摘要
莲是我国重要的水生蔬菜,具有很大的经济和观赏价值。目前人们已经从莲的形态学、分类学、生理学、药物学等方面进行了广泛的研究。随着功能基因组学的迅猛发展,克隆和鉴定各种抗性、品质基因,利用基因工程技术培育高产优质和耐胁迫的新品种,已成为未来解决农业生产问题的重要技术。作为一种重要的经济作物,莲的分子生物学研究却相对比较匮乏。本文通过构建莲cDNA文库,克隆了与莲在胁迫环境下发挥重要作用的一些基因,并对其功能进行了初步的分析,为莲的转基因研究和应用提供了一些分子证据。
1莲cDNA文库的构建
为发掘各种影响莲抗性、品质和具有调控作用的功能基因,我们采用SMART技术成功构建了藕莲和花莲顶芽的cDNA文库。花莲原始cDNA文库滴度达到4.2×106pfu/mL,重组率为100%,97%的文库插入片断在0.5kb以上,扩增文库的总容量达到8.5×108pfu/mL,保障了高比例全长cDNA的获得。通过已知的NnUBC3基因对花莲文库进行筛选验证,成功得到约586bp的全长目的基因,其基因编码区域为459bp,3'UTR序列为127bp。这些数据表明,我们构建的花莲顶芽cDNA文库的有效性,适合以后的测序和基因功能研究。
2NnGPX1的克隆及在胁迫环境中的功能分析
GPX是一种抗氧化酶,能催化H202或者过氧化氢物形成水或者乙醇,使机体免受氧化损伤。我们在莲中克隆了NnGPX1的全长cDNA,通过对莲进行不同Na2Se03浓度处理发现,NnGPX1与大多数植物GPX一样,是一个不依赖硒的GPX蛋白。不同发育时期的组织表达分析发现,NnGPX1在幼苗时期的叶和开花时期的顶芽中表达水平最高,在成熟的胚芽中也有一定的表达。植物GPX家族的成员分别在不同的胁迫环境中发挥作用。我们对莲进行冷、热、机械损伤、高盐、ABA、PEG等不同时间梯度的胁迫处理,发现NnGPX1表达水平在各种处理后都迅速升高,这与其它植物GPX不同,说明NnGPX1在莲生长发育和面对胁迫环境时起着重要的调节作用。同时我们对NnGPX1进行了表达,纯化和蛋白质结构的预测,有待进行更进一步的功能分析。
3莲泛素转运酶NnUBC的克隆与功能分析
E2(UBC)是真核生物泛素化必需的酶之一,与E1,E3共同催化底物蛋白的泛素化。组蛋白H2B单泛素化影响着开花基因的转录与表达,同时泛素化在植物应对胁迫环境时也发挥着重要作用。我们克隆了莲的4个NnUBC全长cDNA,荧光定量PCR的结果显示这4个基因在莲不同组织中有着不同的表达模式,其中NnUBC3在植株的各个组织中表达量最高。于是我们进一步获得了NnUBC3的全长基因组DNA,并通过NnUBC3全长CDS对拟南芥ubcl,2双突变体进行转化,使突变体表型得以恢复。NnUBC3的亚细胞定位结果与拟南芥UBC一致,分布于细胞质和细胞核内,证明NnUBC是泛素转运酶。在对莲进行了不同的处理后发现,NnUBC3在NaCl, ABA, PEG胁迫中发挥着重要的调控的作用。
4VvPYL在ABA信号通路中的功能分析
ABA是植物应对生物和非生物胁迫的重要调节因子,特别是干旱和高盐环境下,通过调控ABA应答基因表达来控制植物生长。我们克隆了3个葡萄的VvPYL,氨基酸序列比对和聚类分析结果显示与拟南芥PYL家族有很高的同源性。对VvPYL进行组织特异性表达分析发现,VvPYL1在叶,茎和根中的表达量都比较高,且在ABA处理后表达水平上调。ITC实验结果表明,在体外VvPYL1能与ABA结合,解离常数为78uM。磷酸酶活性测试结果显示,VvPYL1在ABA存在时,几乎完全抑制拟南芥ABI1的磷酸酶活性。通过对小烟草的转化发现,VvPYL1亚细胞定位于细胞质和细胞核中,结果与拟南芥PYL的定位一致。所有结果证明VvPYL1是ABA的受体,这也是关于葡萄ABA受体研究的首次报道。
Nelumbo nucifera is one of the ancient species in plant and has enormous economic value as aquatic vegetable in China. Research of Nelumbo nucifera has been mainly focused on its morphology, taxology and physiology that is related to clinic medicine. With rapid development in functional genomics, it is important to breed new species of high production, high quality and stress tolerance, however few studies of molecular cloning and transgenic engineering were reported in N. nucifera so far. In this thesis work, we have constructed cDNA libraryin Nelumbo nucifera and cloned stress-related genes and analyzed the fuctions of these genes in Nelumbo nucifera. Our results, for the first time, provide molecular evidence for transgenenic research and application in Nelumbo nucifera.
