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作物学报 ›› 2023, Vol. 49 ›› Issue (11): 3029-3041.doi: 10.3724/SP.J.1006.2023.22061

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

水稻卷叶等位突变体e202的鉴定和基因精细定位

周文期1,2(), 强晓霞3, 李思雨1, 王森4, 卫万荣1,*()   

  1. 1西南野生动植物资源保护重点实验室 / 西华师范大学生命科学学院, 四川南充 637000
    2甘肃省农业科学院作物研究所, 甘肃兰州 730070
    3甘肃省兰州市第四中学, 甘肃兰州 730050
    4陕西省畜牧产业试验示范中心, 陕西咸阳 713702
  • 收稿日期:2022-10-28 接受日期:2023-05-24 出版日期:2023-11-12 网络出版日期:2023-06-15
  • 通讯作者: 卫万荣, E-mail: weiwr18@126.com
  • 作者简介:E-mail: zhouwenqi850202@163.com
  • 基金资助:
    2020年甘肃省科协青年科技人才托举工程项目资助

Identification of a rolling leaf allelic mutant e202 and fine mapping of E202 gene in rice

ZHOU Wen-Qi1,2(), QIANG Xiao-Xia3, LI Si-Yu1, WANG Sen4, WEI Wan-Rong1,*()   

  1. 1Key Laboratory of Southwest China Wildlife Resources Conservation / College of Life Sciences, China West Normal University, Nanchong 637000, Sichuan, China
    2Crops Research Institute, Gansu Academy of Agricultural Sciences, Lanzhou 730070, Gansu, China
    3Lanzhou No. 4 High School, Lanzhou 730050, Gansu, China
    4Shaanxi Province Animal Husbandry Industry Experimental Demonstration Center, Xianyang 713702, Shaanxi, China
  • Received:2022-10-28 Accepted:2023-05-24 Published:2023-11-12 Published online:2023-06-15
  • Supported by:
    Young Scientific and Technological Talents Support Project of Gansu Association for Science and Technology in 2020

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摘要：

良好的叶片形态是保证作物产量因素之一, 一定程度的卷曲能够使水稻叶片保持直立、不下垂, 有利于改善群体的受光面积, 使其接收更多的光能, 提高光合利用率。利用EMS诱变水稻中花11成熟种子, 从M2后代中筛选到一个叶片内卷、挺直的突变体, 命名为e202。通过表型鉴定, 发现e202表现出多种缺陷表型, 与野生型中花11相比, 不定根数目减少、根长变短, 圆锥花序发育异常, 小花形态、花药、花粉形态有畸形, 育性极显著降低, 叶绿素含量显著升高, 气孔密度降低。树脂切片观察叶片结构, 与对照相比, 由于泡状细胞数目的增加和体积的减小, 导致叶片的卷曲。同时切片观察花药, e202中仅形成极少数花粉粒, 导致育性降低。我们通过图位克隆技术将候选基因定位到第10染色体440 kb的物理区域内, 对区间内基因分析和测序发现Os10g0562700 (LOC_Os10g41310)基因第4个外显子上, 第1935位置缺失了1个碱基G, 导致后续氨基酸错乱, 蛋白翻译提前终止。综上表明, e202是REL2的一个新等位突变体, 本研究为揭示REL2参与水稻叶片卷曲和花的发育等提供了一定的理论基础。

关键词: 水稻, 卷叶, 抗旱性, 花发育, 基因精细定位

Abstract:

Good leaf morphology is one of the factors for ensuring crop yield. A certain degree of curling can keep rice leaves standing and not drooping, which is beneficial to improve the light area of the population, enabling it to receive more light energy and improve the photosynthetic efficiency. In this study, the mature seeds of ZH11 in rice were mutated by EMS, and a mutant named e202 with curly and straight leaves was screened from M2 progeny. Phenotypic identification showed that e202 exhibited a variety of defective phenotypes. Compared with the wild type Zhonghua 11 (ZH11), the length and number of adventitious roots were decreased, and the panicle development was abnormal. The morphology of floret, anther, and pollen was deformed, the fertility was significantly reduced, the chlorophyll content was significantly increased, and the stomatal density was decreased. The structure of the leaves was observed by resin sectioning. Compared with the control, the number of vesicular cells increased and the volume decreased, resulting in leaf curling. Meanwhile, the anthers were sliced and observed. Normal pollen could not be formed in e202 or only very few pollen grains were formed, resulting in reduced fertility. The candidate genes were located in the physical region of 440 kb on chromosome 10 by map-based cloning. Gene sequencing analysis within the interval revealed that one base G was missing in the 1935 position of the fourth exon of Os10g0562700 (LOC_Os10g41310) gene, which resulted in the subsequent amino acid confusion. Protein translation is terminated prematurely, and e202 is a novel allelic mutant of REL2. This study provides a theoretical basis for revealing the involvement of REL2 in leaf rolling and flower development in rice.

