水稻颖壳异常突变体ah1的鉴定与候选基因分析

doi:10.3864/j.issn.0578-1752.2024.03.001

中国农业科学 ›› 2024, Vol. 57 ›› Issue (3): 429-441.doi: 10.3864/j.issn.0578-1752.2024.03.001

• 作物遗传育种·种质资源·分子遗传学 • 上一篇下一篇

水稻颖壳异常突变体ah1的鉴定与候选基因分析

张必东¹^,²(), 林泓¹, 朱思颖¹, 李忠成¹, 庄慧¹, 李云峰¹()

¹ 西南大学水稻研究所/西南大学农业科学研究院/转基因植物与安全控制重庆市重点实验室，重庆 400715
² 西南大学创新创业学院/含弘学院，重庆 400715

收稿日期:2023-07-05 接受日期:2023-08-28 出版日期:2024-02-01 发布日期:2024-02-05
通信作者:
李云峰，Tel：023-68251264；E-mail：liyf1980@swu.edu.cn
联系方式: 张必东，E-mail：vacom30@163.com。
基金资助:
国家自然科学基金(32172044); 国家自然科学基金(31971919); 重庆市杰出青年基金(cstc2020jcyj-jqX0020); 重庆市英才计划(cstc2021ycjh-bgzxm0066); 重庆市研究生科研创新项目(CYS21123); 重庆市级大学生创新创业训练计划(S202210635218)

Identification and Candidate Gene Analysis of the ABNORMAL HULL 1 (ah1) Mutant in Rice (Oryza sativa L.)

ZHANG BiDong¹^,²(), LIN Hong¹, ZHU SiYing¹, LI ZhongCheng¹, ZHUANG Hui¹, LI YunFeng¹()

¹ Rice Research Institute, Southwest University/Academy of Agricultural Sciences, Southwest University/Chongqing Key Laboratory of Application and Safety Control of Genetically Modified Crops, Chongqing 400715
² Institute of Innovation and Entrepreneurship, Southwest University/Hanhong College, Chongqing 400715

Received:2023-07-05 Accepted:2023-08-28 Published:2024-02-01 Online:2024-02-05

摘要/Abstract

摘要：

【目的】水稻是世界性的粮食作物，其籽粒形态直接影响水稻的最终产量和营养品质，进而影响其经济价值。此外，水稻花发育与籽粒形态又具有复杂的相关性，探究新的水稻花发育调控基因及其分子调控机制可以为培育籽粒更大、更饱满的水稻品种奠定基础。【方法】在利用甲基磺酸乙酯（ethyl methyl sulfonate，EMS）诱变西大1B（籼稻保持系）得到的突变体库中鉴定到一个颖壳和浆片发育异常且矮化的突变体abnormal hull 1（ah1）；观察并统计野生型和突变体的农艺性状；选取各个时期的小穗，对突变体的组织学、形态学变化进行分析；采用ah1和籼稻温敏不育系56S构建F₂分离群体，并将其用于遗传分析和基因定位；提取野生型和突变体的幼穗RNA并将其反转录为cDNA，对花发育调控基因和ABA合成关键基因的表达量进行RT-qPCR分析。【结果】农艺性状分析表明，突变体ah1各节间的大幅缩短直接导致其矮化；同时，突变体的小穗畸形严重，结实率极低。组织学和形态学分析发现，ah1小穗主要表现为内外稃、浆片和雄蕊等花器官发生了不同程度的退化，部分严重的小穗出现了花器官特征和花分生组织确定性的改变，且伴随大面积的白化，根据其退化程度的不同可分为一般和严重2种类型。遗传分析发现分离群体中野生型和突变体的比值为3﹕1，表明ah1突变性状受1个隐性基因控制。AH1定位于第1染色体上的分子标记RM6716和RM128之间，物理距离近8 Mb，对突变体进行重测序分析后发现该区间内的LOC_Os01g53450和LOC_Os01g51860在野生型和突变体之间出现了变异，因此，将这两个基因暂定为候选基因。RT-qPCR分析表明，在突变体幼穗早期发育过程中，各花器官发育调控基因的表达量发生了显著的变化；同时，ABA合成关键基因OsNCED1/OsNCED2/OsNCED3/OsNCED4/OsNCED5的表达严重受阻。【结论】AH1对于维持水稻内外稃等花器官的形态建成起到至关重要的作用，将LOC_Os01g53450和LOC_Os01g51860暂定为候选基因。

