中国水稻科学 ›› 2022, Vol. 36 ›› Issue (2): 139-149.DOI: 10.16819/j.1001-7216.2022.210308
王永祥, 燕海刚, 徐含聪, 傅玉双, 单壮壮, 胡晓晴, 张文伟*(), 江玲
收稿日期:
2021-03-19
修回日期:
2021-05-12
出版日期:
2022-03-10
发布日期:
2022-03-11
通讯作者:
张文伟
基金资助:
WANG Yongxiang, YAN Hangang, XU Hancong, FU Yushuang, SHAN Zhuangzhuang, HU Xiaoqing, ZHANG Wenwei*(), JIANG Ling
Received:
2021-03-19
Revised:
2021-05-12
Online:
2022-03-10
Published:
2022-03-11
Contact:
ZHANG Wenwei
摘要:
【目的】 研究水稻EARLY STARVATION1 (OsESV1)基因对水稻淀粉代谢的影响。【方法】 通过CRISPR-Cas9技术获得osesv1突变体,考查osesv1的表型及胚乳淀粉的理化特性,分析OsESV1的表达特性及相关功能。【结果】 OsESV1蛋白在植物界中十分保守。osesv1突变体株高、穗长、每穗粒数低于野生型,分蘖数显著多于野生型,叶片中淀粉含量显著下降,籽粒中的直链淀粉含量上升,而总淀粉含量无明显变化。OsESV1呈组成型表达,并且具有昼夜节律表达的特征。OsESV1蛋白定位在叶绿体内且呈点状分布。酵母双杂交和双分子荧光互补实验结果表明OsESV1蛋白可以与ADP-葡萄糖焦磷酸化酶小亚基(OsAGPS) 2a和OsAGPS1互作。【结论】 OsESV1基因影响水稻叶片的淀粉合成途径,而对胚乳淀粉合成的影响不明显。
王永祥, 燕海刚, 徐含聪, 傅玉双, 单壮壮, 胡晓晴, 张文伟, 江玲. OsESV1基因对水稻淀粉合成的影响[J]. 中国水稻科学, 2022, 36(2): 139-149.
WANG Yongxiang, YAN Hangang, XU Hancong, FU Yushuang, SHAN Zhuangzhuang, HU Xiaoqing, ZHANG Wenwei, JIANG Ling. Effects of OsESV1 on Starch Synthesis in Rice[J]. Chinese Journal OF Rice Science, 2022, 36(2): 139-149.
引物 Primer | 正向引物序列 Forward primer sequence (5′-3′) | 反向引物序列 Reverse primer sequence (5′-3′) |
---|---|---|
Crispr-OsESV1 | GGCAAGCACCTGGTACAGGGAGAG | AAACCTCTCCCTGTACCAGGTGCT |
OsESV1-TI | AGCTCCTCAATCCCGTGATA | CTCATACGAAAATGTGGT |
PAN580-OsESV1-GFP | TCCGGAGCTAGCTCTAGAATGGCCGCGTGCTCCAGG | CGCCCTTGCTCACCATGGATCCTTCCAGAGGCGAAGGAGGCA |
1305-OsESV1-GFP | TCCGGAGCTAGCTCTAGAATGGCCGCGTGCTCCAGG | CGCCCTTGCTCACCATGGATCCTTCCAGAGGCGAAGGAGGCA |
QRT-OsESV1 | GTGATACTCCGCCGAAGAGA | TCGTCTCCACTCTCCCTGTA |
Actin | TGCTATGTACGTCGCCATCCAG | AATGAGTAACCACGCTCCGTCA |
AD-OsESV1 | CATGGAGGCCGAATTCATGGCCGCGTGCTCCAGG | CGAGCTCGATGGATCCTCATTCCAGAGGCGAAGGAGG |
BD-OsESV1 | CATGGAGGCCGAATTCATGGCCGCGTGCTCCAG | GCAGGTCGACGGATCCTCATTCCAGAGGCGAAGGAGG |
AD-OsAGPS1 | GGAGGCCAGTGAATTCATGGCGATGATGGCGATG | CGAGCTCGATGGATCCTTATATGACTGTTCCGCTAG |
BD-OsAGPS1 | CATGGAGGCCGAATTCATGGCGATGATGGCGATG | GCAGGTCGACGGATCCTTATATGACTGTTCCGCTAG |
AD-OsAGPS2a | GGAGGCCAGTGAATTCATGGCGATGGCGGCAGCCAT | CGAGCTCGATGGATCCTCATATAACTGTTCCGCTAG |
AD-OsAGPS2b | CATGGAGGCCGAATTCATGAATGTATTGGCATCTAAG | CGAGCTCGATGGATCCTCATATAACTGTTCCGCTAGG |
