中国水稻科学 ›› 2022, Vol. 36 ›› Issue (2): 150-158.DOI: 10.16819/j.1001-7216.2022.210606
裘霖琳1, 刘窍1, 付亚萍2, 刘文真2, 胡国成2, 翟玉凤1, 庞礴1, 汪得凯1,*()
收稿日期:
2021-06-17
修回日期:
2021-09-14
出版日期:
2022-03-10
发布日期:
2022-03-11
通讯作者:
汪得凯
基金资助:
QIU Linlin1, LIU Qiao1, FU Yaping2, LIU Wenzhen2, HU Guocheng2, ZHAI Yufeng1, PANG Bo1, WANG Dekai1,*()
Received:
2021-06-17
Revised:
2021-09-14
Online:
2022-03-10
Published:
2022-03-11
Contact:
WANG Dekai
摘要:
【目的】 水稻株高和穗型在产量形成中发挥着重要作用。鉴定与克隆水稻株高和穗型发育相关基因,可以丰富水稻株高穗型发育调控的分子机理,为分子设计育种奠定理论基础和提供基因资源。【方法】 在粳稻日本晴T-DNA插入群体中筛选到矮化小穗突变体dsp2-D(dwarf and small panicle 2-Dominant),对其主要农艺性状进行了分析;采用图位克隆法结合T-DNA标签分离法进行了基因定位和克隆;利用半定量PCR和qRT-PCR进一步确定dsp2-D的候选基因;遗传转化实验验证了DSP2的功能。【结果】 与野生型日本晴相比,dsp2-D突变体表现为半矮化、穗轴和枝梗明显缩短、穗型直立、千粒重降低等特征;遗传分析表明该突变体受一对不完全显性单基因控制;利用图位克隆将DSP2定位于第2染色体标记RM208和RM7337之间,与RM3850共分离;随后遗传分析发现,T-DNA插入位点与dsp2-D表型共分离,利用TAIL-PCR分离T-DNA插入序列,显示T-DNA插入到上述RM208和RM7337之间的两个基因之间。RT-PCR检测发现,位于T-DNA插入位点下游的一个编码LOB家族转录因子基因的表达量明显增加,而其他5个基因的表达量变化不明显,表明该基因可能为DSP2的候选基因;在野生型日本晴中过量表达DSP2基因,转化植株出现半矮化、小穗的表型,与dsp2-D表型类似,从而验证了DSP2的功能。【结论】 DSP2基因的过量表达是产生dsp2-D突变表型的原因;DSP2基因对水稻株高和穗长发育具有负向调控作用;为进一步丰富株高和穗型的遗传调控网络打下了基础。
裘霖琳, 刘窍, 付亚萍, 刘文真, 胡国成, 翟玉凤, 庞礴, 汪得凯. 水稻矮化小穗基因DSP2的鉴定与克隆[J]. 中国水稻科学, 2022, 36(2): 150-158.
QIU Linlin, LIU Qiao, FU Yaping, LIU Wenzhen, HU Guocheng, ZHAI Yufeng, PANG Bo, WANG Dekai. Identification and Gene Cloning of DSP2 in Rice (Oryza sativa L.)[J]. Chinese Journal OF Rice Science, 2022, 36(2): 150-158.
引物名称 | 正向引物序列 | 反向引物序列 | 实验目的 |
---|---|---|---|
Primer name | Forward primer(5'-3') | Reverse primer(5'-3') | Purpose |
ORF1-qPCR | AGATCAACCTCCAACTACTCTC | AAGCTCAGTAGTACACAGGC | RT-PCR |
ORF2-qPCR | GTGGTTCCAGAAGATCGTGTC | TTGTCGATGGACAGGAGCTC | RT-PCR |
ORF3-qPCR | CACGCCTACGACAACATGAAC | GTACTCGAACGCGTTGACATC | RT-PCR |
ORF4-qPCR | CCGGTGATACAAAGAGGTGC | ATATGCTTCGGCCACCTTG | RT-PCR |
ORF5-qPCR | AGCGGTTGTCTGGTGATATC | AGACAGAAAACCCCTTGACG | RT-PCR |
ORF6-qPCR | CAGCTTCTGATCCTGCAGTAG | ATCTCCCTTACTGATGCTGAC | RT-PCR |
UBQ5-qPCR | ACCACTTCGACCGCCACTACT | ACGCCTAAGCCTGCTGGTT | RT-PCR |
HPTⅡ | CAGAAGAAGATGTTGGCGAC | ATGTCCTGCGGGTAAATAGC | 共分离分析 Co-segregation |
DSP2-ORF | ATGTCGACATGGCTGGTGCTACGGCTGC | ATCTGCAGCTGAGAGTAGTTGGAGGTTGAT | 全长ORF Full length ORF |
NPTⅡ | TATGTCCTGATAGCGGTCCG | GTGCCCTGAATGAACTCCAG | 转化植株检测 Detection of transgenic plants |
表1 PCR鉴定及定量PCR所用的引物
Table 1 Primers used for PCR identification and quantitative real-time PCR (qRT-PCR) analysis.