1Construction of cDNA library in Nelumbo nucifera
To discover functional gene that is related to stress tolerance, quality and transcriptional regulation, we successfully constructed the cDNA library of apical bud in Nelumbo nucifera. The titer of primary cDNA library is4.2×106pfu/mL and the ratio of combinant is100%. The inserted fragment over0.5kb occupied97%of the whole library and the titer of amplified cDNA library is8.5×10pfu/mL, which guarantee the acquisition of full length cDNA. We screened the library through the known NnUBC3gene and successfully obtained the586bp full length cDNA sequence with459bp of coding sequence and127bp of3'UTR. These data suggest that the cDNA library is effective and suitable for sequencing and functional analysis in the future.
2Cloning of NnGPXl and its functional analysis in response to abiotic stress
GPX is an antioxidant enzyme that is able to catalyze the reduction of H2O2or organic hydroperoxides to water or corresponding alcohols and protect organism from oxidant. We cloned full length cDNA of GPX in Nelumbo nucifera named NnGPX1and found that NnGPX1, unlike GPX protein in other species, is not a selenium-dependent glutathione peroxidase. Through expression analysis in various tissues at different developmental stages, we found that NnGPX1is up-regulated in leaf of seedling and apical bud at flowring while it is highly expressed after treatment of chilling, hot, mechanical wounding, salty, ABA and PEG. This is not consistent with previous data in Arabidopsis that different GPX functions in different stress condition. All of data indicate that NnGPX1play a critical role in development, biotic and abiotic stress. Moreover, we purified the NnGPX1protein and predicted its structure for further functional analysis.
3Cloning of NnUBC and its functional analysis
UBC is ubquitin conjugating enzyme catalyzing ubiquitination with E1and E3in eukaryote. Histone H2B monoubiquitination regulate plant flowering time and ubiquitination has also been found important in defense of biotic and abiotic stress. We cloned full length cDNA of four NnUBC genes and found that they show different expression pattern, especially NnUBC3expressed highly in all tested organs through expression analysis in various tissues. Consequently, we cloned the full length genomic DNA sequence of NnUBC3and discovered that it is able to restore the phenotype of mutant by introducing full CDS of NnUBC3into ubcl,2mutant of Arabidopsis. The subcellular localization of NnUBC3resembles its location in Arabidopsis. All the results suggest that NnUBC3is a E2. In conclusion, NnUBC3may play important role in response to NaCl, ABA and PEG.
4Functional analysis of VvPYLl in ABA signalling pathway
ABA is an important regulator in response to biotic and abiotic stress, and control plant growth through regulating ABA responsive gene, especially in drought and salty condition. We cloned three VvPYL genes from Vitis Vinifera and discovered they are homolog of PYL by alignment of amino acid sequence and analysis of phylogenetic tree. Expression analysis in various tissues showed that VvPYLl is highly expressed in all tested tissues and upregulated in response to ABA treatment. ITC assays indicate that VvPYL1can interact with ABA at78uM of Kd value. Phosphatase activity assay demonstrates that VvPYL1is able to completely inhibit the phosphatase activity of ABI1in an ABA-dependent manner. VvPYL1is present both in cytosol and nucleus by Agroinfiltration in tobacco(Nicotiana benthamiana), which resembles the subcellular location of PYL in Arabidopsis. All these data, for the first time, suggest that VvPYL1is an ABA receptor in Vitis Vinifera.
引文
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