不结球白菜BcHSP81-4基因在Po1CMS中的表达分析及BcFLC基因的功能验证
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摘要
细胞质雄性不育系是作物杂交育种的重要材料,也是研究花粉发育、细胞质遗传、核质互作以及时序性表达的理想材料。因此,对细胞质雄性不育分子机理的研究具有非常重要的意义。不结球白菜(Brassica campestris ssp. chienesis Makino)原产于中国,是我国南方普遍种植的大众化蔬菜,在蔬菜周年供应中起着举足轻重的作用。近年来,随着分子生物学技术的发展,对CMS的研究取得了较大的进展,但雄性不育的机理研究尚不清楚。故本研究以Pol胞质雄性不育系及其保持系为材料,从分子生物学方面入手,对不结球白菜Pol雄性不育系和保持系花期发育之间的差异表达基因进行了研究分析,有助于全面地了解高等植物中雄性不育发生的机理。同时,不结球白菜先期抽薹现象降低了产品的产量和质量,晚抽薹基因FLC的引入可避免该现象,但FLC对植物发育存在消极影响,因而,对抽薹和植物发育关系的研究具有重要意义。本文主要研究结果如下：
1.不结球白菜Pol胞质不育系花期抑制性消减文库的构建
以不结球白菜Pol胞质雄性不育系及其保持系为材料,采用抑制性消减杂交(supperssion subtraetive hybridization, SSH)技术,构建Pol胞质雄性不育系抑制消减cDNA文库。菌落PCR检测显示,有效重组率为91%,插入片段大小主要集中在100bp到1000bp之间。斑点杂交筛选得到32个阳性克隆,序列测定和同源性比对分析表明,15个克隆功能未知,其余涉及到激素信号转导、第二信使、光合作用、脂肪酸代谢及衰老等多个方面。
2.不结球白菜BcHSP81-4基因的克隆及其在PolCMS中的表达分析
从不结球白菜Pol胞质雄性不育系抑制消减cDNA文库中筛选出一条BcHSP81-4基因,经鉴定是热激蛋白家族的成员之一。从BcHSP81-4基因的氨基酸比对中,发现其与其他物种的HSP90家族具有很高的同源性。同时,本文还研究了BcHSP81-4在不同花发育时期和不同胁迫条件下的表达差异。结果表明,BcHSP81-4在35℃的热胁迫条件下,表达无显著差异,但是在盐胁迫和冷胁迫条件下,都能诱导表达。同时本文也发现BcHSP81-4基因在不育系的花蕾中高表达,因此,本研究推测BcHSP81-4基因在polCMS不育系中也可能起一定的作用。将BcHSP81-4基因片段连接到原核表达载体PGEX-4T-3,转化大肠杆菌BL21(DE3)后诱导重组蛋白的表达。经IPTG诱导后进行SDS-PAGE电泳,产生了预期大小的重组蛋白,进一步证明了该基因属于热激蛋白90家族。理化性质分析结果显示该蛋白含699个氨基酸,分子量为80.05KD,理论等电点pI值为4.95。TMpred跨膜分析结果显示,它含有1个显著的跨膜螺旋区,这个区域与ProtScale预测的疏水区基本对应。用TargetP server和WOLFPSORT server程序预测该蛋白为细胞质蛋白,二级结构以a-螺旋和随机卷曲为主。
3.不结球白菜晚抽薹基因BcFLC过表达影响拟南芥的育性和抗寒性
FLC在拟南芥中对开花是一个剂量依赖型抑制因子。本实验室从不结球白菜中克隆到一条FLC相关序列,命名为BcFLC,本研究将该基因在拟南芥中组成型过表达,结果发现,BcFLC可以显著延迟抽薹时间10天左右,表明BcFLC是抑制抽薹开花的一个基因。在BcFLC过表达植株中,花期的SOC1的表达量降低,但LFY和FT的表达量显著高于对照。因此本文认为BcFLC过表达植株比对照植株需要更多剂量的LFY和FT来促进开花。同时,本文还发现过表达BcFLC的拟南芥育性受到影响,主要是由于RGA和RGL等花丝抑制因子的表达量上升和SEP3等花和胚珠发育相关基因的下调造成的。在BcFLC过表达植株中,CBF1,2,3和COR15a等抗寒相关基因的表达量上调,植株抗寒性增强,表明BcFLC的过表达激活了CBF1,2,3和COR15a抗寒基因的表达,从而提高植物的抗寒性。
4.外源硝酸盐和NaCl胁迫对拟南芥开花的调节
植物开花受到内源和外源信号的双重调节。很多植物在一定的逆境胁迫下(如低硝酸盐和盐胁迫)都能提前开花,但有关这些胁迫是如何调节植物开花的机理尚不清楚。本研究发现低浓度的硝酸盐和NaCl胁迫能够增强GA生物合成酶GA1的活性,激活GA的生物合成。从而使得CO和SOC1的表达量增强,促进开花。但是硝酸盐和NaCl对开花光周期途径的影响是不同的。光周期相关基因CCA1和LHY在低浓度NaCl胁迫下受到抑制；但在低浓度硝酸盐胁迫下却没有显著变化。先前的研究表明,SOC1介导开花所有的内源途径：春化,自主开花,长日照和GA途径。因此,本文推测SOC1基因在介导植物内外源信号调节植物开花中起着重要作用。
Cytoplasmic male sterile (CMS) lines do not only play a major role in cross breeding of crops, but also they are taken as ideal materials for studying pollen development, cytoplasmic inheritance, nucleo-cytoplasmic interaction and time-dependent expression. Accordingly, studies on the molecular mechanism of cytoplamic male sterile are very significant. Non-heading Chinese cabbage(Brassica campestris ssp. chinensis Makino) which originated from China, is one of the most widely cultivated vegetables in China, especially in the south of China. In recent years, with the development of molecular biotechnology, studies about cytoplasmic male sterility increased gradually. However, there is little kownledge about gene recombination and its expression. The buds of newly bred Pol CMS of non-heading Chinese cabbage and its maintainer line as plant materials, genes differentially expressed transcripts were analyzed by molecular biology. Therefore, this study laid a good foundation on revealing the molecular mechanism of CMS line. Meanwhile, earlier bolting reduces the yield and quality of the products in non-heading chinese cabbage. The introduction of late bolting gene FLC could avoid this phenomenon. but FLC has a passive effect on plant fertility. Therefore, studying the relations of bolting and plant fertility has a great significance.The main results are as follows:
