FRUITFULL调控番茄果实发育与成熟的机理研究
摘要
果实是植物繁衍后代的重要器官,也是人类膳食结构的重要组成部分,其发育与成熟过程一直备受研究者关注。番茄(Solanum lycopersicum)属于研究浆果类植物发育成熟的模式植物,通过对番茄果实发育成熟突变体的研究,已经鉴定出多个调控果实发育成熟的基因。在这些基因中,MADS-box家族转录因子RIN是一个研究重点,它位于乙烯合成途径上游来调控果实发育成熟,且这种调控在呼吸跃变和非呼吸跃变型果实中呈保守机制。RIN可以通过与互作蛋白形成二聚体或者多聚体来调控成熟相关基因表达,最终调控果实成熟。
本研究通过酵母双杂交筛选出与RIN的互作蛋白FUL1和FUL2,转基因功能分析发现它们在番茄植物体及果实发育成熟中起着重要的作用。主要研究结果如下：
1、酵母双杂交和双分子荧光互补实验表明,FUL1和FUL2是RIN的互作蛋白。FUL1和FUL2序列上表现出高度同源性,都具有植物MADS-box蛋白的典型MIKC结构。组织表达谱结果显示,FUL1同RIN一样,更偏向于在果实成熟时期表达；FUL2在除了根系以外所有组织中均有表达。说明FUL1和FUL2在番茄植株生长发育过程中的调控可能存在时空上的异同。
2、转基因功能分析显示：单独干涉FUL1或FUL2的转基因番茄株系均无明显表型。同时干涉FUL1和FUL2的转基因番茄株系果实成熟后不能正常变红,表现为亮黄色。FUL1/FUL2-RN Ai转基因果实果皮中的类胡萝卜素含量下降一倍以上,八氢番茄红素和番茄红素含量极低。果实中质体数目和体积都比野生型要小,且红熟期转基因果实中质体已经瓦解。分析发现类胡萝卜素合成途径中,催化合成八氢番茄红素的ZDS和催化合成番茄红素的PSY1的表达量在转基因果实中分别降低了一倍以上。
3、FUL1/FUL2-RNAi转基因果实与乙烯调控的关系分析表明,成熟过程中转基因果实乙烯释放量大量减少,且没有乙烯呼吸高峰。分析乙烯合成途径关键基因的表达,发现ACO1、ACS2和ACS4的表达量在转基因果实中均明显下调。
4、通常乙烯生成量低的果实贮藏性好,然而FUL1/FUL2-RNAi转基因果实乙烯释放量下降,其采后耐贮性却降低,且失水率比野生型高。分析发现乙烯信号转导途径的受体蛋白ETR4在转基因果实中表达量下调,且对野生型果实施加外源乙烯或者乙烯抑制剂会影响FUL1和FUL2的表达。FUL2-OE转基因果实采后失水率较低,分析发现外果皮表面的角质层密度明显大于野生型。
5、超量表达FUL1植株无明显表型变化,而超量表达FUL2影响了番茄植株生长发育,包括茎杆变细、叶片变小、果皮变薄及果实形状发生变化。FUL2-OE转基因植株的茎杆及果皮细胞体积均小于野生型。FUL2-OE转基因果实中细胞扩张蛋白SlEXP1的表达量明显被抑制。内源启动子驱动GUS (ProFUL2::GUS)的转基因植株分析发现,FUL2主要在果皮薄壁细胞和茎杆形成层表达。
As the reproductive organ and the important component of human diets, the development and ripening of fruit have been paid significant experimental attention. Many fruit-ripening mutants and ripening-related genes were identified in tomato (Solarium lycopersicum) which is the typical model plant for molecular mechanism research of fresh fruit development. The MADS-box transcription factor MADS-RIN is upstream regulator of ethylene biosynthesis and the regulation is conserved in both climacteric and non-climacteric fruits. Further studies revealed that RIN could form higher order complexes to regulate the expression of ripening related genes during tomato fruit ripening.