AD-OsAGPL1 | CATGGAGGCCGAATTCATGCAGTTCAGCAGTGTGTTT | CGAGCTCGATGGATCCCTATATGACCTTCCCGTCC |
AD-SSⅠ | CATGGAGGCCGAATTCATGGCGACGGCGGCGGGGAT | CGAGCTCGATGGATCCTTACATGACATATGGTTGATC |
AD-SSⅡa | CATGGAGGCCGAATTCATGTCGTCGGCCGTCGTCGCGTC | CGAGCTCGATGGATCCTCACCATTGGTACTTGGCCTT |
AD-SSⅡb | CATGGAGGCCGAATTCTTCACCTCCTCTTCGCCGCG | CGAGCTCGATGGATCCTCACCACTGGTACTTGGCCTT |
AD-BEI | GGAGGCCAGTGAATTCATGCTGTGTCTCACC | CGAGCTCGATGGATCCCTCATTTGCAGTCTTC |
AD-BEⅡb | CATGGAGGCCGAATTCATGGCGGCGCCGGCGTCTG | CGAGCTCGATGGATCCTCATTCCGCTGGAGCATA |
p2YC-OsESV1 | ATTTACGAACGATAGTTAATTAAATGGCGATGGCCGCGTGCTCCAGG | CACTGCCACCTCCTCCACTAGTTTCCAGAGGCGAAGGAGG |
p2YN-OsAGPS2a | ATTTACGAACGATAGTTAATTAAATGGCGATGGCGGCAGCCAT | CACTGCCACCTCCTCCACTAGTTATAACTGTTCCGCTAGGG |
p2YN-OsAGPS1 | ATTTACGAACGATAGTTAATTAAATGGCGATGGCGATGATGGCGATG | CACTGCCACCTCCTCCACTAGTTATGACTGTTCCGCTA |
表1 实验所用引物
Table 1 Primers used in this study.
引物 Primer | 正向引物序列 Forward primer sequence (5′-3′) | 反向引物序列 Reverse primer sequence (5′-3′) |
---|---|---|
Crispr-OsESV1 | GGCAAGCACCTGGTACAGGGAGAG | AAACCTCTCCCTGTACCAGGTGCT |
OsESV1-TI | AGCTCCTCAATCCCGTGATA | CTCATACGAAAATGTGGT |
PAN580-OsESV1-GFP | TCCGGAGCTAGCTCTAGAATGGCCGCGTGCTCCAGG | CGCCCTTGCTCACCATGGATCCTTCCAGAGGCGAAGGAGGCA |
1305-OsESV1-GFP | TCCGGAGCTAGCTCTAGAATGGCCGCGTGCTCCAGG | CGCCCTTGCTCACCATGGATCCTTCCAGAGGCGAAGGAGGCA |
QRT-OsESV1 | GTGATACTCCGCCGAAGAGA | TCGTCTCCACTCTCCCTGTA |
Actin | TGCTATGTACGTCGCCATCCAG | AATGAGTAACCACGCTCCGTCA |
AD-OsESV1 | CATGGAGGCCGAATTCATGGCCGCGTGCTCCAGG | CGAGCTCGATGGATCCTCATTCCAGAGGCGAAGGAGG |
BD-OsESV1 | CATGGAGGCCGAATTCATGGCCGCGTGCTCCAG | GCAGGTCGACGGATCCTCATTCCAGAGGCGAAGGAGG |
AD-OsAGPS1 | GGAGGCCAGTGAATTCATGGCGATGATGGCGATG | CGAGCTCGATGGATCCTTATATGACTGTTCCGCTAG |
BD-OsAGPS1 | CATGGAGGCCGAATTCATGGCGATGATGGCGATG | GCAGGTCGACGGATCCTTATATGACTGTTCCGCTAG |
AD-OsAGPS2a | GGAGGCCAGTGAATTCATGGCGATGGCGGCAGCCAT | CGAGCTCGATGGATCCTCATATAACTGTTCCGCTAG |
AD-OsAGPS2b | CATGGAGGCCGAATTCATGAATGTATTGGCATCTAAG | CGAGCTCGATGGATCCTCATATAACTGTTCCGCTAGG |
AD-OsAGPL1 | CATGGAGGCCGAATTCATGCAGTTCAGCAGTGTGTTT | CGAGCTCGATGGATCCCTATATGACCTTCCCGTCC |
AD-SSⅠ | CATGGAGGCCGAATTCATGGCGACGGCGGCGGGGAT | CGAGCTCGATGGATCCTTACATGACATATGGTTGATC |
AD-SSⅡa | CATGGAGGCCGAATTCATGTCGTCGGCCGTCGTCGCGTC | CGAGCTCGATGGATCCTCACCATTGGTACTTGGCCTT |
