桃花芽休眠解除SSH文库构建及相关基因的功能分析
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摘要
本实验于2008年8月～2010年12月在山东农业大学果树试验站、园艺科学与工程学院中心实验室进行。分别以十年生大田‘曙光油桃’和3年生盆栽‘曙光油桃’花芽为试材分离芽休眠解除的相关基因，研究其分子功能，并研究相关基因在低温、短日照、激素处理条件下对休眠解除的影响。对于揭示芽自然休眠机理，人为调控花期，具有重要的理论意义和实践价值。主要结果如下：
利用抑制性差减杂交（SSH）技术，以休眠解除芽的RNA为‘Tester’，休眠芽的RNA为‘Driver’，构建花芽休眠解除相关基因的差减cDNA文库，共测定分析了180个上调表达的cDNA片段，测序成功128个，除去冗余序列，筛选到28个与自然休眠解除相关的差异基因，根据blast分析结果，进行分子功能预测，结果发现：以防御功能蛋白及新陈代谢类占的比例最大，分别达到25%和17.86%；其次是氧化还原与信号转导类，分别达到14.29%和7.14%，另外结构功能活性基因、功能未知蛋白、没有匹配蛋白、转运活性蛋白基因等占35.71%。
根据候选基因的功能推测和前人关于植物休眠调控的研究结果，选取了8个可能与自然休眠解除相关的基因PpMt，PpDhn，PpHis，PpPod，PpSenescence-associatedprotein，PpCyp450，PpATP-binding cassette transporter protein和unknown，采用qRT-PCR技术分析发现：它们在花芽自然休眠解除期间上调表达，认为曙光油桃花芽自然休眠解除的过程伴随着复杂物质代谢和能量消耗，差异基因主要参与植物生长发育及抗性胁迫等方面的调控，而且低温诱导的核糖体蛋白的表达可能也与花芽自然休眠解除有关。
PpDfn、PpDhn是文库中筛选到的全长cDNA序列，PpDfn、PpDhn基因编码的蛋白质氨基酸序列与其他物种的防御素蛋白（DFN、DHN）的氨基酸序列同源性较高。对二者进行蛋白质信号肽、跨膜结构域及疏水性分析，结果显示：PpDFN蛋白有1个信号肽，为分泌性蛋白。存在1个跨膜区，属于疏水性蛋白。通过对其氨基酸序列的进化树分析，发现与葡萄、油橄榄亲缘关系最近。PpDHN蛋白没有信号肽，无跨膜结构域，属于亲水性蛋白。通过对其氨基酸序列的进化树分析，发现与杏亲缘关系最近，二者聚在一起。采用qRT-PCR技术进一步分析了PpDFN、PpDHN在自然休眠解除期间、低温及短日照处理条件下的表达模式变化，发现PpDfn、PpDhn随自然休眠解除、低温处理时间的延长，基因表达量逐渐增加，冷处理2周时达到最高值；短日照处理后则没有明显变化。推测PpDfn基因随休眠解除及低温处理的上调表达，可能是参与了乙烯信号途径而诱导表达的结果。PpDhn基因随休眠解除及低温处理的上调表达可能与抗冻、清除自由基、保护生物膜免受伤害有关。
在油桃休眠及休眠解除期间，曙光油桃PIP1;1的转录水平呈现持续增高趋势，且1月15日的高水平表达使水分通过液泡膜和细胞质膜流出，减少了芽体水分含量，阻止细胞内冰晶的形成，从而抵御冻害；可溶性糖、可溶性蛋白、脯氨酸含量均达到最高，防止细胞的脱水伤害。低温处理2周后高水平表达说明PIP1;1为冷诱导基因。δTIP1的转录水平在休眠期间呈现波动性变化，至休眠解除时大幅度增高，这可能与休眠解除时，其上调表达被休眠解除信号及植物活性的增强所诱导有关。低温处理2周后，其表达没有升高，说明δTIP1并非冷诱导基因。
在曙光油桃休眠及休眠解除期间，深休眠期(11月15日)和休眠后期(12月15日)，喷布GA3和6-BA研究激素对解除休眠的效果。结果发现：在休眠后期施用GA3和6-BA能不同程度的提高H2O2含量和POD、SOD基因的表达量，降低CAT基因的表达水平。对萌芽率的统计表明，深休眠期施用GA3和6-BA对油桃花芽破眠无效，处理15天后没有花芽萌发。休眠后期施用GA3和6-BA破眠效果较为明显，萌芽较对照提前8d和3d，最终萌芽率达98%、91%，分别比对照提高了37%和30%。GA3的破眠效果好于6-BA。
The experiment was carried out in the horticulture experimental station of ShandongAgricultural University and central laboratory of College of Horticulture Science andEngineering during2008.8~2010.12. In the study, floral buds of ten-year-old field culturedand three-year-old potted nectarine (Prunus persica var. nectariana cv. Shuguang) trees wereused to identify and characterize regulatory genes of dormancy release. Expression changes ofthe genes in the dormancy release process under low temperature, short day and hormonetreatments were also detected. The study could be supplementary of the bud dormancymechanism and have important theoretical and practical value in the dormancy control ofprotected production of fruit trees. The main results were as follow:
The cDNA library of genes associated with bud dormancy release was established usingsuppression subtractive hybridization (SSH) procedure. The RNA of dormancy-released budswas used as Tester and the RNA from dormant buds was used as Driver.180up-regulatedpositive cDNA clones were detected and128of them were successfully sequenced.28uniquegenes related to dormancy release were screened after the elimination of redundant sequences.The results of BLAST analysis and molecular function prediction showed that genes involvedin defense and metabolism pathway made up lager percentages, accounting for25%and17.86%respectively. Genes associated with oxidation reduction and signaling accounted for14.29%and7.14%respectively. And genes related to activation of structure or function,unknown protein, un-match protein and transport activator protein accounted for35.71%.
