菊花近缘种属植物抗寒性评价及抗寒分子机制研究
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摘要
菊花(Chrysanthemum grandiflorum(Ramat.)Kitarn.),作为世界四大切花和我国十大传统名花之一,具有很高的观赏和经济价值,在花卉生产中占有重要地位。然而,我国长江中下游及长江以北地区的冬季低温严重影响了菊花生长发育,导致观赏品质下降,成为限制我国菊花产业发展的重要因素。开展菊花抗寒育种、培育抗寒新品种,是解决这一问题的根本途径。菊花近缘种属野生种质资源常具有栽培品种缺乏的某些抗逆性,对拓宽菊花种质资源和基因库有重要应用价值。因此,开展菊花近缘种属野生种质资源的抗寒筛选,获得典型抗寒材料,探索其抗寒生理及分子机制,有利于加快推动菊花抗寒种质创新和新品种选育进程。本文的研究内容及主要结论如下：
1.对45份菊花近缘种属野生资源的抗寒性进行了评价,结合冬季脚芽叶片半致死温度(LT50),将抗寒性进行分类,强抗寒(LT50<-20℃)：紫花野菊、沈阳野菊、异色菊、太行菊和北京植物园野菊等；中等抗寒(-20℃≤LT50≤-10℃)：黄金艾蒿、成都峰峡野菊、菊花脑等；不抗寒(LT50>-10℃)：若峡滨菊、神龙架野菊和达摩菊等。进一步通过脚芽恢复生长试验进行验证,发现脚芽冷冻处理后的存活率与LT50呈显著负相关,表明半致死温度可作为菊花近缘种属抗寒性评价的一个可靠指标。
2.以强抗寒异色菊和不抗寒若峡滨菊为材料,分析了冷驯化(4℃,1d、7d、14d和21d)条件下的抗寒差异。随着冷驯化时间的延长,两个菊属植物的半致死温度LT50呈现不同程度的下降；其中抗寒性强的异色菊从-7.3℃下降到-23.5℃,而抗寒性弱的若峡滨菊从-2.1℃下降到-7.1℃。测定了冷驯化过程中抗氧化酶(SOD、CAT和APX)活性和脯氨酸含量的变化,根据生物信息学从菊花EST数据库中电子克隆了抗氧化同工酶基因(Cu/Zn SOD、Fe SOD、Mn SOD、CAT和APX)、脯氨酸代谢相关基因(P5CS、OAT和PDH)以及冷调节相关基因(2个DREB家族基因、2个COR413家族基因和2个CSD家族基因),并进行了定量表达分析。结果发现,冷驯化使SOD、CAT和APX活性和脯氨酸含量的提高,其中异色菊提高的幅度显著高于若峡滨菊；基因表达分析显示SOD活性的提高主要来源于Mn SOD的表达增强,CAT和APX表达与其酶活性变化一致,P5CS和PDH表达与脯氨酸含量变化趋势一致；异色菊MnSOD、CAT、APX和P5CS表达的上升幅度显著高于若峡滨菊。冷调节相关基因表达分析结果显示,冷驯化诱导了DREBs、COR413s和CSDs基因表达的上调,其中异色菊的上升幅度均显著高于若峡滨菊。进一步分析发现,冷驯化初期抗寒性的提高来源于抗氧化系统的变化、脯氨酸含量的积累和冷调节基因的表达的综合作用,冷驯化后期抗寒性的提高来源于MnSOD, COR413s和CSDs等抗寒基因的表达的上调。
3. DREB家族基因在植物抗寒调控中发挥重要作用,为了进一步研究DREB基因的上游调控路径,从异色菊中分离到一个DREB家族基因CdDREBa(?)(?)启动子序列,其位于翻译起始位点上游1474bp的区域,PLACE数据库分析发现启动子中包括很多胁迫相关顺式作用元件(MYC-box, MYB-site, GT-1和W-box).通过启动子融合β-葡萄糖苷酸酶基因在拟南芥中表达分析,发现不同5’缺失的CdDREBa启动子在低温和盐胁迫下的活性存在差异。GUS组织化学染色和定量检测结果显示,启动子中一段80bp(-430～-351bp)的序列同时受低温和盐胁迫响应。将这个80bp的片段与35S mini启动子融合表达,显著促进了盐胁迫下GUS基因的表达。进一步通过内部缺失方法,发现一个低温抑制元件位于-430～-390bp区域和一个盐诱导元件位于-385～-351bp区域。结果显示,启动子中除了已知的MYC-box和GT-1元件受低温和盐胁迫响应外,我们发现的两个响应元件区域在胁迫响应过程中同样发挥着重要作用。
4.ICE家族基因在冷驯化调控植物抗寒路径中至关重要,以异色菊为材料,分离到一个ICE家族同源基因Cd1CE1,编码471个氨基酸,序列分析与其他物种ICEs具有高度相似性。通过荧光定量表达分析发现,低温、盐和ABA均诱导了Cd1CE1基因的表达；进一步克隆了Cd1CE1启动子,位于翻译起始位点上游1682bp的区域,序列分析发现了很多胁迫相关作用元件。构建35S::Cd1CE1-GFP表达载体,基因枪轰击洋葱表皮细胞进行瞬时表达,发现Cd1CE1定位于细胞核。采用毕赤酵母真核分泌表达系统,对Cd1CE1蛋白进行表达并纯化,凝胶阻滞(EMSA)试验证明Cd1CE1蛋白能与CdDREBa启动子中的MYC元件特异结合；酵母单杂交试验证明Cd1CE1具有转录激活活性。这些初步表明,异色菊中存在ICE1-DREB调节路径。
