甜瓜果实发育相关基因的鉴定、表达特性及功能分析
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摘要
甜瓜是全世界广泛栽培的园艺作物,是重要的经济作物之一。随着对甜瓜研究的深入和它本身的植物学特性,甜瓜已成为阐明肉质果实发育成熟分子机理的另一种重要的模式植物。
乙烯对跃变型果实的成熟衰老起到重要作用,同时影响着果实的成熟及采后的耐贮藏性。为了分离鉴定甜瓜果实乙烯跃变中关键基因和在转录水平上阐明果实发育的分子机理,本研究以典型的呼吸跃变型甜瓜品种河套蜜瓜为研究对象,以甜瓜果实内源乙烯跃变中上升、峰值、下降三个时期中果皮组织mRNA等量混合物作为乙烯跃变期样品,以乙烯跃变前5天的mRNA为乙烯跃变前期样品,构建了正向和反向抑制性差减文库,并对所分离的EST进行克隆、测序和数据库中寻找同源基因。其中,正向文库中分离得到386条有效的EST序列,总计179个unigenes；反向文库中得到417条有效的EST序列,总计213个unigenes；所获得的EST主要编码物质和能量代谢、转录、蛋白翻译后修饰与转运及分子伴侣、结合蛋白、信号转导、生长发育、转运、机体防御与自救、未知功能等相关基因。对文库中随机选取6个候选基因实时荧光定量PCR检测结果显示,Chibl、L-Asp、CSGP和ACO1基因的1mRNA含量在果实内源乙烯跃变时期(授粉后40-45天)的果实中急剧增高,其峰值期与果实内源乙烯峰值期同步,而DAFP和XTH2基因在果实跃变前期表达水平较高。
分别克隆了甜瓜信号转导组分Cm-CTR1、Cm-EIN2、Cm-EIL1、Cm-EIL2. Cm-ERF1、Cm-ERF2基因的全长cDNA.应用荧光定量PCR技术分析了上述基因在甜瓜果实不同发育时期的表达特性。定量PCR分析结果显示：15DAP到45DAP的果实中,Cm-CTR1基因均具有较高的表达,且转录水平变化较少,在果实内源乙烯跃变时稍减少。cm-EIN2在幼叶和花瓣组织中的表达量相似,约为根中表达量的2倍,茎中表达最低,子房中最高。在果实发育成熟过程中,cm-EIN2基因：mRNA水平从20DAP开始缓慢增高,在授粉后35天达到峰值,随后下降。将幼叶用不同浓度的IAA和ABA处理后,Cm-EIN2基因转录水平减少。Cm-EIL1、Cm-EIL2、Cm-ERF1和Cm-ERF2表达模式与甜瓜果实成熟过程及乙烯生成量显著相关。上述基因表达分析结果表明,Cm-EIL1、Cm-EIL2、 cm-ERF1、cm-ERF2基因可能在甜瓜果实乙烯跃变过程中具有重要作用,而cm-EIN2和Cm-CTR1可能在甜瓜果实发育过程中发挥重要作用。
利用烟草核基质附着区序列和甜瓜果实特异性表达的黄瓜素基因启动子,构建了无载体无选择标记基因的果实特异性稳定表达反义ACO1基因线性表达盒(RB-MAR-CMC-antiACO1-nos-MAR-LB),应用花粉管通道法转化甜瓜。经PCR检测证明外源基因已经整合到受体植物的基因组中。获得了耐贮性强的3个株系,经正常商业采摘期采收果实后,转基因T2果实内源乙烯的含量均明显低于未转基因对照果实,在采后第12天,约为对照果实的4%,没有出现内源乙烯释放高峰；刚采摘时,转基因和未转基因果实硬度基本相同,贮藏到一定天数时,转基因果实硬度基本保持不变,而未转基因果实硬度逐渐下降；转基因与对照果实的可溶性固形物在贮藏过程中基本相同。
Melon is one of the important economic crops and widely cultivated all over the world. However, it gives rise to huge economic losses because of the over ripeness, softening and rottenness within a short postharvest storage. Melon has become another important model plant to clarify the molecular mechanisms of fleshy fruit development and ripening due to it's research in-depth and botanical characteristics. Ethylene regulates the ripening process and affects the ripening rate and postharvest storability in climacteric fruit. Therefore, it has become a hot topic to study the molecular mechanism of melon fruit ripening via ethylene.
