摘要
果蔬从采后运输、贮藏到消费者手中的整个过程中都不可避免地受到不同程度的机械损伤,机械损伤破坏了果实天然的组织结构,引起活性氧的代谢失去平衡,对果蔬组织产生毒害现象。当果蔬体内大量活性氧积累时,抗氧化系统的一系列酶类随即被激活,以清除活性氧为目的,以清除和减少细胞膜的氧化伤害。而活性氧是许多生物过程包括细胞生长,细胞周期、细胞程序化死亡、激素信号转导、细胞应答过程中的重要物质。
植物受到机械损伤时,植物体内的茉莉酸及其衍生物的含量显著增加,而增加的茉莉酸又诱导PLD及PLA的表达,进而激活JA合成过程中的一系列酶,在这些酶中,PLD, LOX和AOS是三个及其重要的水解或合成酶,因此编码这三种酶的基因在转录水平上的变化对植物体内JA的含量起着重要的调控作用。
我们的研究表明,机械损伤使黄瓜果实中POD和APX的活性明显上升,而SOD和CAT虽然在机械损伤后有所上升,但上升幅度不明显,而四种酶的变化趋势都是先上升后下降；SA对四种酶的活性都有一定程度的抑制作用；乙烯在损伤后一定时间内对POD和APX的活性有一定的促进作用,损伤后内对CAT的活性几乎没有影响,而对SOD的活性则有较强烈的抑制作用；MeJA对损伤后黄瓜果实中的SOD、POD在损伤后的前几个小时有微弱的抑制作用,而对CAT、APX的活性则几乎没有影响。
我们用同源克隆技术克隆出黄瓜PLD基因的全长。黄瓜PLD基因cDNA全长为2499 bp,其最大编码框为2427 bp,其DNA全长为3754 bp,其中含2个内含子；可能编码含有808个氨基酸的蛋白质,氨基酸序列具有PLD特有的C2结构域；与其它物种PLD同源基因氨基酸序列的比较和构建的分子进化树发现,黄瓜PLD基因属于PLD基因家族的α亚家族。
我们的半定量和实时荧光定量PCR结果发现,机械损伤后的黄瓜果实中CsPLDa、LOX, AOS基因的表达量都有一定程度的上调,SA对损伤后的黄瓜果实中CsPLDa基因和LOX基因的表达都有微弱的抑制作用,而对AOS基因的表达有明显的抑制作用；乙烯利前处理的损伤黄瓜中CsPLDa基因和LOX基因的表达有明显的上调作用,而对AOS基因的表达则没有明显的影响；MeJA对损伤后黄瓜果实中CsPLDa基因和AOS基因的表达量都有明显的促进作用,而对LOX基因的表达则有明显的抑制作用。
包括生物胁迫因子如病原体、昆虫、杂草等,物理胁迫因子如低氧、低温、干旱、物理损伤、机械胁迫等,化学胁迫因子如盐、金属离子、营养元素等各种不良环境因子都能引起植物细胞的程序性死亡。我们的实验结果表明,2℃低温处理9天后,黄瓜果实的乙烯释放量、呼吸速率、电导率明显上升,硬度下降,细胞核变形,染色质开始降解,DNA降解明显,20℃放置2天后乙烯释放量、呼吸速率、电导率上升更加明显,硬度快速下降,DNA降解更加明显,大多数细胞的细胞核降解,形成没有细胞核的空腔,细胞的程序性死亡特征更加明显；而1-MCP和EGTA处理后的黄瓜果实乙烯释放量、呼吸速率、电导率明显比对照低,硬度下降幅度较小,颜色变化较小,细胞核变形数量较少,细胞核降解后形成的没有空腔的细胞数目也相对减少,DNA降解幅度明显小于低温处理黄瓜果实DNA的降解,表明1-MCP和EGTA处理能减缓低温诱导的黄瓜果实发生PCD且1-MCP处理的效果好于EGTA.