In this study, FUL1and FUL2were screened as the interactor of RIN by yeast two-hybrid systerm. The function analysis suggested that FUL1and FUL2play important roles in tomato development and fruit ripening. The main results were as follows:
1. FUL1and FUL2were identified as encoding proteins that interact with MADS-RIN in Yeast two-hybrid and BIFC assays. FUL1and FUL2share a high degree of sequence similarity and both of them contain conserved MIKC domains that typify plant MADS-box proteins. Expression patterns of FUL1and FUL2showed that FUL1was mainly expressed in fruit ripening stage. However, FUL2was expressed in all tissues except root, which indicated that FUL1and FUL2might have distinct functions in the regulation of tomato plant growth.
2. No visible phenotypes changing were observed on either FUL1-RNAi or FUL2-RNAi transgenic tomato lines, however, doubly silenced FUL1and FUL2could result in the bright yellow fruit during ripening. The abundance of phytoene and lycopene reduced dramatically in FUL1/FUL2-RNAi transgenic tomato. Fewer and smaller plastoglobules were observed in FUL1/FUL2doubly silenced pericarp. The transcript abundance of ZDS and PSY1, which respectively responsible for the accumulation of phytoene and lycopene, was approximately50%in FUL1/FUL2-RNAi transgenic tomato.
3. The fruits from the FUL1/FUL2doubly silenced lines showed a substantial reduction in ethylene production compared with the wild-type, and had no typical peak during ripening. The transcript abundance of ethylene biosynthetic genes ACS2, ACS4and ACO1showed a similar reduction in the FUL1/FUL2doubly silenced fruit.
4. Lower ethylene release could usually prolong shelf-life of tomato fruit. However, FUL1/FUL2doubly silenced fruits which showed a reduced ethylene production had the shorter shelf-life and faster dehydration. Further analysis indicated that the transcript abundance of ethylene receptor ETR1was higher in the FUL1/FUL2doubly silenced fruit. Exogenous ethylene signal substances such as ethephon and1-MCP could influence the expressions of FUL1and FUL2. For comparison, the slower dehydration of FUL2-OE fruit was mostly due to the denser cuticle layer covered on the surface of fruits.
5. No visible phenotypes changing were observed on FUL1-OE transgenic tomato lines, however, overexpression of FUL2could influence the development of tomato plant, such as the reduced stem diameter, smaller leaves, thinner pericarp and altered fruit shape. The cell size of both stem and pecicarp was smaller than that of wild-type. The transcript abundance of expansin SIEXP1in FUL2-OE transgenic tomato was significantly lower than that of wild-type. GUS staining of ProFuL2::GUS transgenic plant revealed that the FUL2gene showed high expression in parenchyma in fruit and cambium in stem.
本研究通过酵母双杂交筛选出与RIN的互作蛋白FUL1和FUL2,转基因功能分析发现它们在番茄植物体及果实发育成熟中起着重要的作用。主要研究结果如下：
1、酵母双杂交和双分子荧光互补实验表明,FUL1和FUL2是RIN的互作蛋白。FUL1和FUL2序列上表现出高度同源性,都具有植物MADS-box蛋白的典型MIKC结构。组织表达谱结果显示,FUL1同RIN一样,更偏向于在果实成熟时期表达；FUL2在除了根系以外所有组织中均有表达。说明FUL1和FUL2在番茄植株生长发育过程中的调控可能存在时空上的异同。
2、转基因功能分析显示：单独干涉FUL1或FUL2的转基因番茄株系均无明显表型。同时干涉FUL1和FUL2的转基因番茄株系果实成熟后不能正常变红,表现为亮黄色。FUL1/FUL2-RN Ai转基因果实果皮中的类胡萝卜素含量下降一倍以上,八氢番茄红素和番茄红素含量极低。果实中质体数目和体积都比野生型要小,且红熟期转基因果实中质体已经瓦解。分析发现类胡萝卜素合成途径中,催化合成八氢番茄红素的ZDS和催化合成番茄红素的PSY1的表达量在转基因果实中分别降低了一倍以上。
3、FUL1/FUL2-RNAi转基因果实与乙烯调控的关系分析表明,成熟过程中转基因果实乙烯释放量大量减少,且没有乙烯呼吸高峰。分析乙烯合成途径关键基因的表达,发现ACO1、ACS2和ACS4的表达量在转基因果实中均明显下调。
4、通常乙烯生成量低的果实贮藏性好,然而FUL1/FUL2-RNAi转基因果实乙烯释放量下降,其采后耐贮性却降低,且失水率比野生型高。分析发现乙烯信号转导途径的受体蛋白ETR4在转基因果实中表达量下调,且对野生型果实施加外源乙烯或者乙烯抑制剂会影响FUL1和FUL2的表达。FUL2-OE转基因果实采后失水率较低,分析发现外果皮表面的角质层密度明显大于野生型。