To screen for genes that are differentially expressed at the burst of ethylene climacteric, and help to elucidate the molecular mechanism of ethylene climacteric at the transcriptional level in melon fruit, we performed suppression subtractive hybridization (SSH) to generate forward and reverse libraries, for which2groups of fruit samples, from the typically respiratory climacteric melon fruit of Hetao, at disparate stages were used. Forward subtraction was performed using ethylene climacteric fruit cDNA (poly A+RNA of the fruits—when ethylene production increased, peaked, and decreased respectively—mixed in equal proportions) as the tester and pre-ethylene climacteric (5days before the ethylene climacteric) fruit cDNA as the driver. For the corresponding reverse libraries, the cDNAs were reciprocated. The isolated EST clones were sequenced, and databases were searched for homologous genes. From the forward library, the386EST sequences were assembled into179unigenes, while the417EST sequences were assembled into213unigenes in the reverse library. The identified ESTs encoded proteins that mediate metabolism of energy and substances, transcription, posttranslational modification, protein turnover and chaperones, binding protein, signal transduction, growth and development, transport, body defense and self-rescue, unknown function etc. The expression patterns of the6randomly selected candidate genes in fruit at various stages of development were analyzed by qRT-PCR. The results of showed that the transcription of the Chib1, L-Asp, CSGP and ACO1genes increased dramatically during fruit ripening (40-45DAP) and the peaks of Chibl, L-Asp, CSGP and ACO1mRNAs coincided with the climacteric peak of ethylene production. The XTH2and DAFP mRNAs levels were higher during pre-climacteric stage of ethylene.
In this study, the full-length cDNAs of the ethylene signal transduction component genes Cm-CTR1, Cm-EIN2, Cm-EIL1, Cm-EIL2, Cm-ERF1and Cm-ERF2were cloned, and the expression patterns of these genes were analyzed by QRT-PCR. Quantitative RT-PCR analysis indicated as follows:The transcript level of the Cm-CTR1was higher and had few changes from15DAP to45DAP, while slightly reduced at ethylene climacteric stage; Cm-EIN2had similar expression patterns in young leaves and petal tissues—approximately2times higher than in root, the minimum and maximum expression were observed in the stem and ovary, respectively. During fruit development, Cm-EIN2mRNA increased slightly from20DAP, peaked at35DAP, and declined. Leaves that were treated with IAA and ABA had lower levels of Cm-EIN2transcripts. The expression patterns of Cm-EILl, Cm-EIL2, Cm-ERF1and Cm-ERF2genes were highly relative to fruit ripening process and ethylene production. The expression patterns of these genes reveals that Cm-EIL1, Cm-EIL2, Cm-ERF1and Cm-ERF2genes may regulate the climacteric ripening, while Cm-EIN2and Cm-CTR1gene may affect the fruit development.
The vector-free and marker-free linear cassettes of a fruit-specific stable expression of ACO1antisense gene was constructed, which was combined with the matrix attachment region sequence of tobacco and fruit-specific expression promoter of Cucumisin gene in melon, for the first time (RB-MAR-CMC-antiACO1-nos-MAR-LB). The melon cultivar Hetao was transformed with these linear gene cassettes via pollen-tube pathway. The PCR data showed that the transgenes were inserted into the recipient's genome.3strains with improved storage capacity have been selected. The fruits were harvested in normal commercial picking stages. The endogenous ethylene contents of the transgenic T2fruits were significantly lower than non-transgenic ones, and were approximately4%of the control at12days post-harvest. It was not found the peak of endogenous ethylene in transgenic fruits. The fruit firmness between the transgenic T2and non-transgenic control was approximately the same when harvest, however, the transgenic fruit firmness remained basically unchanged, while the control one gradually decreased after storage a number of days. The soluble solids contents between the transgenic T2and non-transgenic control were approximately the same during storage.