AD-SSⅡb | CATGGAGGCCGAATTCTTCACCTCCTCTTCGCCGCG | CGAGCTCGATGGATCCTCACCACTGGTACTTGGCCTT |
AD-BEI | GGAGGCCAGTGAATTCATGCTGTGTCTCACC | CGAGCTCGATGGATCCCTCATTTGCAGTCTTC |
AD-BEⅡb | CATGGAGGCCGAATTCATGGCGGCGCCGGCGTCTG | CGAGCTCGATGGATCCTCATTCCGCTGGAGCATA |
p2YC-OsESV1 | ATTTACGAACGATAGTTAATTAAATGGCGATGGCCGCGTGCTCCAGG | CACTGCCACCTCCTCCACTAGTTTCCAGAGGCGAAGGAGG |
p2YN-OsAGPS2a | ATTTACGAACGATAGTTAATTAAATGGCGATGGCGGCAGCCAT | CACTGCCACCTCCTCCACTAGTTATAACTGTTCCGCTAGGG |
p2YN-OsAGPS1 | ATTTACGAACGATAGTTAATTAAATGGCGATGGCGATGATGGCGATG | CACTGCCACCTCCTCCACTAGTTATGACTGTTCCGCTA |
图1 OsESV1蛋白的保守性分析 A—OsESV1及其同源蛋白的进化树分析,蛋白名采用相应的GenBank 序列登录号;B—多氨基酸序列比对,星号表示20个氨基酸的间隔,浅红色区域为富含色氨酸(W)区,保守色氨酸残基下用“:”标注,黄色区域为富含脯氨酸(P)区。
Fig. 1. The conservation analysis of OsESV1. A, Neighbor-joining tree of OsESV1 and its homologs. The proteins are named by GenBank protein accession numbers; B, Multiple sequence alignment of OsESV1 and its homologs. Asterisks represent the interval of twenty amino acids. The tryptophan (W)-rich region is marked by light red and those conserved tryptophan residues are indicated by colons. The proline (P)-rich region is marked by yellow.
图2 osesv1突变体的表型 A—OsESV1的基因结构。深灰和浅灰方块分别表示外显子区域和UTR区域,方块之间的黑色连线表示内含子,下面为野生型和osesv1靶位点基因序列比较,连续的点表示缺失的序列,插入的碱基加粗标示,红色标注为原间隔基序(protospacer adjacent motif , PAM) TGG;B—野生型和突变体的OsESV1蛋白结构。箭头标注富含色氨酸区(Trp-rich region)和富含脯氨酸区(Pro-rich region);C—野生型和osesv1植株,标尺为10 cm;D—野生型和osesv1幼苗在白天和夜晚结束时OsESV1基因的表达水平。数据表示为平均值±标准差(n=3)。*和**分别表示差异达0.05和0.01显著水平(t 检验)。下同。
Fig. 2. Phenotype of the osesv1 mutants. A, Structure of the OsESV1 gene. The dark gray and light gray boxes represent the exon and UTR regions, respectively. The black lines connecting boxes are introns. The corresponding sequences of the wild type (WT) and osesv1 were compared below and a succession of points indicates the deleted sequence in osesv1. The inserted base is shown in bold. TGG (PAM) is marked by red; B, The protein structure of WT and osesv1; Arrows indicate the tryptophan-rich region and proline-rich region; C, WT and osesv1 plants. Bar=10cm; D, Expression level of OsESV1 in WT and osesv1 seedlings at end of day(ED) and end of night(EN). Values are mean ± SD (n=3). * and **indicate significant difference at 0.05 and 0.01 level, respectively (t-test). The same below.