5.为了进一步证明Cd1CE16(?)抗寒调控功能,将Cd1CE1基因转到拟南芥中,并进行抗冻表型分析和分子机制探索。结果发现,不同温度驯化均提高了拟南芥抗冻性,异源表达异色菊Cd1CE1基因显著提高了不同温度驯化下抗冻性。23-4℃驯化下,Cd1CE1促进了CBFs及下游COR15a、COR6.6和RD29a三个抗冻基因的表达,与拟南芥At1CE1基因功能相似；而在23-16℃驯化条件下,Cd1CE1促进了三个抗冻基因的小幅上调,而对CBFs表达没有影响。23-16℃驯化诱导了miR398(?)(?)下调,促进了CSD1和CSD2的上调,进一步研究发现miR398负向调控拟南芥抗冻性,而CSDs正向调控抗冻性；miR398-CSDs路径参与了23-16℃驯化诱导的抗冻性调节。Cd1CE1过量表达显著下调了23-16℃驯化后miR398的表达；Cd1CE1介导了miR398-CSDs路径,提高抗冻性。因此,不同温度驯化下,Cd1CE1通过两条不同路径调控拟南芥抗冻性。
6.大量研究证明ICE1蛋白存在转录后的翻译修饰,为了深入研究Cd1CE1的调控机制,应用酵母双杂交体系对Cd1CE1互作蛋白进行了筛选。结果表明,采用Clontech试剂盒,成功构建了对照和冷驯化下菊花叶片cDNA混合酵母文库,以pGBKT7-Cd1CE1-85为诱饵质粒,筛选了64个候选阳性克隆并测序,在NCBI上进行BLAST比对分析,获得与光合作用、氧化胁迫和翻译修饰等相关的候选蛋白；其功能预测表明Cd1CE1可能参与了多条信号转导途径,在抗逆调控中具有重要作用。
Chrysanthemum(Chrysanthemum grandiflorum (Ramat.) Kitam.), as one of the four most popular cut flower over the world and one of the ten most famous flowers in China, provides very high ornamental and economic value, which possess the important status on flower industry. However, low temperature in winter in the North and Region of Yangtze in China seriously affects growth and development of Chrysanthemum and leads to lower ornamental characteristics, which has become a limitive factor on Chrysanthemum industry development. The research on cold-tolerance and new cultivar breeding is a fundamental approach to solve the problem. Chrysanthemum related genera carrying stress-resistent ability that is in absence in cultivar can play important worthiness for broadening germplasm and gene pool of Chrysanthemum. Therefor, cold-tolerance selection of Chrysanthemum related species is performed to achieve resistant germs based on the further analysis about their physiological and molecular mechanisms for cold-tolerance, which is benefit for pushing forward the germplasm enhancement and new cultivar breeding of Chrysanthemum. The contents and results on this research are as follows:
1. Evaluation for cold-tolerance of45Chrysanthemum related genera was analyzed based on the method of Semi-lethal temperature (LT50) of rhizomes leaf. Cold tolerance was classified, strong cold tolerance (LT50<-20℃):C.zawadskii, C.indicum (shenyang), C. dichrum, O. taihangensis and C.indicum (beijing); moderate cold tolerance (-20℃-10℃):C.makinoi, C.indicum (shennongjia), A.spathulifolius. Further confirmation via rhizomes recovery growth exhibited negative correlation between survival rate after freezing-treatment and LT50, which indicated that LT50could be a reliable index for cold-tolerance evaluation of Chrysanthemum related genera.