乙烯对跃变型果实的成熟衰老起到重要作用,同时影响着果实的成熟及采后的耐贮藏性。为了分离鉴定甜瓜果实乙烯跃变中关键基因和在转录水平上阐明果实发育的分子机理,本研究以典型的呼吸跃变型甜瓜品种河套蜜瓜为研究对象,以甜瓜果实内源乙烯跃变中上升、峰值、下降三个时期中果皮组织mRNA等量混合物作为乙烯跃变期样品,以乙烯跃变前5天的mRNA为乙烯跃变前期样品,构建了正向和反向抑制性差减文库,并对所分离的EST进行克隆、测序和数据库中寻找同源基因。其中,正向文库中分离得到386条有效的EST序列,总计179个unigenes；反向文库中得到417条有效的EST序列,总计213个unigenes；所获得的EST主要编码物质和能量代谢、转录、蛋白翻译后修饰与转运及分子伴侣、结合蛋白、信号转导、生长发育、转运、机体防御与自救、未知功能等相关基因。对文库中随机选取6个候选基因实时荧光定量PCR检测结果显示,Chibl、L-Asp、CSGP和ACO1基因的1mRNA含量在果实内源乙烯跃变时期(授粉后40-45天)的果实中急剧增高,其峰值期与果实内源乙烯峰值期同步,而DAFP和XTH2基因在果实跃变前期表达水平较高。
分别克隆了甜瓜信号转导组分Cm-CTR1、Cm-EIN2、Cm-EIL1、Cm-EIL2. Cm-ERF1、Cm-ERF2基因的全长cDNA.应用荧光定量PCR技术分析了上述基因在甜瓜果实不同发育时期的表达特性。定量PCR分析结果显示：15DAP到45DAP的果实中,Cm-CTR1基因均具有较高的表达,且转录水平变化较少,在果实内源乙烯跃变时稍减少。cm-EIN2在幼叶和花瓣组织中的表达量相似,约为根中表达量的2倍,茎中表达最低,子房中最高。在果实发育成熟过程中,cm-EIN2基因：mRNA水平从20DAP开始缓慢增高,在授粉后35天达到峰值,随后下降。将幼叶用不同浓度的IAA和ABA处理后,Cm-EIN2基因转录水平减少。Cm-EIL1、Cm-EIL2、Cm-ERF1和Cm-ERF2表达模式与甜瓜果实成熟过程及乙烯生成量显著相关。上述基因表达分析结果表明,Cm-EIL1、Cm-EIL2、 cm-ERF1、cm-ERF2基因可能在甜瓜果实乙烯跃变过程中具有重要作用,而cm-EIN2和Cm-CTR1可能在甜瓜果实发育过程中发挥重要作用。
利用烟草核基质附着区序列和甜瓜果实特异性表达的黄瓜素基因启动子,构建了无载体无选择标记基因的果实特异性稳定表达反义ACO1基因线性表达盒(RB-MAR-CMC-antiACO1-nos-MAR-LB),应用花粉管通道法转化甜瓜。经PCR检测证明外源基因已经整合到受体植物的基因组中。获得了耐贮性强的3个株系,经正常商业采摘期采收果实后,转基因T2果实内源乙烯的含量均明显低于未转基因对照果实,在采后第12天,约为对照果实的4%,没有出现内源乙烯释放高峰；刚采摘时,转基因和未转基因果实硬度基本相同,贮藏到一定天数时,转基因果实硬度基本保持不变,而未转基因果实硬度逐渐下降；转基因与对照果实的可溶性固形物在贮藏过程中基本相同。
Melon is one of the important economic crops and widely cultivated all over the world. However, it gives rise to huge economic losses because of the over ripeness, softening and rottenness within a short postharvest storage. Melon has become another important model plant to clarify the molecular mechanisms of fleshy fruit development and ripening due to it's research in-depth and botanical characteristics. Ethylene regulates the ripening process and affects the ripening rate and postharvest storability in climacteric fruit. Therefore, it has become a hot topic to study the molecular mechanism of melon fruit ripening via ethylene.