1-Suppression subtractive library was constructed using flowers of Pol cytoplasmic male sterile lines of Non-heading Chinese cabbage
Using suppression subtractive hybridization, the suppression subtractive cDNA library was builded, Pol cytoplasmic male sterile lines of Non-heading Chinese cabbage as the tester, and maintain as the driver. Colony were detected by PCR, the effective recombination rate is91%, and the size of inserts mainly between100bp and1000bp. Through dot blot screening,32positive clones were sequenced. Homology comparison analysis showed that15clones were unknown function, and the rest related to multiple aspects of the hormonal signal transduction, such as second messenger, photosynthesis, fatty acid metabolism and aging.
2. Cloning and expression analysis of BcHSP81-4gene in the PolCMS of Non-heading Chinese Cabbage
BcHSP81-4gene, a member of heat shock proteins, was identified from a suppression subtractive hybridization cDNA library in non-heading Chinese cabbage (Brassica campestris ssp. chinensis Makino). The deduced amino acid sequence of the BcHSP81-4cDNA revealed that it has high homology to other plant organelle isoforms and similar homology to both cytoplasmic and prokaryotic HSP90s. To study the regulation of gene expression, BcHSP81-4genes in maintainer and sterility lines were monitored at different development stages and at different stress treatments. Real-time PCR was used for quantification of BcHSP81-4mRNA. These results indicate that BcHSP81-4is not responsive to heat shock at least at35℃, while it is very responsive to salt and cold stress. And high expression of BcHSP81-4in the bud of sterile line suggests that it may play prominent roles in sterility of pol CMS in non-heading Chinese cabbage. Then the ORF had been cloned into PGEX-4T-3vector and transformed into the host BL21(DE3). Results of SDS-PAGE showed that the specific fusion protein was successfully induced to express by IPTG, and further proved that the gene belongs to heat shock protein90family。The heat shock protein BcHSP81-4gene of Brassica campestris ssp. Chinensis was analyzed by using a series of online software in the following aspects:physical and chemical characters, hydrophobicity, transmembrane topologieal structure, subcellular localization, and structure of the protein. The results indicated that the protein contains699amino acid residues. Its molecular weight is80.05KD, and the theoretical pI is4.95. TMpred analysis shows that it contained one distinguished transmembrane helice which correspond with the hydrophobic region that predicted by ProtScale. It is located in cytoplasm by Target P server. Structure analysis shows that a-helix and random coil are the main secondary structures of this protein.