2. The difference of cold-tolerance between strong cold-tolerant C. dichrum and weak cold-tolerant C.makinoi under cold acclimation was analyzed. The results showed LT50presented different degree decrease under cold acclimation between the two Chrysanthemum species, C. dichrum varing from-7.3℃to-23.5℃and C.makinoi from-2.1℃to-7.1℃. Activities of antioxidant enzymes (SOD, CAT and APX) and proline content were measured, and antioxidant isoenzymes gene (Cu/Zn SOD、Fe SOD、Mn SOD、 CAT and APX), proline related-metabolism gene (P5CS, OAT and PDH) and cold-regulated gene (two DREB family genes, two COR413family gene and two CSD family gene) obtained from Chrysanthemum EST library were used for quantitive expression analysis. The results showed that cold acclimation promoted the activities increase of SOD, CAT and APX and proline accumulation, and C. dichrum presented higher varing degree than C.makinoi. Gene expression assay indicated that SOD activity promotion resulted from Mn SOD expression increase, CAT and APX expression exhibited consistent changes with their enzymes activities, P5CS and PDH expression showed similar alteration with proline content. Higher extent was detected on Mn SOD、CAT、APX and P5CS expression in C. dichrum comparing to in C.makinoi. Cold-regulated gene analysis indicated DREBs, COR413s and CSDs expression up-regulated under cold acclimation, and C. dichrum presented higher increase degree than C.makinoi. Further analysis revealed cold-tolerance improvement during the early days resulted from the functions of antioxidant enzymes, proline content and cold-regulated gene expression, and improvement during the late days origined from Mn SOD, COR413s and CSDs expression.
3. DREB family genes play important roles in regulating cold-tolerance. To investigate the upstream regulating passway of DREB gene, a1474bp stress-inducible CdDREBa promoter was identified from Chrysanthemum dichrum, revealing several candidate stress-related cis-acting elements (MYC-box, MYB-site, GT-1and W-box) within it. In Arabidopsis leaf tissues transformed with a CdDREBa promoter-β-glucuronidase (GUS) gene fusion, serially5'-deleted CdDREBa promoters were differentially activated by cold and salinity. Histochemical and quantitative assays of GUS expression allowed us to localize a critical part of the promoter located between upstream430and351bp. This80bp fragment enhanced GUS expression under salinity stress when fused to-90/+8CaMV35S minimal promoter. Further promoter internal-deletion assays indicated that a low temperature responsive element was located between positions-430and-390, and a salinity inducible one between-385and-351. Our results showed that there was a novel stress-related critical region except for the known cis-acting element (MYC-box, GT-1) in CdDREBa promoter.
4. ICE family genes are very important for regulating cold-tolerance passway. An ICE family homologue gene CdICE1was separated from Chrysanthemum dichrum, encoded471amino acids, which showed high similarity with ICE homologues from other species. Quantitive analysis revealed that CdICE1expression was induced under low temperature, salt and ABA treatment. CdICE1promoter within1682bp region upstream translation initiation site was identified, including some stress-responsve elements by sequence analysis. The assay that35S::CdICE1-GFP expression vector was bombarded into onion epidermal cells showed CdICE1was located in nucleus. CdICE1were expressed in Pichia pastoris system and purified, and then the specific binding of CdICE1to MYC element within the CdDREBa promoter was found through EMSA assay. Yeast one-hybrid assay presented that CdICE1had transcriptional activation activity. These data showed ICE1-DREB passway would exist in Chrysanthemum dichrum.
