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作物学报 ›› 2025, Vol. 51 ›› Issue (6): 1467-1479.doi: 10.3724/SP.J.1006.2025.42052

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

水稻内卷叶突变体acl3的鉴定及调控基因的功能分析

雷松翰(), 范骏扬, 车艳奕, 代永东, 郑雨萌, 田维江, 桑贤春(), 王晓雯()   

  1. 西南大学农学与生物科技学院, 重庆 400715
  • 收稿日期:2024-12-02 接受日期:2025-03-26 出版日期:2025-06-12 网络出版日期:2025-04-09
  • 通讯作者: *王晓雯, E-mail: xwwang78@126.com;桑贤春, E-mail: sangxianchun@163.com
  • 作者简介:E-mail: Leisonghan11@126.com
  • 基金资助:
    本研究由重庆市现代农业产业技术体系项目(CQMAITS202401);国家重点研发计划项目(2022YFD1201600);国家级大学生创新创业训练计划项目(202410635004)

Identification of an adaxially-curled-leaf mutant acl3 and function analysis of the regulated gene in rice (Oryza sativa L.)

LEI Song-Han(), FAN Jun-Yang, CHE Yan-Yi, DAI Yong-Dong, ZHENG Yu-Meng, TIAN Wei-Jiang, SANG Xian-Chun(), WANG Xiao-Wen()   

  1. College of Agronomy and Biotechnology, Southwest University, Chongqing 400715, China
  • Received:2024-12-02 Accepted:2025-03-26 Published:2025-06-12 Published online:2025-04-09
  • Contact: *E-mail: xwwang78@126.com;E-mail: sangxianchun@163.com
  • Supported by:
    the Chongqing Modern Agricultural Industry Technology System(CQMAITS202401);the National Key Research and Development Program of China(2022YFD1201600);the College Students’ Innovation and Entrepreneurship Training Scientific Research Project(202410635004)

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摘要：

适度卷叶有利于异交结实和提高作物群体光合效率, 在杂交水稻制种及生产中具有重要的应用价值。我们从甲基磺酸乙酯EMS诱变体库中鉴定到1个稳定遗传的内卷叶突变体acl3 (adaxially curled leaf 3), 该突变体表现出株高和结实率降低、千粒重极显著提高。石蜡切片表明, 叶片上表皮维管束间的泡状细胞数量减少和面积变小是导致acl3极度内卷的主要原因。遗传分析表明, acl3的突变性状受1对单隐性核基因调控, 初步定位在第7染色体Indel标记07g89和07g498之间409 kb的物理范围内。对突变体acl3及其野生型西大1B进行全基因组测序, 利用IGV软件查找acl3定位区间的序列差异, 发现注释基因LOC_Os07g01240/SRL1/CLD1在第6外显子上存在1个G到A的碱基替换, 初步确定为候选基因。通过构建互补载体转化突变体acl3, 表型鉴定结果显示互补植株的卷叶、株高、籽粒大小等突变性状均恢复至野生型水平, 表明ACL3是SRL1的一个新等位基因, 并具有“一因多效”的特性。qRT-PCR分析结果显示, 卷叶相关基因REL2和NAL7极显著上调; 进一步分析生长素(IAA)途径相关基因的表达变化发现, 生长素合成基因YUCCA2、应答基因ARF1和ARF7、原初响应基因IAA10、IAA21、IAA22等均有不同程度的上调, 这表明ACL3可能通过生长素途径调控水稻株型的发育过程。

关键词: 水稻(Oryza sativa L.), 泡状细胞, 卷叶, SRL1, 生长素

Abstract:

Moderate leaf curling is beneficial for cross-pollination, fruit set, and enhancing the photosynthetic efficiency of crop canopies, making it highly valuable for hybrid rice seed production and cultivation. To elucidate the molecular mechanisms underlying leaf rolling in rice, we identified a stably inherited adaxial leaf-rolling mutant, acl3 (adaxially curled leaf 3), from an ethyl methanesulfonate (EMS)-mutagenized population. In addition to leaf curling, acl3 exhibits reduced plant height and seed-setting rate, along with a highly significant increase in 1000-grain weight. Histological analysis using paraffin sections revealed that the extreme curling in acl3 primarily results from a reduction in both the number and area of bulliform cells between vascular bundles in the adaxial epidermis. Genetic analysis indicated that the acl3 mutant phenotype is controlled by a single recessive nuclear gene, which was preliminarily mapped to a 409 kb physical region between Indel markers 07g89 and 07g498 on chromosome 7. Whole-genome sequencing of acl3 and the wild-type Xida 1B was performed, and sequence variations within the acl3 mapping interval were identified using IGV software. A G-to-A base substitution was detected in the sixth exon of the annotated gene LOC_Os07g01240/SRL1/CLD1, leading to a structural change in the encoded protein. This gene was preliminarily identified as the candidate gene for ACL3. Complementary vector construction and transformation of the acl3 mutant restored the mutant traits—leaf curling, plant height, and grain size to wild-type levels, confirming that ACL3 is a new allele of SRL1 with pleiotropic effects. Quantitative real-time PCR (qRT-PCR) analysis revealed highly significant upregulation of leaf-rolling-related genes REL2 and NAL7. Further investigation of auxin IAA pathway-related genes showed that auxin synthesis gene YUCCA2, auxin response genes ARF1 and ARF7, and primary auxin response genes IAA10, IAA21, and IAA22 were all upregulated to varying degrees, suggesting that ACL3 may regulate rice plant architecture development through the auxin signaling pathway.