According to functional predict of candidate genes and research progress of plantdormancy,8doemacy related genes, PpMt, PpDhn, PpHis, PpPod, PpSenescence-associatedprotein, PpCyp450, PpATP-binding cassette transporter protein and unknown, wereanalysised with qRT-PCR. The results showed that expression patterns of the genes wereup-regulated during dormancy release. Complex metabolism and energy consumption wereassociated with dormancy release, and the genes may be mainly involved in growth,development and tolerance to stress. The expression of ribosomal proteins induced by lowtemperature may be related to dormancy release of floral buds.
PpDfn、PpDhn were full-length cDNA sequence screened from the library. Amino acidsequences of PpDFN、PpDHN were highly homologous with defensins (DFN, DHN) of other species. Analysis of signal peptide, trans-membrane domain and hydrophobicity of the twogenes suggested that PpDFN was a secretory and hydrophobic protein containning a signalpeptide and a trans-membrane domain. PpDHN was hydrophilic protein without signalpeptide and trans-membrane domain. Cladogram analysis of amino acid sequences showedthat PpDFN of ‘Shuguang’ nectarine had closer genetic relationship with those of grape andolive and PpDHN had closer genetic relationship with that of Armeniaca. Expression analysisof PpDFN and PpDHN showed that expression of the two genes were up-regulated along withchilling accumulation during dormancy release, and peaked at the2thweek of chillingtreatment. Short day showed no obvious effect on the expression of the two genes. PpDFNmight be involved in ethylene signaling pathway and PpDHN might related tofreeze-resistance, eliminating free radicals and protecting bio-membrane from injury.
At stage of dormancy and dormancy release, transcripts of PIP1;1increased consistently.The high expression on15thJanuary led to outflow of water through vacuolar membrane andcytoplasm membrane, which caused water content decrease in buds and prevented frozeninjury. Contents of soluble sugars, proteins and proline peaked at the same time, whichprevented cell from dehydration injury. The high level expression after2weeks of lowtemperature treatment suggested that PIP1;1was a cold-inducible gene. Transcription ofδTIP1fluctuated during dormancy but increased significantly at stage of dormancy release.The increase might be related to dormancy release signal and plant activity enhancement.Expression of δTIP1did not increase after2weeks of low temperature treatment, indicatingthat δTIP1was not a cold-inducible gene.
GA3and6-BA were used at15thNovember (endo-dormancy) and15thDecember (latestage of endo-dormancy) to study effects of hormone on dormancy release. Application ofGA3and6-BA at late stage of dormancy increased H2O2level and expression of POD andSOD, but decrease the expression of CAT. Application of GA3and6-BA duringendo-dormancy did not increase budburst of dormant buds. However, at the late stage ofendo-dormancy, budburst time was brought forward by8and3days respectively by GA3and6-BA treatment, budburst rate were also increased by37%and30%respectively. Effect ofGA3on dormancy release was more significant than6-BA.