5. For identifying the cold-regulated functions of CdICE1, CdICE1was transformed to Arabidopsis and freezing-tolerance was detected by phenotype analysis, then molecular mechanism of freezing-tolerance was further explored. The results indicated that Arabidopsis freezing-tolerance was enhanced after different temperature acclimation, and heterologous expression CdICE1improved the freezing-tolerance under cold acclimation. Under4℃acclimation, CdICE1promoted expression of CBFs and their downstream freezing-tolerance gene including COR15a, COR6.6and RD29a, which revealed similar function with Arabidopsis AtICE1. However, under16℃acclimation, CdICE1only increased mildly three freezing-tolerant genes, while no changes on CBFs expression.16℃acclimation downregulated miR398and upregulated CSD1and CSD2expression, miR398negatively modulated freezing-tolerance and CSDs positively regulated freezing-tolerance, which indicated miR398-CSDs pathway played a role in16℃acclimation inducing freezing-tolerance. Overexpression CdICE1plants revealed significant reduction of miR398under16℃acclimation, comparing to WT plants. CdICE1mediated miR398-CSDs passway and then led to freezing-tolerance. Therefor, CdICE1affected freezing-tolerance via two different passway under two different cold acclimation conditions.
6. ICE1modification was found in previous research. To improve the research on CdICE1regulating mechanism, CdICE1-interacting proteins were screened by yeast two-hybrid system. A cDNA yeast mixture library was successfully constructed according to Clontech kit. A total of64candidate positive clones were sequenced and analyzed through homology analysis using the BLAST in NCBI, which showed that those candidate proteins were related to photosynthesis, oxidative stress and post-translational modification. Function prediction of the candidate proteins suggested that CdICE1was possibly involved in several stress signal transduction pathways and played an important role in regulation stress resistance.
1.对45份菊花近缘种属野生资源的抗寒性进行了评价,结合冬季脚芽叶片半致死温度(LT50),将抗寒性进行分类,强抗寒(LT50<-20℃)：紫花野菊、沈阳野菊、异色菊、太行菊和北京植物园野菊等；中等抗寒(-20℃≤LT50≤-10℃)：黄金艾蒿、成都峰峡野菊、菊花脑等；不抗寒(LT50>-10℃)：若峡滨菊、神龙架野菊和达摩菊等。进一步通过脚芽恢复生长试验进行验证,发现脚芽冷冻处理后的存活率与LT50呈显著负相关,表明半致死温度可作为菊花近缘种属抗寒性评价的一个可靠指标。
2.以强抗寒异色菊和不抗寒若峡滨菊为材料,分析了冷驯化(4℃,1d、7d、14d和21d)条件下的抗寒差异。随着冷驯化时间的延长,两个菊属植物的半致死温度LT50呈现不同程度的下降；其中抗寒性强的异色菊从-7.3℃下降到-23.5℃,而抗寒性弱的若峡滨菊从-2.1℃下降到-7.1℃。测定了冷驯化过程中抗氧化酶(SOD、CAT和APX)活性和脯氨酸含量的变化,根据生物信息学从菊花EST数据库中电子克隆了抗氧化同工酶基因(Cu/Zn SOD、Fe SOD、Mn SOD、CAT和APX)、脯氨酸代谢相关基因(P5CS、OAT和PDH)以及冷调节相关基因(2个DREB家族基因、2个COR413家族基因和2个CSD家族基因),并进行了定量表达分析。结果发现,冷驯化使SOD、CAT和APX活性和脯氨酸含量的提高,其中异色菊提高的幅度显著高于若峡滨菊；基因表达分析显示SOD活性的提高主要来源于Mn SOD的表达增强,CAT和APX表达与其酶活性变化一致,P5CS和PDH表达与脯氨酸含量变化趋势一致；异色菊MnSOD、CAT、APX和P5CS表达的上升幅度显著高于若峡滨菊。冷调节相关基因表达分析结果显示,冷驯化诱导了DREBs、COR413s和CSDs基因表达的上调,其中异色菊的上升幅度均显著高于若峡滨菊。进一步分析发现,冷驯化初期抗寒性的提高来源于抗氧化系统的变化、脯氨酸含量的积累和冷调节基因的表达的综合作用,冷驯化后期抗寒性的提高来源于MnSOD, COR413s和CSDs等抗寒基因的表达的上调。