关键词: 水稻, 颖壳, 花发育, 候选基因

Abstract:

【Objective】Rice is the staple grain crop worldwide, and the morphology of its grains directly influences its ultimate yield, nutritional excellence, and economic significance. Moreover, the intricate interplay between floral development and grain morphology adds further significance to this relationship. Thus, exploring novel rice floral development regulatory genes and molecular regulatory mechanisms lays the foundation for larger and more plump grains rice varieties. 【Method】Ethyl methyl sulfonate (EMS) was used to mutate XD1B (xian-type maintainer line), and a dwarf mutant abnormal hull 1 (ah1) with abnormal formation of glume and lodicule was identified. The agronomic traits of both the wild-type and mutant were observed and recorded. Spikelets from various flowering stage were collected to histological and morphological analysis. The F₂ segregating population was established by ah1 and 56S (xian-type thermo-sensitive sterility line), and utlized for genetic analysis and gene mapping. RNA was isolated from young panicles of both the wild-type and mutant, then reverse transcribed into cDNA. The RT-qPCR analysis was performed to analyze the relative expression levels of the genes regulating floral development and the key genes in the ABA synthesis pathway. 【Result】The observation of agronomic traits revealed that the dwarfed plant was caused by the dramatic shortening of the internodes. At the same time, the mutant is also accompanied by severe spikelet abnormalities and low fruit setting rate. Histological and morphological analysis revealed that the ah1 mutant spikelets exhibited varying degrees of degeneration in floral organs such as palea, lemma, lodicules, and stamens. Some severely affected spikelets displayed altered floral organ characteristics and determinacy of floral meristems, often accompanied by extensive whitening. Based on the extent of degeneration, these spikelets could be classified as slight or severe mutant phenotypes. Genetic analysis showed a segregation ratio of 3﹕1 for the wild-type and mutant within the segregating population, indicating that the mutant traits of ah1 were controlled by a single recessive locus. The AH1 was mapped between the molecular markers RM6716 and RM128 on the chromosome 1, with a physical distance of approximately 8 Mb. Resequencing analysis of the mutant revealed that the LOC_Os01g53450 and LOC_Os01g51860 within this interval showed variation between wild-type and mutant, thus these two genes were provisionally identified as candidate genes. RT-qPCR analysis revealed significant alterations in the relative expression levels of floral organ development regulatory genes during the early developmental stages of mutant panicles; meanwhile, the relative expression levels of OsNCED1/OsNCED2/ OsNCED3/OsNCED4/OsNCED5, the ABA synthesis pathway key genes, were severe inhibited.【Conclusion】AH1 plays a crucial role in the morphological formation of floral organs, such as palea and lemma in rice. LOC_Os01g53450 and LOC_Os01g51860 were provisionally identified as candidate genes in this work.

Key words: rice (Oryza sativa L.), glume, floral development, candidate genes

张必东, 林泓, 朱思颖, 李忠成, 庄慧, 李云峰. 水稻颖壳异常突变体ah1的鉴定与候选基因分析[J]. 中国农业科学, 2024, 57(3): 429-441.

ZHANG BiDong, LIN Hong, ZHU SiYing, LI ZhongCheng, ZHUANG Hui, LI YunFeng. Identification and Candidate Gene Analysis of the ABNORMAL HULL 1 (ah1) Mutant in Rice (Oryza sativa L.)[J]. Scientia Agricultura Sinica, 2024, 57(3): 429-441.