引物名称 | 正向引物序列 | 反向引物序列 | 实验目的 |
---|---|---|---|
Primer name | Forward primer(5'-3') | Reverse primer(5'-3') | Purpose |
ORF1-qPCR | AGATCAACCTCCAACTACTCTC | AAGCTCAGTAGTACACAGGC | RT-PCR |
ORF2-qPCR | GTGGTTCCAGAAGATCGTGTC | TTGTCGATGGACAGGAGCTC | RT-PCR |
ORF3-qPCR | CACGCCTACGACAACATGAAC | GTACTCGAACGCGTTGACATC | RT-PCR |
ORF4-qPCR | CCGGTGATACAAAGAGGTGC | ATATGCTTCGGCCACCTTG | RT-PCR |
ORF5-qPCR | AGCGGTTGTCTGGTGATATC | AGACAGAAAACCCCTTGACG | RT-PCR |
ORF6-qPCR | CAGCTTCTGATCCTGCAGTAG | ATCTCCCTTACTGATGCTGAC | RT-PCR |
UBQ5-qPCR | ACCACTTCGACCGCCACTACT | ACGCCTAAGCCTGCTGGTT | RT-PCR |
HPTⅡ | CAGAAGAAGATGTTGGCGAC | ATGTCCTGCGGGTAAATAGC | 共分离分析 Co-segregation |
DSP2-ORF | ATGTCGACATGGCTGGTGCTACGGCTGC | ATCTGCAGCTGAGAGTAGTTGGAGGTTGAT | 全长ORF Full length ORF |
NPTⅡ | TATGTCCTGATAGCGGTCCG | GTGCCCTGAATGAACTCCAG | 转化植株检测 Detection of transgenic plants |
图1 突变体dsp2-D的表型 AA、Aa、aa分别代表野生型日本晴植株、杂合体和dsp2-D纯合体。不同字母代表差异极显著(P<0.01,Tukey检验),n=10。
Fig. 1. Phenotypes of the dsp2-D mutant. AA, Wild-type ‘Nipponbare’ plants; Aa, dsp2-D heterozygous plants; aa, dsp2-D homozygous plants. Values are mean ± SD of ten biological replicates. Samples with different letters are significantly different (P < 0.01; Tukey’s test).
图2 DSP2基因在第2染色体上的定位和T-DNA共分离分析 A―DSP2基因的初步定位;B―T-DNA插入位点共分离分析。M, 1 kb ladder; P为质粒阳性对照; A1~A3为野生型;B1~B7为杂合体;C1~C4为纯合体。
Fig. 2. Location of DSP2 on chromosome 2 and co-segregation analysis of T-DNA. A, Preliminary localization of DSP2 gene; B, Co-segregation analysis of T-DNA and dsp2-D phenotypes. M, 1 kb ladder, P, Plasmid positive control; A1-A3, Wild type; B1-B7, Heterozygote; C1-C4, Homozygous mutant.
图3 候选基因DSP2的表达分析 A―T-DNA插入位点;B, C―6个基因在突变体dsp2-D和野生型中的表达水平,水稻OsUBQ5为内参基因。Bar值表示3个生物学重复的平均值±SD;**表示0.01极显著水平。
Fig. 3. Expression analysis of candidate gene DSP2. A, Schematic diagram of T-DNA insertion sites; B and C, Expression levels of six candidate genes in dsp2-D and wild type analyzed by semi-quantitative RT-PCR and real time PCR; OsUBQ5 was amplified as a control. Bars represent standard deviation (n=3). ** indicates significant difference between Nipponbare and dsp2-D by t-test (P<0.01).
图4 DSP2基因的过量表达和功能验证 A―DSP2基因在野生型和转基因植株中的表达水平,其中WT代表野生型日本晴,CK代表空载体转化植株,OE-1#~OE-3#代表DSP2-OE转化植株;Bar值表示3个生物学重复的平均值±SD;**表示0.01极显著水平。B, C―DSP2过表达转基因植株的表型,CK为空载体转化植株。B图中标尺表示10 cm; C图中标尺表示5 cm。
Fig. 4. Overexpression of DSP2 gene and functional identification. A, Expression levels of DSP2 gene in wild-type and transgenic plants. WT, Wild type Nipponbare; CK, Control plasmid transgenic plants; OE-1#-OE-3#, DSP2-OE transgenic plants; Bars represent standard deviation (n=3). **indicates significant difference between Nipponbare and dsp2-D by t-test (P<0.01). B and C, Phenotype of the dsp2-D mutant and transgenic plant. B, Bar=10 cm; C, Bar=5 cm.
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