3. DREB家族基因在植物抗寒调控中发挥重要作用,为了进一步研究DREB基因的上游调控路径,从异色菊中分离到一个DREB家族基因CdDREBa(?)(?)启动子序列,其位于翻译起始位点上游1474bp的区域,PLACE数据库分析发现启动子中包括很多胁迫相关顺式作用元件(MYC-box, MYB-site, GT-1和W-box).通过启动子融合β-葡萄糖苷酸酶基因在拟南芥中表达分析,发现不同5’缺失的CdDREBa启动子在低温和盐胁迫下的活性存在差异。GUS组织化学染色和定量检测结果显示,启动子中一段80bp(-430～-351bp)的序列同时受低温和盐胁迫响应。将这个80bp的片段与35S mini启动子融合表达,显著促进了盐胁迫下GUS基因的表达。进一步通过内部缺失方法,发现一个低温抑制元件位于-430～-390bp区域和一个盐诱导元件位于-385～-351bp区域。结果显示,启动子中除了已知的MYC-box和GT-1元件受低温和盐胁迫响应外,我们发现的两个响应元件区域在胁迫响应过程中同样发挥着重要作用。
4.ICE家族基因在冷驯化调控植物抗寒路径中至关重要,以异色菊为材料,分离到一个ICE家族同源基因Cd1CE1,编码471个氨基酸,序列分析与其他物种ICEs具有高度相似性。通过荧光定量表达分析发现,低温、盐和ABA均诱导了Cd1CE1基因的表达；进一步克隆了Cd1CE1启动子,位于翻译起始位点上游1682bp的区域,序列分析发现了很多胁迫相关作用元件。构建35S::Cd1CE1-GFP表达载体,基因枪轰击洋葱表皮细胞进行瞬时表达,发现Cd1CE1定位于细胞核。采用毕赤酵母真核分泌表达系统,对Cd1CE1蛋白进行表达并纯化,凝胶阻滞(EMSA)试验证明Cd1CE1蛋白能与CdDREBa启动子中的MYC元件特异结合；酵母单杂交试验证明Cd1CE1具有转录激活活性。这些初步表明,异色菊中存在ICE1-DREB调节路径。
5.为了进一步证明Cd1CE16(?)抗寒调控功能,将Cd1CE1基因转到拟南芥中,并进行抗冻表型分析和分子机制探索。结果发现,不同温度驯化均提高了拟南芥抗冻性,异源表达异色菊Cd1CE1基因显著提高了不同温度驯化下抗冻性。23-4℃驯化下,Cd1CE1促进了CBFs及下游COR15a、COR6.6和RD29a三个抗冻基因的表达,与拟南芥At1CE1基因功能相似；而在23-16℃驯化条件下,Cd1CE1促进了三个抗冻基因的小幅上调,而对CBFs表达没有影响。23-16℃驯化诱导了miR398(?)(?)下调,促进了CSD1和CSD2的上调,进一步研究发现miR398负向调控拟南芥抗冻性,而CSDs正向调控抗冻性；miR398-CSDs路径参与了23-16℃驯化诱导的抗冻性调节。Cd1CE1过量表达显著下调了23-16℃驯化后miR398的表达；Cd1CE1介导了miR398-CSDs路径,提高抗冻性。因此,不同温度驯化下,Cd1CE1通过两条不同路径调控拟南芥抗冻性。
6.大量研究证明ICE1蛋白存在转录后的翻译修饰,为了深入研究Cd1CE1的调控机制,应用酵母双杂交体系对Cd1CE1互作蛋白进行了筛选。结果表明,采用Clontech试剂盒,成功构建了对照和冷驯化下菊花叶片cDNA混合酵母文库,以pGBKT7-Cd1CE1-85为诱饵质粒,筛选了64个候选阳性克隆并测序,在NCBI上进行BLAST比对分析,获得与光合作用、氧化胁迫和翻译修饰等相关的候选蛋白；其功能预测表明Cd1CE1可能参与了多条信号转导途径,在抗逆调控中具有重要作用。
Chrysanthemum(Chrysanthemum grandiflorum (Ramat.) Kitam.), as one of the four most popular cut flower over the world and one of the ten most famous flowers in China, provides very high ornamental and economic value, which possess the important status on flower industry. However, low temperature in winter in the North and Region of Yangtze in China seriously affects growth and development of Chrysanthemum and leads to lower ornamental characteristics, which has become a limitive factor on Chrysanthemum industry development. The research on cold-tolerance and new cultivar breeding is a fundamental approach to solve the problem. Chrysanthemum related genera carrying stress-resistent ability that is in absence in cultivar can play important worthiness for broadening germplasm and gene pool of Chrysanthemum. Therefor, cold-tolerance selection of Chrysanthemum related species is performed to achieve resistant germs based on the further analysis about their physiological and molecular mechanisms for cold-tolerance, which is benefit for pushing forward the germplasm enhancement and new cultivar breeding of Chrysanthemum. The contents and results on this research are as follows:
1. Evaluation for cold-tolerance of45Chrysanthemum related genera was analyzed based on the method of Semi-lethal temperature (LT50) of rhizomes leaf. Cold tolerance was classified, strong cold tolerance (LT50<-20℃):C.zawadskii, C.indicum (shenyang), C. dichrum, O. taihangensis and C.indicum (beijing); moderate cold tolerance (-20℃