Key words: rice (Oryza sativa L.), bulliform cells, curled leaf, SRL1, auxin

表1

基因定位及qRT-PCR所用引物序列"

引物名称
Primer name		 正向引物
Forward primer (5′-3′)		 反向引物
Reverse primer (5′-3′)
07g20 GCCACTTATCCAAATGCCCTT GCTGGGCCAAAGGAGAGAGG
07g89 GCTGCTACTCGATGATGATGATC TCTTCTTCCAGGACCAGCTTGC
07g498 CGGATTGAAACCCATCCACC ATGTTGGGTGGACGTGGATG
07g798 TGATGTCCGAAGTCAAGGTGC GCTTAACCGCAAACGTGGATT
ACTIN GACCCAGATCATGTTTGAGACCT CAGTGTGGCTGACACCATCAC
ADL1 AATGGAATGGTCCGTGGTCAG CACCTGCACTATAACCACGCCA
ACL1 TCTAGGGGCATCGACCTGAA GCATGACGTAGATGAAGCAGCGA
ROC5 AGGTGGAAATGTGAAGTGTAGG AGGCGAGACCAGACCGAAG
RL14 CTATCCACCGTGCTTGAA AGAACCATTTGCCATCCT
REL2 CTTGCTATCGTCGCTGTCGT GCTCTTCCCTTCCATTGACT
NAL7 CAGGAGGACGGTGAGTTGTTC GCAGTAGAGCCCGTTTGGAC
SLL1 CGGCGACATGGACCAGTCGT GCTCGATCTGCACGGTTCCAA
AVB CACAGTCTTCTCCGGGCAGCAA CATTACCACCACCTAAGCCAGCTCT

图1

野生型(WT)和突变体(acl3)的表型鉴定 A~C: 三叶一心期、拔节期和成熟期野生型(WT)和突变体acl3的植株表型; A中标尺为5 cm, B和C中标尺为10 cm。D: 成熟期野生型和突变体acl3倒一叶表型, 标尺为1 cm。E: 成熟期野生型和突变体acl3叶片卷曲指数(LRI)。F: 成熟期野生型和突变体acl3的穗及节间表型, 左边为野生型, 右边为突变体acl3, 标尺为5 cm。G: 成熟期野生型和突变体acl3的穗及节间统计。*、**分别表示在0.05、0.01水平差异显著。"

图2

野生型(WT)和突变体(acl3)产量性状统计分析 A: 野生型和突变体acl3的穗, 红色箭头指的为未结实的空壳, 标尺为3.5 cm; B: 野生型和突变体acl3的一次枝梗和二次枝梗数量统计; C: 野生型和突变体acl3的花结构, ①和②为相应的花粉粒KI-I2染色, 标尺为2 mm; D: 野生型和突变体acl3的籽粒, 标尺为6 mm; E: 野生型和突变体acl3的结实率统计; F: 野生型和突变体acl3的千粒重统计; G: 野生型和突变体acl3的籽粒长度统计; H: 野生型和突变体acl3的籽粒宽度统计; I: 野生型和突变体acl3的籽粒厚度统计; J: 野生型和突变体acl3的有效穗数量统计。**表示在0.01水平差异显著。"

表2

野生型与突变体acl3的主要农艺性状比较"

性状Trait 野生型Wild type 突变体Mutant
株高 Plant height (cm) 89.00±1.41 71.00±1.22**
穗长 Panicle length (cm) 25.80±0.45 24.00±1.00*
有效穗数 Panicles per plant 5.80±0.84 6.80±0.45
结实率 Seed-setting rate (%) 92.71±2.40 40.33±7.11**
粒长 Grain length (mm) 9.10±0.10 8.78±0.08**
粒宽 Grain width (mm) 2.76±0.05 2.82±0.04
粒厚 Grain thickness (mm) 1.92±0.04 2.13±0.04**
千粒重 1000-grain weight (g) 23.25±0.28 3.13±0.62**

图3

野生型(WT)和突变体(acl3)叶片细胞学观察和光合效率测定 A和B: 抽穗期野生型和突变体acl3倒一叶横切面石蜡切片, 标尺为500 μm; a和b: A和B图中红色线框的放大图, 标尺为100 μm; C: 两小维管束之间的泡状细胞数量统计; D: 泡状细胞面积统计; E: 净光合速率统计; F: 气孔导度统计; G: 蒸腾速率统计; H: 胞间二氧化碳浓度统计。BC: 泡状细胞。*、**分别表示在0.05、0.01水平差异显著。"

表3

卷叶突变体acl3的遗传分析"

组合名称
Combination		 F2群体株数
Total number		 野生型株数
Wild type number		 突变株数
Mutant number		 理论值
Theory value		 χ2 0.05=3.84
acl3/缙恢10 acl3/Jinhui 10 636 463 173 3:1 1.64

图4

ACL3基因的图位克隆 A: ACL3基因的分子定位; B: 定位区间差异位点的IGV图谱, 红色箭头表示碱基差异位点; C: 野生型WT和突变体acl3突变位点的测序验证, 红色方框表示突变碱基; D: 候选基因的结构图, 白色方块表示5′UTR, 黑色方块表示外显子, 白色箭头表示3′UTR, 黑实线表示内含子, 红色折线表示突变位点; E: WT、acl3和互补转基因植株(COM)表型, 标尺为20 cm; F: WT、acl3和COM剑叶叶片表型, 标尺为1 cm。"

图5

ACL3蛋白结构及生物信息学分析 A: 野生型和突变体acl3中的ACL3蛋白结构, 箭头所指为变异区; B: ACL3系统进化树分析; C: ACL3保守结构域(红色方框)和氨基酸突变位点(红色星号)。"

图6

卷叶及IAA相关基因的qRT-PCR分析 A: 卷叶相关基因的qRT-PCR分析; B: 生长素途径相关基因的qRT-PCR分析。*、**分别表示在0.05、0.01水平差异显著。"

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