5、超量表达FUL1植株无明显表型变化,而超量表达FUL2影响了番茄植株生长发育,包括茎杆变细、叶片变小、果皮变薄及果实形状发生变化。FUL2-OE转基因植株的茎杆及果皮细胞体积均小于野生型。FUL2-OE转基因果实中细胞扩张蛋白SlEXP1的表达量明显被抑制。内源启动子驱动GUS (ProFUL2::GUS)的转基因植株分析发现,FUL2主要在果皮薄壁细胞和茎杆形成层表达。
As the reproductive organ and the important component of human diets, the development and ripening of fruit have been paid significant experimental attention. Many fruit-ripening mutants and ripening-related genes were identified in tomato (Solarium lycopersicum) which is the typical model plant for molecular mechanism research of fresh fruit development. The MADS-box transcription factor MADS-RIN is upstream regulator of ethylene biosynthesis and the regulation is conserved in both climacteric and non-climacteric fruits. Further studies revealed that RIN could form higher order complexes to regulate the expression of ripening related genes during tomato fruit ripening.
In this study, FUL1and FUL2were screened as the interactor of RIN by yeast two-hybrid systerm. The function analysis suggested that FUL1and FUL2play important roles in tomato development and fruit ripening. The main results were as follows:
1. FUL1and FUL2were identified as encoding proteins that interact with MADS-RIN in Yeast two-hybrid and BIFC assays. FUL1and FUL2share a high degree of sequence similarity and both of them contain conserved MIKC domains that typify plant MADS-box proteins. Expression patterns of FUL1and FUL2showed that FUL1was mainly expressed in fruit ripening stage. However, FUL2was expressed in all tissues except root, which indicated that FUL1and FUL2might have distinct functions in the regulation of tomato plant growth.
2. No visible phenotypes changing were observed on either FUL1-RNAi or FUL2-RNAi transgenic tomato lines, however, doubly silenced FUL1and FUL2could result in the bright yellow fruit during ripening. The abundance of phytoene and lycopene reduced dramatically in FUL1/FUL2-RNAi transgenic tomato. Fewer and smaller plastoglobules were observed in FUL1/FUL2doubly silenced pericarp. The transcript abundance of ZDS and PSY1, which respectively responsible for the accumulation of phytoene and lycopene, was approximately50%in FUL1/FUL2-RNAi transgenic tomato.
3. The fruits from the FUL1/FUL2doubly silenced lines showed a substantial reduction in ethylene production compared with the wild-type, and had no typical peak during ripening. The transcript abundance of ethylene biosynthetic genes ACS2, ACS4and ACO1showed a similar reduction in the FUL1/FUL2doubly silenced fruit.
4. Lower ethylene release could usually prolong shelf-life of tomato fruit. However, FUL1/FUL2doubly silenced fruits which showed a reduced ethylene production had the shorter shelf-life and faster dehydration. Further analysis indicated that the transcript abundance of ethylene receptor ETR1was higher in the FUL1/FUL2doubly silenced fruit. Exogenous ethylene signal substances such as ethephon and1-MCP could influence the expressions of FUL1and FUL2. For comparison, the slower dehydration of FUL2-OE fruit was mostly due to the denser cuticle layer covered on the surface of fruits.
5. No visible phenotypes changing were observed on FUL1-OE transgenic tomato lines, however, overexpression of FUL2could influence the development of tomato plant, such as the reduced stem diameter, smaller leaves, thinner pericarp and altered fruit shape. The cell size of both stem and pecicarp was smaller than that of wild-type. The transcript abundance of expansin SIEXP1in FUL2-OE transgenic tomato was significantly lower than that of wild-type. GUS staining of ProFuL2::GUS transgenic plant revealed that the FUL2gene showed high expression in parenchyma in fruit and cambium in stem.
引文
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