利用抑制性差减杂交（SSH）技术，以休眠解除芽的RNA为‘Tester’，休眠芽的RNA为‘Driver’，构建花芽休眠解除相关基因的差减cDNA文库，共测定分析了180个上调表达的cDNA片段，测序成功128个，除去冗余序列，筛选到28个与自然休眠解除相关的差异基因，根据blast分析结果，进行分子功能预测，结果发现：以防御功能蛋白及新陈代谢类占的比例最大，分别达到25%和17.86%；其次是氧化还原与信号转导类，分别达到14.29%和7.14%，另外结构功能活性基因、功能未知蛋白、没有匹配蛋白、转运活性蛋白基因等占35.71%。
根据候选基因的功能推测和前人关于植物休眠调控的研究结果，选取了8个可能与自然休眠解除相关的基因PpMt，PpDhn，PpHis，PpPod，PpSenescence-associatedprotein，PpCyp450，PpATP-binding cassette transporter protein和unknown，采用qRT-PCR技术分析发现：它们在花芽自然休眠解除期间上调表达，认为曙光油桃花芽自然休眠解除的过程伴随着复杂物质代谢和能量消耗，差异基因主要参与植物生长发育及抗性胁迫等方面的调控，而且低温诱导的核糖体蛋白的表达可能也与花芽自然休眠解除有关。
PpDfn、PpDhn是文库中筛选到的全长cDNA序列，PpDfn、PpDhn基因编码的蛋白质氨基酸序列与其他物种的防御素蛋白（DFN、DHN）的氨基酸序列同源性较高。对二者进行蛋白质信号肽、跨膜结构域及疏水性分析，结果显示：PpDFN蛋白有1个信号肽，为分泌性蛋白。存在1个跨膜区，属于疏水性蛋白。通过对其氨基酸序列的进化树分析，发现与葡萄、油橄榄亲缘关系最近。PpDHN蛋白没有信号肽，无跨膜结构域，属于亲水性蛋白。通过对其氨基酸序列的进化树分析，发现与杏亲缘关系最近，二者聚在一起。采用qRT-PCR技术进一步分析了PpDFN、PpDHN在自然休眠解除期间、低温及短日照处理条件下的表达模式变化，发现PpDfn、PpDhn随自然休眠解除、低温处理时间的延长，基因表达量逐渐增加，冷处理2周时达到最高值；短日照处理后则没有明显变化。推测PpDfn基因随休眠解除及低温处理的上调表达，可能是参与了乙烯信号途径而诱导表达的结果。PpDhn基因随休眠解除及低温处理的上调表达可能与抗冻、清除自由基、保护生物膜免受伤害有关。
在油桃休眠及休眠解除期间，曙光油桃PIP1;1的转录水平呈现持续增高趋势，且1月15日的高水平表达使水分通过液泡膜和细胞质膜流出，减少了芽体水分含量，阻止细胞内冰晶的形成，从而抵御冻害；可溶性糖、可溶性蛋白、脯氨酸含量均达到最高，防止细胞的脱水伤害。低温处理2周后高水平表达说明PIP1;1为冷诱导基因。δTIP1的转录水平在休眠期间呈现波动性变化，至休眠解除时大幅度增高，这可能与休眠解除时，其上调表达被休眠解除信号及植物活性的增强所诱导有关。低温处理2周后，其表达没有升高，说明δTIP1并非冷诱导基因。
在曙光油桃休眠及休眠解除期间，深休眠期(11月15日)和休眠后期(12月15日)，喷布GA3和6-BA研究激素对解除休眠的效果。结果发现：在休眠后期施用GA3和6-BA能不同程度的提高H2O2含量和POD、SOD基因的表达量，降低CAT基因的表达水平。对萌芽率的统计表明，深休眠期施用GA3和6-BA对油桃花芽破眠无效，处理15天后没有花芽萌发。休眠后期施用GA3和6-BA破眠效果较为明显，萌芽较对照提前8d和3d，最终萌芽率达98%、91%，分别比对照提高了37%和30%。GA3的破眠效果好于6-BA。
The experiment was carried out in the horticulture experimental station of ShandongAgricultural University and central laboratory of College of Horticulture Science andEngineering during2008.8~2010.12. In the study, floral buds of ten-year-old field culturedand three-year-old potted nectarine (Prunus persica var. nectariana cv. Shuguang) trees wereused to identify and characterize regulatory genes of dormancy release. Expression changes ofthe genes in the dormancy release process under low temperature, short day and hormonetreatments were also detected. The study could be supplementary of the bud dormancymechanism and have important theoretical and practical value in the dormancy control ofprotected production of fruit trees. The main results were as follow:
The cDNA library of genes associated with bud dormancy release was established usingsuppression subtractive hybridization (SSH) procedure. The RNA of dormancy-released budswas used as Tester and the RNA from dormant buds was used as Driver.180up-regulatedpositive cDNA clones were detected and128of them were successfully sequenced.28uniquegenes related to dormancy release were screened after the elimination of redundant sequences.The results of BLAST analysis and molecular function prediction showed that genes involvedin defense and metabolism pathway made up lager percentages, accounting for25%and17.86%respectively. Genes associated with oxidation reduction and signaling accounted for14.29%and7.14%respectively. And genes related to activation of structure or function,unknown protein, un-match protein and transport activator protein accounted for35.71%.
