中国水稻科学 ›› 2021, Vol. 35 ›› Issue (5): 427-438.DOI: 10.16819/j.1001-7216.2021.210303
褚晓洁1, 芦涛1, 叶涵斐1, 王盛1, 林晗1, 吴先美2, 何瑞2, 严钢1, 王跃星2, 李三峰2, 路梅1, 胡海涛1,*(), 杨窑龙2,*(), 饶玉春1,*()
收稿日期:
2021-03-04
修回日期:
2021-04-12
出版日期:
2021-09-10
发布日期:
2021-09-10
通讯作者:
胡海涛,杨窑龙,饶玉春
基金资助:
Xiaojie CHU1, Tao LU1, Hanfei YE1, Sheng WANG1, Han LIN1, Xianmei WU2, Rui HE2, Gang YAN1, Yuexing WANG2, Sanfeng LI2, Mei LU1, Haitao HU1,*(), Yaolong YANG2,*(), Yuchun RAO1,*()
Received:
2021-03-04
Revised:
2021-04-12
Online:
2021-09-10
Published:
2021-09-10
Contact:
Haitao HU, Yaolong YANG, Yuchun RAO
摘要:
【目的】早衰突变体是研究早衰机制的良好载体,对于探究早衰的遗传机理与作用机制及提高水稻的产量和品质具有重要作用。【方法】本研究利用EMS诱变获得了一个早衰突变体lps1,并对该突变体及其野生型进行表型观察、细胞学及组织化学分析、生理生化分析、遗传分析、基因定位和激素处理。【结果】lps1的叶片从3叶期开始发黄,成熟期株高、有效分蘖数、结实率、千粒重等极显著降低。电镜观察发现lps1叶表面光滑,硅质化突起和叶绿体数目减少、片层结构紊乱。生理生化分析表明lps1中有大量的活性氧积累,同时伴有蛋白质的降解、细胞膜的损伤以及大规模的细胞死亡。遗传分析表明该早衰表型受单隐性核基因控制,并且其在第5染色体上编码了一个泛素结合酶。亚细胞定位结果证明LPS1蛋白在细胞质与核中均有表达。外源激素处理发现,lps1对外源激素的处理更为敏感,且LPS1突变促进了ABA合成相关基因的表达。【结论】LPS1 突变使水稻ABA合成信号途径异常,进而引发H2O2等一系列与衰老相关生理指标的异常变动,导致lps1过早衰老,最终造成水稻产量严重降低。
褚晓洁, 芦涛, 叶涵斐, 王盛, 林晗, 吴先美, 何瑞, 严钢, 王跃星, 李三峰, 路梅, 胡海涛, 杨窑龙, 饶玉春. 水稻叶片衰老基因LPS1的克隆与功能研究[J]. 中国水稻科学, 2021, 35(5): 427-438.
Xiaojie CHU, Tao LU, Hanfei YE, Sheng WANG, Han LIN, Xianmei WU, Rui HE, Gang YAN, Yuexing WANG, Sanfeng LI, Mei LU, Haitao HU, Yaolong YANG, Yuchun RAO. Cloning and Functional Analysis of Leaf Senescence Gene LPS1 in Oryza sativa[J]. Chinese Journal OF Rice Science, 2021, 35(5): 427-438.