To screen for genes that are differentially expressed at the burst of ethylene climacteric, and help to elucidate the molecular mechanism of ethylene climacteric at the transcriptional level in melon fruit, we performed suppression subtractive hybridization (SSH) to generate forward and reverse libraries, for which2groups of fruit samples, from the typically respiratory climacteric melon fruit of Hetao, at disparate stages were used. Forward subtraction was performed using ethylene climacteric fruit cDNA (poly A+RNA of the fruits—when ethylene production increased, peaked, and decreased respectively—mixed in equal proportions) as the tester and pre-ethylene climacteric (5days before the ethylene climacteric) fruit cDNA as the driver. For the corresponding reverse libraries, the cDNAs were reciprocated. The isolated EST clones were sequenced, and databases were searched for homologous genes. From the forward library, the386EST sequences were assembled into179unigenes, while the417EST sequences were assembled into213unigenes in the reverse library. The identified ESTs encoded proteins that mediate metabolism of energy and substances, transcription, posttranslational modification, protein turnover and chaperones, binding protein, signal transduction, growth and development, transport, body defense and self-rescue, unknown function etc. The expression patterns of the6randomly selected candidate genes in fruit at various stages of development were analyzed by qRT-PCR. The results of showed that the transcription of the Chib1, L-Asp, CSGP and ACO1genes increased dramatically during fruit ripening (40-45DAP) and the peaks of Chibl, L-Asp, CSGP and ACO1mRNAs coincided with the climacteric peak of ethylene production. The XTH2and DAFP mRNAs levels were higher during pre-climacteric stage of ethylene.
In this study, the full-length cDNAs of the ethylene signal transduction component genes Cm-CTR1, Cm-EIN2, Cm-EIL1, Cm-EIL2, Cm-ERF1and Cm-ERF2were cloned, and the expression patterns of these genes were analyzed by QRT-PCR. Quantitative RT-PCR analysis indicated as follows:The transcript level of the Cm-CTR1was higher and had few changes from15DAP to45DAP, while slightly reduced at ethylene climacteric stage; Cm-EIN2had similar expression patterns in young leaves and petal tissues—approximately2times higher than in root, the minimum and maximum expression were observed in the stem and ovary, respectively. During fruit development, Cm-EIN2mRNA increased slightly from20DAP, peaked at35DAP, and declined. Leaves that were treated with IAA and ABA had lower levels of Cm-EIN2transcripts. The expression patterns of Cm-EILl, Cm-EIL2, Cm-ERF1and Cm-ERF2genes were highly relative to fruit ripening process and ethylene production. The expression patterns of these genes reveals that Cm-EIL1, Cm-EIL2, Cm-ERF1and Cm-ERF2genes may regulate the climacteric ripening, while Cm-EIN2and Cm-CTR1gene may affect the fruit development.
The vector-free and marker-free linear cassettes of a fruit-specific stable expression of ACO1antisense gene was constructed, which was combined with the matrix attachment region sequence of tobacco and fruit-specific expression promoter of Cucumisin gene in melon, for the first time (RB-MAR-CMC-antiACO1-nos-MAR-LB). The melon cultivar Hetao was transformed with these linear gene cassettes via pollen-tube pathway. The PCR data showed that the transgenes were inserted into the recipient's genome.3strains with improved storage capacity have been selected. The fruits were harvested in normal commercial picking stages. The endogenous ethylene contents of the transgenic T2fruits were significantly lower than non-transgenic ones, and were approximately4%of the control at12days post-harvest. It was not found the peak of endogenous ethylene in transgenic fruits. The fruit firmness between the transgenic T2and non-transgenic control was approximately the same when harvest, however, the transgenic fruit firmness remained basically unchanged, while the control one gradually decreased after storage a number of days. The soluble solids contents between the transgenic T2and non-transgenic control were approximately the same during storage.
引文
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