Mechanical wounding is an environmental stress which stimulates plants to produce the necessary signals to prepare further defenses. Oxidative burst is a common early response of plants to elicitor treatments, which induces the enhancement of plant antioxidative enzyme activity to scavenge the generated active oxide species. Wound-induced responses activate those genes involved in the production of wound-signals which then switch-on genes involved in systemic defense against stress. JA signaling can be induced by a range of abiotic stresses, including mechanical wounding. Phospholipase D (PLD), lipoxygenase (LOX) and allene oxide synthase (AOS) are key enzymes which involved in the JA synthesis.
This study was to investigate the effects of salicylic acid, ethephon, or methyl jasmonate on antioxidant system and on the transcription levels of wounded-induced PLDα, LOX, AOS genes in postharvest cucumber (Cucumis sativus L. cv. Biyu-3) fruit.
We identified the CsPLD gene with homology cloning. Potential orthologous PLD amino acid sequences were collected based on BLAST-P homology searches using the CsPLD gene sequence in the NCBI database. Multiple sequences alignments were performed with the ClustalX programme. A phylogenetic tree was constructed applying the neighbour-joining (NJ) method. We found that the maxisum open reading frame (ORF) of the CsPLD gene cDNA was 2,427 nt-long and encoded a predicted protein of 808 amino acids with a C2 domain related to the C2 superfamily. Phylogenetic analysis result indicated that CsPLD gene belonged to the a sub-family of PLD genes family.
We found that mechanical wounding stimulated the activities of SOD, POD, APX and CAT, activities of POD and APX distinctly increased, however, activities of SOD and CAT both had slightly increase. The activities of all the four antioxidative enzymes were increased to a maxisim value and then declined. SA pre-treated inhibited activities of SOD, POD, CAT and APX in wounded cucumber. Ethephon pre-treated could slightly enhanced activites of POD and APX in several hours after wounded, but the pre-treated had no effect on activity of CAT and intensively inhibited the activity of SOD in wounded fruits. The activities of SOD and POD were slightly inhibited a couple of hours after wounding, but there were nearly no changes on activities of APX and CAT in wounded cucumber.
The relative levels of expression of the PLDa gene induced by wounding following SA, ETH, or MeJA pre-treatment were detected by semi-quantitative PCR and quantitative real-time PCR analysis. Wounding increased the levels of expression of the PLDa, LOX and AOS genes. SA pre-treated slightly surpressed the relative levels of expression of PLDa and LOX genes, but strongly surpressed the relative level of expression of AOS gene. Transcript levels of PLDa and LOX genes apparently increased but transcript level of AOS gene had no change in ETh pre-treated cucumber fruits. MeJA treatment activated expression of the PLDa and AOS gene, and the stimulatory effect of ETH was greater than that of MeJA, however, this treatment surpressed the transcript level of LOX gene.
Programmed cell death (PCD) is an essential physiological process occurring during plant development and in response to biotic and abiotic stresses such as pathogen, insects feeding, low oxygen concentration, chilling, draought, mechanical wounding, osmosis stress, metallic ion and so on. Our results showed that ethylene production rate, respiration rate and electrolyte leakage distinctly increased in cucumber with 9 days at 2℃chilling treatment, the hardness declined and the color tended to become yellow in cucumber after chilling treatment, DNA degraded and nucleus became deformed; when the chilling fruits were put at 20℃for 2 days, ethylene production rate, respiration rate and electrolyte leakage increased more evidently, firmness declined more quickly and the color became yellowish green; the degradation of DNA in the treatment fruits were remarkable; most of the chromatin condensed and the nucleus disappeared; the cells of chilling treatment fruits presented apparent characteristics of PCD. Ethylene production rate, respiration rate and electrolyte leakage in chilling treatment cucumber fruits with 1-MCP and EGTA pre-treated for 24 h increased slower than those in chilling treatment, the firmness declined also slowly and the color almost green in pre-treated fruits; DNA of the pre-treated fruits had a little bit degradation and the number of cell shrinkage and nucleus deformation distinctly reduced than that in chilling fruits; those indicated that 1-MCP and EGTA pre-treated could relieve the occurrence of PCD in chilling treatment cucumber fruits and the effects of 1-MCP pre-treatment was better than EGTA pre-treatment.
引文
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