引物名称 Primer name | 正向引物 Forward primer(5′-3′) | 反向引物 Reverse primer(5′-3′) | 用途 Use |
---|---|---|---|
M1 | CACACATTCTAAATTTGGAAAAAGG | TACGTGGCTACTTGGCGTTC | 精细定位Fine mapping |
M2 | GCTGAAGAGCCTCCTCGAA | GGTATCATGAGAGCGAGTCTGA | 精细定位Fine mapping |
M3 | ACCCATGACCATGAGACGAT | GGGTACTAGCCGTGCTTATCC | 精细定位Fine mapping |
M4 | AGCCATTAGGGGCTTAGGAA | CCCCTGAGTGATATGCTTGG | 精细定位Fine mapping |
M5 | TAAATTACATCGGCCGGAGA | CCCACCAAAGAAATCTCCAA | 精细定位Fine mapping |
M6 | ATTTGGGGGAAAGTTTGCTT | AATAGTATGCGTGCGCTGTG | 精细定位Fine mapping |
LPS1 | TCCTAAGAAGGGCTCGGAAA | CTGCCTCAACCACACTTACA | 载体构建Vector construction |
LPS1-GFP | GCCCAGATCAACTAGTATGGAT CTATATGCAATTGAC | TCGAGACGTCTCTAGACGGGCTGC AGGGGATGCCGG | 载体构建Vector construction |
Q-YGL1 | AACCTTACCGTCCTATTCCTT | CCATACATCTAACAGAGCACC | qRT-PCR |
Q-CAO1 | GATCCATACCCGATCGACAT | CGAGAGACATCCGGTAGAGC | qRT-PCR |
Q-NYC3 | TCTATCTAGGTGCCAAAGGC | ATTCTGGCACCTGCTGTTTC | qRT-PCR |
Q-DVR | CGAGCCCAGGTTCATCAAGGTGC | CCTCCCGATCTTGCCGAACTC | qRT-PCR |
Q-RLS1 | CTTGGGCTGTTGATGCAGC | CTTCAACACCCGCCTCGC | qRT-PCR |
Q-OsABA1 | GGATGCCATTGAGTTTGGTT | TGGCTGACTGAAGTCTCTCG | qRT-PCR |
Q-OsABA2 | AGCAAACCTGAAAGGTGTGGA | AAAGCCACCATCCACCATGA | qRT-PCR |
Q-OsABA3 | GGGCAAGATTTTGTTCGGCA | AAGGGTACACTTGTTGCCCC | qRT-PCR |
Q-OsNCED1 | ACCATGAAGTCCATGAGGCT | TCTCGTAGTCTTGGTCTTGG | qRT-PCR |
Q-OsNCED2 | ATGGAAACGAGGATAGTGGT | CTTATTGTTGTGCGAGAAGT | qRT-PCR |
Q-OsNCED3 | CTCCCAAACCATCCAAACCG | TGAGCATATCCTGGCGTCGT | qRT-PCR |
Q-OsNCED5 | TCCGAGCTCCTCGTCGTGAA | AGGTGTTTTGGAATGAACCA | qRT-PCR |
Q-OsZEP | GGATGCCATTGAGTTTGGTT | TGGCTGACTGAAGTCTCTCG | qRT-PCR |
Q-OsZDS | CACGTGTTCTTCGGGTGTTA | ATGTAACGGAGCTCCCACAG | qRT-PCR |
Q-OsABA80x1 | AAGCTGGCAAAACCAACATC | CCGTGCTAATACGGAATCCA | qRT-PCR |
Q-OsABA80x2 | CTACTGCTGATGGTGGCTGA | CCCATGGCCTTTGCTTTAT | qRT-PCR |
Q-OsABA80x3 | AGTACAGCCCATTCCCTGTG | ACGCCTAATCAAACCATTGC | qRT-PCR |
Actin | CAGGCCGTCCTCTCTCTGTA | AAGGATAGCATGGGGGAGAG | qRT-PCR |
表1 本研究所用引物及序列
Table 1 Primers and sequences used in this study.
引物名称 Primer name | 正向引物 Forward primer(5′-3′) | 反向引物 Reverse primer(5′-3′) | 用途 Use |
---|---|---|---|
M1 | CACACATTCTAAATTTGGAAAAAGG | TACGTGGCTACTTGGCGTTC | 精细定位Fine mapping |
M2 | GCTGAAGAGCCTCCTCGAA | GGTATCATGAGAGCGAGTCTGA | 精细定位Fine mapping |
M3 | ACCCATGACCATGAGACGAT | GGGTACTAGCCGTGCTTATCC | 精细定位Fine mapping |
M4 | AGCCATTAGGGGCTTAGGAA | CCCCTGAGTGATATGCTTGG | 精细定位Fine mapping |
M5 | TAAATTACATCGGCCGGAGA | CCCACCAAAGAAATCTCCAA | 精细定位Fine mapping |
M6 | ATTTGGGGGAAAGTTTGCTT | AATAGTATGCGTGCGCTGTG | 精细定位Fine mapping |
LPS1 | TCCTAAGAAGGGCTCGGAAA | CTGCCTCAACCACACTTACA | 载体构建Vector construction |
LPS1-GFP | GCCCAGATCAACTAGTATGGAT CTATATGCAATTGAC | TCGAGACGTCTCTAGACGGGCTGC AGGGGATGCCGG | 载体构建Vector construction |
