型。表达模式分析结果显示, 该基因在花药中特异表达, OsAP90蛋白主要定位在内质网上。qPCR检测结果表明, ap90突变体中许多雄性不育相关基因的表达量受到了该突变位点的影响, 进一步证明OsAP90在水稻花药发育的乌氏体形成、花粉壁发育中有重要作用。

关键词: 水稻, 隐性核雄性不育, ap90, Mut-Map, 表达模式分析

Abstract:

In this study, we obtained a stable male sterile mutant abortive pollen 90 (ap90) from the mutant library of Zhonghui 8015 (wild-type, WT), an indica restorer line, induced by Ethyl Methy Sulfone (EMS). Compared to the WT, the ap90 mutant displayed no significant differences in plant height, plant type, tiller number, heading date and other agronomic traits, but the anthers were thinner, light creamy yellow and pollen grains were completely abortive. Semi-thin sections observation of anther development at different stages showed that the ap90 mutant carried an abnormal development process of anther wall cells. Namely, the tapetum cell degradation was obviously abnormal, the microspore cells could not form a normal pollen wall structure during mitosis and the starch filling process was blocked which eventually resulted in the degradation of microspores into threadlets and failure of anther dehiscence. Scanning electron microscopic (SEM) observations of the anther surface and pollen exine suggested that the anther epidermis of mutant ap90 were shrunk and covered by more compactly arranged cuticles. The shape of Ubisch distributed on the inner surface of anther locule were irregular, closely arranged and disordered. The pollen grains were shriveled and the sporopollenin on the pollen exine were abnormally arranged. Genetic analysis showed that the ap90 phenotype was controlled by a pair of recessive nuclear genes. Gene preliminary mapping located the mutation site into a 491.73 kb interval between RM21421 and RM21435 on the long arm of rice chromosome 7. Further Mut-Map sequencing analysis confirmed that there was a 37 bp deletion and a following single base substitution in the second exon region of LOC_Os07g22850 in the ap90 mutant, which resulted in the shifted coding sequence, prematurely terminated transcription and translation, leading to the entire abortive pollen and sterile spikelet in the ap90 mutant. Expression pattern analysis results demonstrated that the OsAP90 gene was specifically expressed in anthers and the OsAP90 protein was mainly located in the endoplasmic reticulum (ER). The qPCR results suggested that the relative expression level of many male sterility-related genes in the ap90 mutant was affected by the mutation site, which further proved that OsAP90 played an important role during the formation of Ubisch and pollen wall in rice anther development.

Key words: rice, recessive genic male sterility, ap90, Mut-Map, the relative expression pattern

图1

野生型中恢8015 (WT)与突变体ap90的表型分析 A: 野生型中恢8015和ap90突变体抽穗期的植株; 标尺为10 cm。B: 野生型和ap90突变体的小穗; C: 野生型和ap90突变体去除内外桴的颖花形态; D: 野生型和ap90突变体的花药表型; 标尺为2 mm。E: 野生型的花粉镜检形态; F: ap90突变体的花药压片; 标尺为0.1 mm。"

图2

野生型和ap90突变体不同发育时期花药的半薄横切观察 A~E: 野生型花药横切图; F~J: 突变体ap90花药横切图。A, F: 花药发育第8a 时期; B, G: 花药发育第9时期; C, H: 第10时期; D, I: 花药发育第11时期; E, J: 花药发育第13时期。Ep: 表皮; En: 内皮层; ML: 中层; T: 绒毡层; Dy: 二分体; Msp: 小孢子; Bp: 二孢花粉; Mp: 成熟花粉。"

图3

野生型和ap90花药壁和花粉细胞的扫描电镜观察 A, B: 野生型和突变体第12时期花药的整体形态; C~F: 花药外壁; G, H: 花药内壁; I, J: 花粉细胞; K~N: 花粉外壁。aUb: 不正常的乌氏体; Ub: 乌氏体; dSp: 畸形的孢粉素; Sp: 孢粉素。放大倍数已在图中标注。"

表1

ap90 突变位点的遗传分析"

组合
Combination		 F1单株结实率
Seed-setting rate of F1		 F2 χ(3:1) χ20.05
野生型 No. of wild type plants 突变体 No. of mutant plants
ap90/ZH8015 82.32 365 115 0.28 3.84
ap90/02428 86.59 3322 1078 0.59

图4

雄性不育基因OsAP90的基因定位 A: ap90位点的连锁筛选与初步定位; B: Mut-Map对AP90位点的精细定位; C: 目标基因的基因结构及ap90位点; D: 突变位点引起的氨基酸变化; E: 功能标记在BC2F2中的分离鉴定。"

图5

雄性不育基因OsAP90的组织特异性表达分析 以Ubiquitin作为内参基因, 误差线表示n = 3的标准误。MMCS、MS、YMS、PMS、MPS分别代表花药发育的花粉母细胞形成期、减数分裂期、小孢子细胞时期、有丝分裂期和成熟花粉时期; WT: 野生型; ap90: 突变体。所有数据均为平均值± SD (n = 3)。**在P = 0.01水平上差异显著。*在P = 0.05水平上差异显著。"

图6

AP90 蛋白的亚细胞定位"

图7

实时荧光定量表达分析野生型和ap90突变体花药发育相关基因的表达 以Ubiquitin作为内参基因, 误差线表示n = 3的标准误。MMCS、MS、YMS、PMS、MPS分别代表花药发育的花粉母细胞形成期、减数分裂期、小孢子细胞时期、有丝分裂期和成熟花粉时期; WT: 野生型; ap90: 突变体。所有数据均为平均值± SD (n = 3)。**在P = 0.01水平上差异显著。*在P = 0.05水平上差异显著。"

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