3. Fertility and Cold-resistance were affected in Arabidopsis by the overexpression of BcFLC isolated from Non-heading Chinese cabbage
FLOWERING LOCUS C (FLC) in Arabidopsis encodes a dosage dependent repressor of flowering. In this paper, we isolated a FLC-related sequence from Non-heading Chinese cabbage (Brassica campestris ssp. chinensis Makino). Constitutive expression of the BcFLC genes in Arabidopsis significantly delayed flowering by10days, confirming the requirement of this gene for floral repression. In35S::BcFLC overexpression line, the level of SOC1mRNA expression has decreased, but LEAFY (LFY) and FLOWERING LOCUS T (FT) expression in35S::BcFLC were significantly higher than that in wild-type when flowering. The analysis revealed that the35S::BcFLC overexpression line in Arabidopsis requires a greater dose of LFY and FT to flower than the wild type. Here, we reported that the reduced fertility in Arabidopsis by the overexpression of BcFLC may be related with the enhanced expression of anther filaments suppressor, RGA and RGL gene, and slightly declined expression of SEPALLATA3(SEP3) gene, encoding a MADS-box transcription factor involved flower and ovule development. We also found that, CBF1,2,3and COR15a expression were gradually increased in35S::BcFLC overexpression line, suggesting that increased BcFLC expression could activate the expression of CBF1,2,3and COR15a, and then improving the cold resistance.
4. Exogenous Nitrate or NaCl regulates floral induction in Arabidopsis thaliana
Flowering, the transition from the vegetative to reproductive phase in plants, is regulated by both endogenous and environmental signals. Exposure to an extended period of stress (such as low nitrate or NaCl) can also promote flowering in many species, but little is known about how these forms of stress regulate floral induction. We found that stress induced by low concentrations of nitrate or NaCl activated the biosynthesis of gibberellin (GA) as evidenced by increased expression of the GA biosynthetic enzyme GA1. Expression of CO and SOC1were also enhanced, leading to an acceleration of flowering. The effects of nitrate and NaCl on the photoperiod pathway were distinct, however. Two genes related to the photoperiod pathway, CCA1and LHY, were repressed only under low NaCl treatment, while expression was unaltered by nitrate. Previous studies have shown that SOC1integrated all endogenous pathways:vernalization, autonomous, long-day, and GA pathways. We suggest that the SOC1MADS-box gene may play an important role in integrating signals induced by exogenous stress with endogenous signals to regulate flowering in Arabidopsis.
1.不结球白菜Pol胞质不育系花期抑制性消减文库的构建
以不结球白菜Pol胞质雄性不育系及其保持系为材料,采用抑制性消减杂交(supperssion subtraetive hybridization, SSH)技术,构建Pol胞质雄性不育系抑制消减cDNA文库。菌落PCR检测显示,有效重组率为91%,插入片段大小主要集中在100bp到1000bp之间。斑点杂交筛选得到32个阳性克隆,序列测定和同源性比对分析表明,15个克隆功能未知,其余涉及到激素信号转导、第二信使、光合作用、脂肪酸代谢及衰老等多个方面。
2.不结球白菜BcHSP81-4基因的克隆及其在PolCMS中的表达分析