2. The difference of cold-tolerance between strong cold-tolerant C. dichrum and weak cold-tolerant C.makinoi under cold acclimation was analyzed. The results showed LT50presented different degree decrease under cold acclimation between the two Chrysanthemum species, C. dichrum varing from-7.3℃to-23.5℃and C.makinoi from-2.1℃to-7.1℃. Activities of antioxidant enzymes (SOD, CAT and APX) and proline content were measured, and antioxidant isoenzymes gene (Cu/Zn SOD、Fe SOD、Mn SOD、 CAT and APX), proline related-metabolism gene (P5CS, OAT and PDH) and cold-regulated gene (two DREB family genes, two COR413family gene and two CSD family gene) obtained from Chrysanthemum EST library were used for quantitive expression analysis. The results showed that cold acclimation promoted the activities increase of SOD, CAT and APX and proline accumulation, and C. dichrum presented higher varing degree than C.makinoi. Gene expression assay indicated that SOD activity promotion resulted from Mn SOD expression increase, CAT and APX expression exhibited consistent changes with their enzymes activities, P5CS and PDH expression showed similar alteration with proline content. Higher extent was detected on Mn SOD、CAT、APX and P5CS expression in C. dichrum comparing to in C.makinoi. Cold-regulated gene analysis indicated DREBs, COR413s and CSDs expression up-regulated under cold acclimation, and C. dichrum presented higher increase degree than C.makinoi. Further analysis revealed cold-tolerance improvement during the early days resulted from the functions of antioxidant enzymes, proline content and cold-regulated gene expression, and improvement during the late days origined from Mn SOD, COR413s and CSDs expression.
3. DREB family genes play important roles in regulating cold-tolerance. To investigate the upstream regulating passway of DREB gene, a1474bp stress-inducible CdDREBa promoter was identified from Chrysanthemum dichrum, revealing several candidate stress-related cis-acting elements (MYC-box, MYB-site, GT-1and W-box) within it. In Arabidopsis leaf tissues transformed with a CdDREBa promoter-β-glucuronidase (GUS) gene fusion, serially5'-deleted CdDREBa promoters were differentially activated by cold and salinity. Histochemical and quantitative assays of GUS expression allowed us to localize a critical part of the promoter located between upstream430and351bp. This80bp fragment enhanced GUS expression under salinity stress when fused to-90/+8CaMV35S minimal promoter. Further promoter internal-deletion assays indicated that a low temperature responsive element was located between positions-430and-390, and a salinity inducible one between-385and-351. Our results showed that there was a novel stress-related critical region except for the known cis-acting element (MYC-box, GT-1) in CdDREBa promoter.