材料 Materials | 株高 Plant height /cm | 穗长 Length of panicle /cm | 分蘖数 Tiller number | 每穗粒数 Grain number per panicle | 一次枝梗数 Primary rachis branch number | 千粒重 1000-grain weight /g |
---|---|---|---|---|---|---|
野生型WT | 94.80±3.10 | 19.31±0.77 | 26.13±8.55 | 127.58±9.33 | 8.60±0.83 | 22.57±1.07 |
osesv1-1 | 90.96±4.44** | 17.82±0.52** | 33.43±10.45* | 112.11±15.42* | 8.44±0.96 | 21.50±0.56 |
osesv1-2 | 90.47±5.40** | 17.69±0.82** | 43.08±13.85** | 113.63±10.71** | 9.00±0.95 | 23.01±0.85 |
表2 野生型和osesv1的农艺性状
Table 2 Agronomic traits of the wild type (WT) and osesv1.
材料 Materials | 株高 Plant height /cm | 穗长 Length of panicle /cm | 分蘖数 Tiller number | 每穗粒数 Grain number per panicle | 一次枝梗数 Primary rachis branch number | 千粒重 1000-grain weight /g |
---|---|---|---|---|---|---|
野生型WT | 94.80±3.10 | 19.31±0.77 | 26.13±8.55 | 127.58±9.33 | 8.60±0.83 | 22.57±1.07 |
osesv1-1 | 90.96±4.44** | 17.82±0.52** | 33.43±10.45* | 112.11±15.42* | 8.44±0.96 | 21.50±0.56 |
osesv1-2 | 90.47±5.40** | 17.69±0.82** | 43.08±13.85** | 113.63±10.71** | 9.00±0.95 | 23.01±0.85 |
图3 野生型和osesv1的叶片淀粉水平 A—白天和夜晚结束时野生型(WT)和osesv1叶片的碘染观察; B—白天和夜晚结束时野生型(WT)和osesv1叶片的淀粉含量测定。
Fig. 3. Starch contents in leaves of WT and osesv1. A and B show iodine staining and starch contents of WT and osesv1 leaves at end of day and end of night, respectively.
图4 osesv1种子表型及淀粉理化性质分析 A—野生型(WT)和osesv1的成熟种子表型,标尺为5 mm;B, C—野生型(WT)和osesv1种子总淀粉含量、直链淀粉含量;D—野生型和osesv1胚乳淀粉的黏度曲线。
Fig. 4. Phenotype of osesv1 grains and physicochemical properties of endosperm starch. A, Phenotype of WT and osesv1 mature seeds. Bar=5 mm; B, Total starch contents of WT and osesv1 grains; C, Amylose content of WT and osesv1 grains; D, Viscosity profiles of endosperm starch of WT and osesv1.
图5 OsESV1的表达模式分析 A—OsESV1在野生型不同组织及胚乳发育不同时期的表达水平;B—OsESV1的昼夜节律表达。黑框和白框分别表示黑夜和白天的时间轴。
Fig. 5. Expression pattern of OsESV1. A, Expression level of OsESV1 in various tissues and endosperm at different developmental stages of WT; B, Diurnal expression pattern of OsESV1. The black and white boxes represent the timelines of night and day, respectively.
图6 OsESV1的亚细胞定位 A—OsESV1-GFP融合蛋白在水稻原生质体中的表达,pAN580-GFP空载体作为对照,标尺为20 μm;B—OsESV1-GFP融合蛋白在烟草表皮细胞中的表达,p1305-GFP空载体作为对照,标尺为20 μm。
Fig. 6. Subcellular localization of OsESV1. A, Transient expression of OsESV1-GFP fusion protein in rice protoplasts. The empty vector pAN580-GFP was used as control. Bars=20 μm; B, Transient expression of OsESV1-GFP fusion protein in tobacco epidermal cells. The empty vector p1305-GFP was used as control. Bars=20 μm.
图7 酵母双杂实验表明OsESV1可以与OsAGPS2a和OsAGPS1互作 二缺:SD-Leu/-Trp;四缺:SD-Leu/-Trp/-His/-Ade。
Fig. 7. Yeast two-hybrid assays show that OsESV1 can interact with OsAGPS2a and OsAGPS1. DDO(double dropout supplements), SD-Leu/-Trp; QDO(quadruple dropout supplements), SD-Leu/-Trp/ -His/-Ade.
图8 双分子荧光互补实验证明OsESV1可以与OsAGPS2a和OsAGPS1在烟草表皮细胞中互作
Fig. 8. BiFC assays show that OsESV1 can interact with OsAGPS2a and OsAGPS1 in tobacco epidermal cells. Bars = 10 μm.
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