Q-YGL1 | AACCTTACCGTCCTATTCCTT | CCATACATCTAACAGAGCACC | qRT-PCR |
Q-CAO1 | GATCCATACCCGATCGACAT | CGAGAGACATCCGGTAGAGC | qRT-PCR |
Q-NYC3 | TCTATCTAGGTGCCAAAGGC | ATTCTGGCACCTGCTGTTTC | qRT-PCR |
Q-DVR | CGAGCCCAGGTTCATCAAGGTGC | CCTCCCGATCTTGCCGAACTC | qRT-PCR |
Q-RLS1 | CTTGGGCTGTTGATGCAGC | CTTCAACACCCGCCTCGC | qRT-PCR |
Q-OsABA1 | GGATGCCATTGAGTTTGGTT | TGGCTGACTGAAGTCTCTCG | qRT-PCR |
Q-OsABA2 | AGCAAACCTGAAAGGTGTGGA | AAAGCCACCATCCACCATGA | qRT-PCR |
Q-OsABA3 | GGGCAAGATTTTGTTCGGCA | AAGGGTACACTTGTTGCCCC | qRT-PCR |
Q-OsNCED1 | ACCATGAAGTCCATGAGGCT | TCTCGTAGTCTTGGTCTTGG | qRT-PCR |
Q-OsNCED2 | ATGGAAACGAGGATAGTGGT | CTTATTGTTGTGCGAGAAGT | qRT-PCR |
Q-OsNCED3 | CTCCCAAACCATCCAAACCG | TGAGCATATCCTGGCGTCGT | qRT-PCR |
Q-OsNCED5 | TCCGAGCTCCTCGTCGTGAA | AGGTGTTTTGGAATGAACCA | qRT-PCR |
Q-OsZEP | GGATGCCATTGAGTTTGGTT | TGGCTGACTGAAGTCTCTCG | qRT-PCR |
Q-OsZDS | CACGTGTTCTTCGGGTGTTA | ATGTAACGGAGCTCCCACAG | qRT-PCR |
Q-OsABA80x1 | AAGCTGGCAAAACCAACATC | CCGTGCTAATACGGAATCCA | qRT-PCR |
Q-OsABA80x2 | CTACTGCTGATGGTGGCTGA | CCCATGGCCTTTGCTTTAT | qRT-PCR |
Q-OsABA80x3 | AGTACAGCCCATTCCCTGTG | ACGCCTAATCAAACCATTGC | qRT-PCR |
Actin | CAGGCCGTCCTCTCTCTGTA | AAGGATAGCATGGGGGAGAG | qRT-PCR |
图1 水稻早衰突变体lps1及其野生型的表型及农艺性状A-3叶期植株表型(标尺=3 cm);B-3叶期叶片表型(标尺=1 cm);C-分蘖期表型(标尺=6 cm);D-成熟期表型(标尺=8 cm);E-籽粒表型(标尺=10 cm); F~I-野生型与lps1的农艺性状(株高、分蘖、结实率、千粒重);J~L, 粒厚、粒长和粒宽。*和**分别表示野生型与lps1在0.05和0.01水平上差异显著。
Fig. 1. Phenotype and agronomic traits of the wild type and lps1. A, Plant phenotype(Bar=3 cm); B, Leaf phenotype at the three-leaf stage(Bar=1 cm); C, Tillering stage(Bar=6 cm); D, Maturity stage(Bar=8 cm); E, Seed phenotype of wild type and lps1(Bar=10 cm); F-I, Agronomic traits of wild type and lps1; J-L, Statistics of seed thickness, length and width of the wild type and lps1. * and ** indicate that the wild-type and mutant lps1 are significantly different at the levels of 0.05 and 0.01, respectively.
图2 水稻早衰突变体lps1及其野生型分蘖期叶绿素含量及相关基因表达分析A-分蘖期野生型与lps1不同部位叶表型,标尺为2 cm;B~D-野生型与lps1叶绿素a、叶绿素b和类胡萝卜素含量测定;E-叶绿素合成与代谢相关基因表达水平。L1―倒1叶;L2―倒2叶;L3―倒3叶。*和**分别表示野生型与lps1在0.05和0.01水平上差异显著
Fig. 2. Chlorophyll contents and related gene expression analysis at the tillering stage.A, Leaf phenotype of the wild type and lps1 in the tillering stage, bar=2 cm; B-D, Chlorophyll a, chlorophyll b and carotenoid contents of the wild type and lps1; E, Chlorophyll synthesis and metabolism-related gene expression level. L1, First leaf from the top; L2, Second leaf from the top; L3, Third leaf from the top. * and ** indicate that the wild type and lps1 are significantly different at the levels of 0.05 and 0.01, respectively.