4. ICE family genes are very important for regulating cold-tolerance passway. An ICE family homologue gene CdICE1was separated from Chrysanthemum dichrum, encoded471amino acids, which showed high similarity with ICE homologues from other species. Quantitive analysis revealed that CdICE1expression was induced under low temperature, salt and ABA treatment. CdICE1promoter within1682bp region upstream translation initiation site was identified, including some stress-responsve elements by sequence analysis. The assay that35S::CdICE1-GFP expression vector was bombarded into onion epidermal cells showed CdICE1was located in nucleus. CdICE1were expressed in Pichia pastoris system and purified, and then the specific binding of CdICE1to MYC element within the CdDREBa promoter was found through EMSA assay. Yeast one-hybrid assay presented that CdICE1had transcriptional activation activity. These data showed ICE1-DREB passway would exist in Chrysanthemum dichrum.
5. For identifying the cold-regulated functions of CdICE1, CdICE1was transformed to Arabidopsis and freezing-tolerance was detected by phenotype analysis, then molecular mechanism of freezing-tolerance was further explored. The results indicated that Arabidopsis freezing-tolerance was enhanced after different temperature acclimation, and heterologous expression CdICE1improved the freezing-tolerance under cold acclimation. Under4℃acclimation, CdICE1promoted expression of CBFs and their downstream freezing-tolerance gene including COR15a, COR6.6and RD29a, which revealed similar function with Arabidopsis AtICE1. However, under16℃acclimation, CdICE1only increased mildly three freezing-tolerant genes, while no changes on CBFs expression.16℃acclimation downregulated miR398and upregulated CSD1and CSD2expression, miR398negatively modulated freezing-tolerance and CSDs positively regulated freezing-tolerance, which indicated miR398-CSDs pathway played a role in16℃acclimation inducing freezing-tolerance. Overexpression CdICE1plants revealed significant reduction of miR398under16℃acclimation, comparing to WT plants. CdICE1mediated miR398-CSDs passway and then led to freezing-tolerance. Therefor, CdICE1affected freezing-tolerance via two different passway under two different cold acclimation conditions.
6. ICE1modification was found in previous research. To improve the research on CdICE1regulating mechanism, CdICE1-interacting proteins were screened by yeast two-hybrid system. A cDNA yeast mixture library was successfully constructed according to Clontech kit. A total of64candidate positive clones were sequenced and analyzed through homology analysis using the BLAST in NCBI, which showed that those candidate proteins were related to photosynthesis, oxidative stress and post-translational modification. Function prediction of the candidate proteins suggested that CdICE1was possibly involved in several stress signal transduction pathways and played an important role in regulation stress resistance.