从不结球白菜Pol胞质雄性不育系抑制消减cDNA文库中筛选出一条BcHSP81-4基因,经鉴定是热激蛋白家族的成员之一。从BcHSP81-4基因的氨基酸比对中,发现其与其他物种的HSP90家族具有很高的同源性。同时,本文还研究了BcHSP81-4在不同花发育时期和不同胁迫条件下的表达差异。结果表明,BcHSP81-4在35℃的热胁迫条件下,表达无显著差异,但是在盐胁迫和冷胁迫条件下,都能诱导表达。同时本文也发现BcHSP81-4基因在不育系的花蕾中高表达,因此,本研究推测BcHSP81-4基因在polCMS不育系中也可能起一定的作用。将BcHSP81-4基因片段连接到原核表达载体PGEX-4T-3,转化大肠杆菌BL21(DE3)后诱导重组蛋白的表达。经IPTG诱导后进行SDS-PAGE电泳,产生了预期大小的重组蛋白,进一步证明了该基因属于热激蛋白90家族。理化性质分析结果显示该蛋白含699个氨基酸,分子量为80.05KD,理论等电点pI值为4.95。TMpred跨膜分析结果显示,它含有1个显著的跨膜螺旋区,这个区域与ProtScale预测的疏水区基本对应。用TargetP server和WOLFPSORT server程序预测该蛋白为细胞质蛋白,二级结构以a-螺旋和随机卷曲为主。
3.不结球白菜晚抽薹基因BcFLC过表达影响拟南芥的育性和抗寒性
FLC在拟南芥中对开花是一个剂量依赖型抑制因子。本实验室从不结球白菜中克隆到一条FLC相关序列,命名为BcFLC,本研究将该基因在拟南芥中组成型过表达,结果发现,BcFLC可以显著延迟抽薹时间10天左右,表明BcFLC是抑制抽薹开花的一个基因。在BcFLC过表达植株中,花期的SOC1的表达量降低,但LFY和FT的表达量显著高于对照。因此本文认为BcFLC过表达植株比对照植株需要更多剂量的LFY和FT来促进开花。同时,本文还发现过表达BcFLC的拟南芥育性受到影响,主要是由于RGA和RGL等花丝抑制因子的表达量上升和SEP3等花和胚珠发育相关基因的下调造成的。在BcFLC过表达植株中,CBF1,2,3和COR15a等抗寒相关基因的表达量上调,植株抗寒性增强,表明BcFLC的过表达激活了CBF1,2,3和COR15a抗寒基因的表达,从而提高植物的抗寒性。
4.外源硝酸盐和NaCl胁迫对拟南芥开花的调节
植物开花受到内源和外源信号的双重调节。很多植物在一定的逆境胁迫下(如低硝酸盐和盐胁迫)都能提前开花,但有关这些胁迫是如何调节植物开花的机理尚不清楚。本研究发现低浓度的硝酸盐和NaCl胁迫能够增强GA生物合成酶GA1的活性,激活GA的生物合成。从而使得CO和SOC1的表达量增强,促进开花。但是硝酸盐和NaCl对开花光周期途径的影响是不同的。光周期相关基因CCA1和LHY在低浓度NaCl胁迫下受到抑制；但在低浓度硝酸盐胁迫下却没有显著变化。先前的研究表明,SOC1介导开花所有的内源途径：春化,自主开花,长日照和GA途径。因此,本文推测SOC1基因在介导植物内外源信号调节植物开花中起着重要作用。
Cytoplasmic male sterile (CMS) lines do not only play a major role in cross breeding of crops, but also they are taken as ideal materials for studying pollen development, cytoplasmic inheritance, nucleo-cytoplasmic interaction and time-dependent expression. Accordingly, studies on the molecular mechanism of cytoplamic male sterile are very significant. Non-heading Chinese cabbage(Brassica campestris ssp. chinensis Makino) which originated from China, is one of the most widely cultivated vegetables in China, especially in the south of China. In recent years, with the development of molecular biotechnology, studies about cytoplasmic male sterility increased gradually. However, there is little kownledge about gene recombination and its expression. The buds of newly bred Pol CMS of non-heading Chinese cabbage and its maintainer line as plant materials, genes differentially expressed transcripts were analyzed by molecular biology. Therefore, this study laid a good foundation on revealing the molecular mechanism of CMS line. Meanwhile, earlier bolting reduces the yield and quality of the products in non-heading chinese cabbage. The introduction of late bolting gene FLC could avoid this phenomenon. but FLC has a passive effect on plant fertility. Therefore, studying the relations of bolting and plant fertility has a great significance.The main results are as follows:
1-Suppression subtractive library was constructed using flowers of Pol cytoplasmic male sterile lines of Non-heading Chinese cabbage
Using suppression subtractive hybridization, the suppression subtractive cDNA library was builded, Pol cytoplasmic male sterile lines of Non-heading Chinese cabbage as the tester, and maintain as the driver. Colony were detected by PCR, the effective recombination rate is91%, and the size of inserts mainly between100bp and1000bp. Through dot blot screening,32positive clones were sequenced. Homology comparison analysis showed that15clones were unknown function, and the rest related to multiple aspects of the hormonal signal transduction, such as second messenger, photosynthesis, fatty acid metabolism and aging.