Key words: rice, rolled-leaf, drought resistance, flower development, fine mapping

表1

本研究使用的部分引物"

引物名称
Primer name		 引物序列
Primer sequence (5°-3°)		 用途
Use
e202-F ATGGGGTGCACGGCGTCGAAGG 基因克隆Gene cloning
e202-R CTACCGCACCGATCCAGCTCGCCT
InDel-1F TTATGTCAACCTTACAATT 基因定位Fine mapping
InDel-1R GCATACTGTATTCCTATATC
InDel-2F TTGCTTAGTGGTAAGCTATG
InDel-2R GGAGTTCTTGAAGAAACTCC
InDel-3F GTATGTTTGTGGAGTAATAT
InDel-3R CATGCAATGTGCCAAGAACC
InDel-4F GCATACGAATGTACTTGTGT
InDel-4R TGGAAGTATGCGTGGAATAT
InDel-5F CTCAAATCACTTATATTATG
InDel-5R GTGTAACCAGTAGAAGAATG
InDel-6F TGTATAATTTGTTGAGATGAG
InDel-6R GTAGATCTTGTGATATTGGAG
C10-1-F GATTCACGGGTCGATCTACT 基因定位Fine mapping
C10-1-R GTGTGGTGCCATGCCTAATG
C10-22-F CAAGTATAATAATACTTAGAG
C10-22-R GAGCAAGGCTAATAATACATC
C10-38-F GTTGAAGTGCTACAGGTATG
C10-38-R GCTTCCTCAGCCTTGATCTC
C10-45-F GTATTCGCACTCTCAAATGAG
C10-45-R CAAATTAAGTCACAGATAAGTG
C10-57-F CAAGAATTCTAAGAGCTCG
C10-57-R CGAGCTTATGCATAAGCTTTG
C10-72-F TAGGAGGAGCAGAAGGAATAG
C10-72-R CGATCGATCGTCTTCTTCCAC
TUB-F TTTCACTCTTGGTGTGAAGCAGAT 实时荧光定量PCR技术RT-PCR
TUB-R GACTTCCTTCACGATTTCATCGTAA
REL2-F TGATCATCGTGACTTCACAGGC 实时荧光定量PCR技术RT-PCR
REL2-R TCTACCAGACCACGGACTTGC

图1

e202和ZH11植株表型分析 A: 生长早期, e202比ZH11较矮, 叶片卷曲; B: e202与对照开花期植株高度一样; C: ZH11幼苗茎与叶之间夹角较大; D: e202茎与叶之间的夹角比ZH11小; E: e202苗期根系不发达, 不定根数目减少且变短; F: 左为ZH11叶, 中为e202微卷叶, 右为e202呈卷筒状叶片; G: F2代植株, e202矮化, 卷叶。A, B, G: 标尺为10 cm; C, D, E, F: 标尺为1 cm."

图2

e202部分稻穗的突变表型 A, E: ZH11完整的穗花结构; B~D, F~L: e202; A: ZH11拨开外稃和內稃的小穗结构; B~D: e202拨开颖壳, 外稃和内稃数均增加; F~H: e202稻壳的多种畸变表型; I~L: e202缺失雄蕊和雌蕊的畸变结构, 用星号标出。标尺为0.1 cm。"

图3

e202部分花及花粉的突变表型 A, I: ZH11; B~H, J~L: e202; A: ZH11外稃和內稃的花的解剖图; B~D: e202, 2个雌蕊; E~H: e202雄蕊、柱头和桨片的畸变结构; I: ZH11的花药结构; J~L: e202花粉畸变结构, e202不同形态花药, 育性降低。A~H: 标尺为 0.05 cm; I~L: 标尺为0.01 cm。"

图4

各种花器官的数目统计 A: 雄蕊; B: 雌蕊; C: 柱头; D: 桨片; E: 外稃; F: 内稃。对照统计50个花, e202统计150个花。n = 50/150。"

图5

ZH11与e202花粉染色及花药的横切图 A: ZH11开花; B: e202不开花, 散粉期颖壳不开; C: ZH11花药及花粉粒; D: e202花药及花粉, 少量花粉粒; E: ZH11成熟种子饱满; F:大多数e202种子干瘪、不饱满; G, H: ZH11和e202经过0.1%的碘-碘化钾染色的花粉; e202能被染色花粉极少; I, J: ZH11花药横切图; K, L: e202花药横切图, e202中几乎没有饱满的花粉粒。A~D: 标尺为1 cm; E~H: 标尺为100 μm。"

图6

e202和ZH11叶片横切面 A~C: ZH11; D~F: e202。A, D: 剑叶中大维管束结构, 箭头表示薄壁组织细胞, 小三角型表示e202中缺失或减少的厚壁组织细胞层; B, E: 剑叶中小维管束结构, 箭头表示维管束; C, F: 泡状细胞, e202中泡状细胞较对照数目增多体积变小。"

图7

ZH11和e202叶片叶绿素含量测定 *表示与对照有显著性差异, P < 0.05; **表示与对照有极显著性差异, P < 0.01。"

表2

F2群体中正常植株和卷叶植株的分离比检测"

群体
Population		 总株数
Total plant number		 正常表型
Normal phenotype		 矮化表型
Dwarfing phenotype		 期望分离比
Segregation ratio		 P值
P-value		 χ2
(ZH11×e202) F2 560 433 127 3﹕1 0.21 1.731

图8

候选基因初步定位和精细定位图 A: 候选基因初步定位和精细定位遗传图谱; B: 候选基因结构图; C: REL2基因在ZH11和e202中的相对表达量。“n”表示定位分析所用到F2突变单株数。"

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