图3 水稻野生型(A~C)和早衰突变体lps1(D~F)叶片下表皮的扫描电镜(SEM)观察
Fig. 3. Scanning electron microscope (SEM) observations of the epidermis of the wild type (A-C) and lps1(D-F).
图4 水稻野生型(A~C)和早衰突变体lps1(D~F)叶片的透射电镜(TEM)观察N-细胞核; C-叶绿体; Thy-类囊体; S-淀粉颗粒; Og-嗜锇颗粒。
Fig. 4. Transmission electron microscopy (TEM) observation of the wild type (A-C) and lps1(D-F) leaves. N, Nucleus; C, Chloroplast; Thy, Thylakoid; S, Starch granules; Og, Osmophilic granules.
图5 水稻早衰突变体lps1及其野生型生理生化检测A-DAB染色; 标尺为2 cm;B-NBT染色, 标尺为2 cm;C-衰老相关生理指标(过氧化氢、丙二醛、可溶性蛋白含量、过氧化氢酶、过氧化物酶、超氧化物歧化酶活性)的测定。*和**分别表示野生型与lps1在0.05和0.01水平上差异显著。
Fig. 5. Physiological and biochemical detection of the wild type and lps1.A, DAB staining; bar=2 cm; B, NBT staining, bar=2 cm; C, Determination of aging-related physiological indicators (H2O2, MDA, SP, CAT, POD, SOD). * and ** indicate that the wild-type and lps1 are significantly different at the levels of 0.05 and 0.01, respectively.
图6 野生型(A、B)和lps1(C、D)叶片的TUNEL处理 A-D,标尺为100 μm。蓝色荧光代表正常细胞,绿色荧光代表凋亡细胞。
Fig. 6. TUNEL treatment results of the wild type (A and B) and lps1 (C and D) leaves. A-D, bar=100 μm. Blue fluorescence represents normal cells, green fluorescence represents apoptotic cells.
杂交组合 Combination | F1表型 Phenotype of F1 | F2表型 Phenotype of F2 | χ2(3:1) | ||
---|---|---|---|---|---|
正常表型 Normal | 突变表型 Mutant | 总株数 Total | |||
lps1/浙辐802 lps1/Zhefu 802 | 正常表型 Normal | 253 | 82 | 335 | 0.049 |
浙辐802/lps1 Zhefu 802/lps1 | 正常表型 Normal | 233 | 76 | 309 | 0.027 |
表2 F2代分离群体统计结果
Table 2 Statistical results of F2 segregating population.
杂交组合 Combination | F1表型 Phenotype of F1 | F2表型 Phenotype of F2 | χ2(3:1) | ||
---|---|---|---|---|---|
正常表型 Normal | 突变表型 Mutant | 总株数 Total | |||
lps1/浙辐802 lps1/Zhefu 802 | 正常表型 Normal | 253 | 82 | 335 | 0.049 |
浙辐802/lps1 Zhefu 802/lps1 | 正常表型 Normal | 233 | 76 | 309 | 0.027 |
图9 外源激素处理对野生型和lps1幼苗的影响 A-外源激素处理对野生型WT(左)和lps1(右)表型的影响,标尺为3 cm;B-激素处理后地上部分长度;C-激素处理后地下部分长度。*和**分别表示野生型与lps1在0.05和0.01水平上差异显著。
Fig. 9. Effect of exogenous hormone treatment on the wild type and lps1 seedlings. A, Effect of exogenous hormone treatment on the phenotype of the wild type(left) and lps1(right), bar=3 cm; B, Length of the aerial part after hormone treatment; C, Length of the underground part after hormone treatment. * and ** indicate significant difference between the wild type and lps1 at 0.05 and 0.01 level, respectively.
图10 ABA合成与降解相关基因表达水平 A-处理前野生型(WT)和lps1中ABA相关基因表达水平;B-处理后野生型(WT)和lps1中ABA相关基因表达水平;*和**分别表示野生型与lps1在0.05和0.01水平上差异显著。
Fig. 10. ABA synthesis and degradation related gene expression levels. A, ABA-related gene expression levels in the wild type and lps1 before treatment; B, ABA-related gene expression levels in the wild type and lps1 after treatment; * and ** indicate significant difference between the wild type and lps1 at 0.05 and 0.01 level, respectively.
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