2. Cloning and expression analysis of BcHSP81-4gene in the PolCMS of Non-heading Chinese Cabbage
BcHSP81-4gene, a member of heat shock proteins, was identified from a suppression subtractive hybridization cDNA library in non-heading Chinese cabbage (Brassica campestris ssp. chinensis Makino). The deduced amino acid sequence of the BcHSP81-4cDNA revealed that it has high homology to other plant organelle isoforms and similar homology to both cytoplasmic and prokaryotic HSP90s. To study the regulation of gene expression, BcHSP81-4genes in maintainer and sterility lines were monitored at different development stages and at different stress treatments. Real-time PCR was used for quantification of BcHSP81-4mRNA. These results indicate that BcHSP81-4is not responsive to heat shock at least at35℃, while it is very responsive to salt and cold stress. And high expression of BcHSP81-4in the bud of sterile line suggests that it may play prominent roles in sterility of pol CMS in non-heading Chinese cabbage. Then the ORF had been cloned into PGEX-4T-3vector and transformed into the host BL21(DE3). Results of SDS-PAGE showed that the specific fusion protein was successfully induced to express by IPTG, and further proved that the gene belongs to heat shock protein90family。The heat shock protein BcHSP81-4gene of Brassica campestris ssp. Chinensis was analyzed by using a series of online software in the following aspects:physical and chemical characters, hydrophobicity, transmembrane topologieal structure, subcellular localization, and structure of the protein. The results indicated that the protein contains699amino acid residues. Its molecular weight is80.05KD, and the theoretical pI is4.95. TMpred analysis shows that it contained one distinguished transmembrane helice which correspond with the hydrophobic region that predicted by ProtScale. It is located in cytoplasm by Target P server. Structure analysis shows that a-helix and random coil are the main secondary structures of this protein.
3. Fertility and Cold-resistance were affected in Arabidopsis by the overexpression of BcFLC isolated from Non-heading Chinese cabbage
FLOWERING LOCUS C (FLC) in Arabidopsis encodes a dosage dependent repressor of flowering. In this paper, we isolated a FLC-related sequence from Non-heading Chinese cabbage (Brassica campestris ssp. chinensis Makino). Constitutive expression of the BcFLC genes in Arabidopsis significantly delayed flowering by10days, confirming the requirement of this gene for floral repression. In35S::BcFLC overexpression line, the level of SOC1mRNA expression has decreased, but LEAFY (LFY) and FLOWERING LOCUS T (FT) expression in35S::BcFLC were significantly higher than that in wild-type when flowering. The analysis revealed that the35S::BcFLC overexpression line in Arabidopsis requires a greater dose of LFY and FT to flower than the wild type. Here, we reported that the reduced fertility in Arabidopsis by the overexpression of BcFLC may be related with the enhanced expression of anther filaments suppressor, RGA and RGL gene, and slightly declined expression of SEPALLATA3(SEP3) gene, encoding a MADS-box transcription factor involved flower and ovule development. We also found that, CBF1,2,3and COR15a expression were gradually increased in35S::BcFLC overexpression line, suggesting that increased BcFLC expression could activate the expression of CBF1,2,3and COR15a, and then improving the cold resistance.
4. Exogenous Nitrate or NaCl regulates floral induction in Arabidopsis thaliana
Flowering, the transition from the vegetative to reproductive phase in plants, is regulated by both endogenous and environmental signals. Exposure to an extended period of stress (such as low nitrate or NaCl) can also promote flowering in many species, but little is known about how these forms of stress regulate floral induction. We found that stress induced by low concentrations of nitrate or NaCl activated the biosynthesis of gibberellin (GA) as evidenced by increased expression of the GA biosynthetic enzyme GA1. Expression of CO and SOC1were also enhanced, leading to an acceleration of flowering. The effects of nitrate and NaCl on the photoperiod pathway were distinct, however. Two genes related to the photoperiod pathway, CCA1and LHY, were repressed only under low NaCl treatment, while expression was unaltered by nitrate. Previous studies have shown that SOC1integrated all endogenous pathways:vernalization, autonomous, long-day, and GA pathways. We suggest that the SOC1MADS-box gene may play an important role in integrating signals induced by exogenous stress with endogenous signals to regulate flowering in Arabidopsis.
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