图/表 7

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引物 Primer	正向序列 Forward sequence (5′-3′)	反向序列 Reverse sequence (5′-3′)	用途 Purpose
RM6716	CCTATGCGATCTCCTATTTGG	CGAGTCAGACACGTACTACTGC	定位 Mapping
RM128	TGATTTCTTGGAAGCGAAGAGTGAGG	CCTCCTTGTGCTCAGCCATGC
RM1268	GCTGTCACTGACCGAGCGTAGG	TCGAGAGATCCAATCCAGTTTGC
ACTIN	GACCCAGATCATGTTTGAGACCT	CAGTGTGGCTGACACCATCAC	RT-qPCR
OsMADS14	CTCATCATCCTCCTTCAT	GCTACATCCTCTATCCTT
OsMADS15	GCTGAATCCGAGAGTGAG	TGAGGTGTTTGTGACATTTG
OsMADS18	GGTGATGAATATGGAATT	CTTATATGCTTCAGAGAAT
OsMADS2	ATGATTGAGGAGGCACTT	TCGTCTTCCAGCATCTTAT
OsMADS4	CCAATCTGCGGGACAAGA	AGCCAAATTGGCAGTGCTC
OsMADS16	TTCGCCACAGGAAGTATCA	CCTCGTAGGAGTGCTTCA
OsMADS3	CGATCATATGGTTAATAACCCA	AGACGAGTTGCCAAAGTTCGA
OsMADS58	ATATCACCACCATGAACCA	TTCTTCCGTGCTCTAATCT
OsMADS13	AGAGCCAGGAAGAATGAA	ATGTTGTCGTTCTGAAGC
OsMADS1	GCTGCAACTACAACTCACAGG	TGATGGTGAGCATGAGGGTG
OsMADS5	ACTGAACTAAGCAATTACCAAGA	CAAGTCCTCGCCAAGAAG
OsMADS7	ATCTTGATTCATTAGGCATAA	TTGTCCTTGTAGTTCTGA
OsMADS8	GCACATTAGATCCACAAG	TTCTTCTGAGGCACTTAT
OsMADS34	AGGAATATGTGAACTTGAA	TTCTGACTACTTGACTCT
DL	CCCATCTGCTTACAACCGCTT	GTTGGAGGTGGAAACCGTCG
OsMADS6	TGATGATGGAACAAGTGGAG	TTGTGCTTGAGTTGCCTAT
OsMADS32	GGAGTTTGTCGAGAGGTACGA	CTGAACGTCAAGCGTCCATC
SL1	AAGCATCCAGGGTTACTAA	GCAGAACCTACACTCGTA
FON1	TGCGTGAAGGAGGACAACATCATC	AGCAGCAGGTTCGTCTCCCTGTT
OsNCED1	AAGCAGACCGAGCAAGAG	ATTCGTCGTAGCAATAACAGTTC
OsNCED2	AGCCAATGTTATGACTCT	TCTGTAGATTACGGTGAG
OsNCED3	AGGATATGCTCACATACAG	AGAATCTCACCGAATTGG
OsNCED4	ACGATGATCCACGACTTC	ATCTCCTGGAGCTTGAAC
OsNCED5	ATCAAGAAGCCATACCTGAA	CTGCTCAAGCTCGATCTC
LOC_Os01g51860	TGAAGGAATCTGCTGAAGT	AGTATCTGAGGACCACAATT
LOC_Os01g53450	ACGGAATGGCACAGTGGGAC	GGTCCCATGCCGTACTCTTGAG

序号 Number	基因名称 Gene name	注释 Annotation	位置 Position (Mb)
1	LOC_Os01g51860	紫黄素脱环氧化酶，推测，表达 Violaxanthin de-epoxidase, putative, expressed	28.44
2	LOC_Os01g53450	转氨酶，Ⅰ类和Ⅱ类，结构域蛋白，表达 Aminotransferase, classesⅠandⅡ, domain containing protein, expressed	29.29

水稻颖壳异常突变体ah1的鉴定与候选基因分析

Identification and Candidate Gene Analysis of the ABNORMAL HULL 1 (ah1) Mutant in Rice (Oryza sativa L.)

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