引文
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管志勇,陈素梅,陈发棣,尹冬梅,刘兆磊,唐娟,杨帆(201 0)32个菊花近缘种属植物耐盐性筛选.中国农业科学43(19):4063-4071
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洪波,史春凤,张晓娇,高俊平(2009)菊花观赏性状和农艺性状基因工程改良研究进展.中国农业科学42(4)：1348-1358
胡春梅,侯喜林,王旻(2008)低温胁迫对不结球自菜光合及叶绿素荧光特性的影响.西北植物学报28(12):2478-2484
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李亚,谢晓金,宣继萍,刘建秀(2003)中国结缕草属(Zoysia spp.)植物抗寒性评价.草地学报11(03):240-245
李争艳,韩艳,张新芳,安黎哲,徐世健(2008)低温条件下高山离子芥SA的产生及NO对SA质量分数的影响.兰州大学学报(自然科学版)44(6):29-33
梁李宏,梅新,林锋,夏军,刘术金,王金辉(2009)低温胁迫对腰果幼苗叶片组织结构和生理指标的影响.生态环境学报18(1):317-320
刘冰浩,徐宁,朱建华,梁文,彭宏祥,黄江流(2006)广西龙眼种质耐寒性的CTR值鉴定.西南农业学报19(4):668-671
刘立军,陈为峰,张志华,田素娟(2010)叶面喷施脱落酸对高羊茅抗寒性的影响.中国草地学报32(6):94-99
刘鹏,赵世杰,孟庆伟,魏佑营,邹琦(2002)冷锻炼对甜椒叶片光合作用及其低温光抑制的影响(英文).植物生理与分子生物学学报28(1):51-58
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马向丽,魏小红,龙瑞军,崔文娟,万引琳(2005)外源一氧化氮提高一年生黑麦草抗冷性机制.生态学报25(6)：1269-1274
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唐立红(2010)不同品种紫斑牡丹叶片结构与抗寒性关系的初步研究.北方园艺23:95-97
田丹青,葛亚英,刘晓静,潘刚敏,沈晓岚,张智,郁永明(2011)叶面喷施水杨酸对红掌植株抗寒性的影响.浙江农业学报23(2):304—308
王静,孙磊,张成军,陈国祥,王萍,施大伟,吕川根,许所扣(2006)杂交稻幼苗期对低温胁迫的生理反应.作物学报32(7):1049-1056
王强,张其德(2003)光合作用光抑制的研究进展.植物学通报20(5):539～548
王易振,杨治国,曾祥琼,张廷惠(2010)低温处理对青蒿叶片脯氨酸、SOD和CAT的含量及蛋白质的影响.基因组学与应用生物学29(5):927-930
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韦弟,李杨瑞,邸南,闭志强,卜朝阳(2009)乙烯利对香蕉抗寒性的影响.热带作物学报30(12)：1789-1792
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许楠,孙广玉(2009)低温锻炼后桑树幼苗光合作用和抗氧化酶对冷胁迫的响应.应用生态学报20(4):761-766
许瑛,陈发棣(2008)菊花8个品种的低温半致死温度及其抗寒适应性.园艺学报35(4):559-564
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杨凤仙,董俊梅,杨晓霞(2001)低温胁迫下棉叶叶绿体、液胞超微结构的变化.山西农业大学学报2:116-117
杨华庚,林位夫(2009)低温胁迫对油棕幼苗光合作用及叶绿素荧光特性的影响.中国农学通报25(24):506-509
姚远,闵义,胡新文,李开绵,郭建春(2010)低温胁迫对木薯幼苗叶片转化酶及可溶性糖含量的影响.热带作物学报31(4):556-560
尹冬梅,管志勇,陈素梅,陈发棣(2009)菊花及其近缘种属植物耐涝评价体系建立及耐涝性鉴定.植物遗传资源学报10(3):399-340
于锡宏,蒋欣梅,刁艳,王丽丽,刘在民,于广建(2010)脱落酸、水杨酸和氯化钙对番茄幼苗抗冷性的影响.东北农业大学学报41(5):42～45
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张帆,颉建明,唐大为,郁继华,冯致,潘香梅(2011)亚精胺对辣椒幼苗抗冷性的影响.甘肃农业大学学报46(1):58-62
张楠,洪永聪,王玉,丁兆堂(2010)脱落酸和水杨酸对越冬期茶树叶片抗寒生理指标的影响.北方园艺2010(22):21-24
张有福,陈银萍,张满效,陈拓,安黎哲(2006)两种圆柏属植物不同季节显微和超微结构变化与耐寒性的关系.应用生态学报17(8):1393-1397
赵梅,周瑞莲,刘建芳,钟旭生(2011)冬季融冻过程中白三叶叶片抗氧化酶活力和渗透调节物含量变化与抗冻性的关系.生态学报31(2):306-315
郑国华,张贺英,钟秀容(2009)低温胁迫下枇杷叶片细胞超微结构及膜透性和保护酶活性的变化.中国生态农业学报17(4)：739-745
郑丽梅,司龙亭,韩贵超(2009)低温处理对不同耐寒性黄瓜幼苗叶片超微结构的影响.西北农业学报18(4):276-279
郑志勇,石进朝,王德芳(2009)长绿期金银木耐寒性与叶片组织结构的关系.华北农学报24(S1):331-333
周建,杨立峰,郝峰鸽,尤扬(2009)低温胁迫对广玉兰幼苗光合及叶绿素荧光特性的影响.西北植物学报29(1)：136-142
周明琦,吴丽华,沈忱,林娟(2010)ABA、MeJA和SA诱导下的荠菜CBF途径冷响应相关基因表达调控研究.中国农业科技导报12(6):75-80