According to functional predict of candidate genes and research progress of plantdormancy,8doemacy related genes, PpMt, PpDhn, PpHis, PpPod, PpSenescence-associatedprotein, PpCyp450, PpATP-binding cassette transporter protein and unknown, wereanalysised with qRT-PCR. The results showed that expression patterns of the genes wereup-regulated during dormancy release. Complex metabolism and energy consumption wereassociated with dormancy release, and the genes may be mainly involved in growth,development and tolerance to stress. The expression of ribosomal proteins induced by lowtemperature may be related to dormancy release of floral buds.
PpDfn、PpDhn were full-length cDNA sequence screened from the library. Amino acidsequences of PpDFN、PpDHN were highly homologous with defensins (DFN, DHN) of other species. Analysis of signal peptide, trans-membrane domain and hydrophobicity of the twogenes suggested that PpDFN was a secretory and hydrophobic protein containning a signalpeptide and a trans-membrane domain. PpDHN was hydrophilic protein without signalpeptide and trans-membrane domain. Cladogram analysis of amino acid sequences showedthat PpDFN of ‘Shuguang’ nectarine had closer genetic relationship with those of grape andolive and PpDHN had closer genetic relationship with that of Armeniaca. Expression analysisof PpDFN and PpDHN showed that expression of the two genes were up-regulated along withchilling accumulation during dormancy release, and peaked at the2thweek of chillingtreatment. Short day showed no obvious effect on the expression of the two genes. PpDFNmight be involved in ethylene signaling pathway and PpDHN might related tofreeze-resistance, eliminating free radicals and protecting bio-membrane from injury.
At stage of dormancy and dormancy release, transcripts of PIP1;1increased consistently.The high expression on15thJanuary led to outflow of water through vacuolar membrane andcytoplasm membrane, which caused water content decrease in buds and prevented frozeninjury. Contents of soluble sugars, proteins and proline peaked at the same time, whichprevented cell from dehydration injury. The high level expression after2weeks of lowtemperature treatment suggested that PIP1;1was a cold-inducible gene. Transcription ofδTIP1fluctuated during dormancy but increased significantly at stage of dormancy release.The increase might be related to dormancy release signal and plant activity enhancement.Expression of δTIP1did not increase after2weeks of low temperature treatment, indicatingthat δTIP1was not a cold-inducible gene.
GA3and6-BA were used at15thNovember (endo-dormancy) and15thDecember (latestage of endo-dormancy) to study effects of hormone on dormancy release. Application ofGA3and6-BA at late stage of dormancy increased H2O2level and expression of POD andSOD, but decrease the expression of CAT. Application of GA3and6-BA duringendo-dormancy did not increase budburst of dormant buds. However, at the late stage ofendo-dormancy, budburst time was brought forward by8and3days respectively by GA3and6-BA treatment, budburst rate were also increased by37%and30%respectively. Effect ofGA3on dormancy release was more significant than6-BA.
引文
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