不同溶质型桃果实成熟软化机理研究
摘要
本研究以软溶质型桃果实‘雨花三号’和硬溶质型桃果实‘加纳岩’为材料,从形态学、生理学和分子生物学水平研究了两种不同溶质型桃果实成熟软化过程中的基本生理变化、果肉细胞壁超微结构的变化、细胞壁多糖降解特性以及细胞壁多糖降解相关的重要酶在桃果实成熟软化过程中的作用,为进一步研究核果类果实成熟软化机理提供理论依据。主要研究结果如下：
1.对两种溶质型桃果实成熟软化过程中果实硬度、内源乙烯释放量及果肉细胞壁超微结构进行了分析。结果表明：‘雨花三号’桃果实树体成熟及采后贮藏过程中随着果实硬度下降,呼吸速率和乙烯释放量明显增加并出现峰值；但‘加纳岩’桃果实在树体成熟过程中一直保持相对较高的硬度,采后贮藏过程中‘加纳岩’在贮藏末期硬度有所下降。‘加纳岩’桃果实在树体成熟及采后贮藏过程中乙烯释放量很低,其最高值不及‘雨花三号’桃果实乙烯释放量的1%。桃果实成熟软化过程中对果肉细胞壁超微结构观察的结果表明,与‘加纳岩’桃果实相比,‘雨花三号’桃果实树体成熟过程中果肉细胞壁降解较早被启动；‘雨花三号’在采后贮藏过程前期细胞壁结构发生显著变化。
2.对细胞壁多糖分析结果表明：两种不同溶质型桃果实在树体上的成熟软化过程中,CDTA-1溶性果胶含量都呈现上升趋势,而Na2CO3-1溶性果胶、Na2CO3-2溶性果胶、KOH.1溶性半纤维素多糖、KOH-2溶性半纤维素多糖和CWM-残渣纤维素多糖含量在两种品种中总体都呈现下降趋势,但在软溶质型‘雨花三号’桃果实成熟软化过程中的下降速度相对‘加纳岩’桃果实中更快。说明桃果实成熟过程中果胶分子发生了降解,共价紧密结合型果胶转变为离子可溶性果胶,半纤维素和纤维素多糖含量也出现下降,细胞壁多糖的降解参与了果实成熟软化进程,但细胞壁多糖的降解速度在不同溶质型桃果实中有明显差异。
3.采用高效液相色谱法测定细胞壁多糖组份分子量分布变化结果表明：CDTA、 Na2CO3和KOH组份多糖的分子量在两种桃果实成熟软化过程中都发生下降,说明细胞壁多糖在桃果实成熟过程中发生了解聚,高分子量物质向低分子量物质转变。但两种溶质型桃在多糖组份分子量的大小,各分子量组份含量及出现解聚的时间都明显不同,说明多糖组份的解聚方式对桃果实软化过程有一定影响。
4.采用气相色谱法对两种溶质型桃果实中细胞壁多糖的单糖组成分析结果表明：桃果实成熟软化过程中,果胶多糖解聚,细胞壁富含半乳糖醛酸的果胶主链断裂,多聚半乳糖醛酸降解,支链阿拉伯糖、半乳糖解离,半纤维素多糖中也伴随着阿拉伯糖、半乳糖等中性糖的不同程度的解离。不同的是软溶质型桃果实‘雨花三号’果胶主链的降解主要在成熟前期,而硬溶质型桃果实‘加纳岩’的降解主要发生在成熟后期,且支链半乳糖的解离主要在软化前期,而支链阿拉伯糖的解离主要在快速软化期。
5.分析了桃果实成熟软化过程中,细胞壁多糖降解相关糖苷酶的基因表达和活性变化。结果表明,β-半乳糖苷酶基因表达量和活性在桃果实成熟软化前期较高,表明其与桃果实成熟软化启动密切相关。α-L-阿拉伯呋哺糖苷酶基因表达和活性在成熟软化后期较高,与桃果实的快速软化期密切相关,且α-L-阿拉伯呋哺糖苷酶的快速变化期稍滞后于乙烯及其合成相关酶的变化,表明该酶的激活与乙烯释放有一定的相关性,且其对果实成熟中后期快速软化影响更为显著。桃果实成熟过程中α-L-阿拉伯呋哺糖苷酶蛋白的积累和活性与a-L-阿拉伯呋哺糖苷酶基因的表达以及果实成熟软化进程具有一定的相关性。进一步证明α-L-阿拉伯呋哺糖苷酶对桃果实软化过程的必要性。
6.1-MCP有效延缓了‘雨花三号’和‘加纳岩’桃果实硬度的下降,抑制了内源乙烯的产生及呼吸速率的增加,使得采后贮藏过程中的细胞壁降解速度延缓。但1-MCP对两个不同品种的显著抑制期有一定差异,对软溶质型桃‘雨花三号’的显著抑制效应主要表现在贮藏前期和中期,而对硬溶质型桃‘加纳岩’的抑制效应主要表现在贮藏后期。1-MCP处理后α-L-阿拉伯呋哺糖苷酶基因、蛋白表达和酶活性受到抑制,果实软化延迟,进一步证明α-L-阿拉伯呋哺糖苷酶作用于细胞壁的降解受乙烯调控。
Fruit softening is a significant event during ripening and has been widely accepted to be the result of the alteration of cell wall components. Studies on two peach cultivars:'Yuhuasanhao'and'Jianayan'showed differences in softening behavior. Each cultivar showed a unique relationship between softening, ethylene production and respiration rate. The changes of physiological, the cell wall ultrastructure in the flesh cells, the degradation characteristics of cell wall polysaccharides as well as correlative glycosidaes for cell wall polysaccharides degration during peach fruit ripening and softening were studied, providing some theoretical basis for further study the role of cell wall changes in stone hard fruit ripening and softening. The main results were as follows:
The modification of firmness, ethylene production and the ultrastructural change in the flesh cells were determined in present study. The results preliminary showed that, respiration rate and ethylene production increased significantly in'Yuhuasanhao'accompanied by the decline of firmness, while relatively higher firmness had been remained during'Jianayan'peach fruit ripening and the firmness decreased little in the end of postharvest storage. The ethylene production in'Jianayan'peach fruit was less than1%of that in'Yuhuasanhao'peach fruit. The results of cell wall ultrastructure showed that the cell wall degradation in 'Yuhuasanhao'peach fruit launched earlier than'Jianayan'peach fruit. A significant change in 'Yuhuasanhao'cell wall was observed in the early stage of postharvest storage.
The results for cell wall polysaccharides showed that the content of CDTA-1pectin polysaccharide increased while the content of Na2CO3-1, Na2CO3-2pectin polysaccharide, KOH-1and KOH-2hemicellulosic polysaccharides and CWM-residue polysaccharide decreased during peach ripening and softening. However, the rate of decline in'Yuhuasanhao'was faster than in'Jianayan'. That may contribute to the process of fruit softening, indicating that pectin experienced degradation and tightly covalently bound pectin transformed into ion bound pectin. However, the degradation rates of cell wall polysaccharides differed significantly in the two different types of peach fruit.
High performance liquid chromatography was used to determine molecular weight distribution in cell wall polysaccharides. The results showed that the molecular mass distributions of CDTA, Na2CCO3and KOH fractions exhibited downshift tendency in both of two types of peach fruit, indicating that cell wall polysaccharides occurred depolymerization and high molecular weight materials changed to the low molecular weight substances during peach fruit ripening. But the molecular weight, the content and the time of depolymerization in polysaccharide components between the two cultivars showed a significant difference, indicating that the pattern of depolymerization had some influence on the peach fruit softening.
Monosaccharide compositions of cell wall polysaccharides were determined by gas chromatography. The results showed that pectic polysaccharides depolymerized, the pectic main chains that riched in galacturonic acid took rupture along with the degradation of galacturonic acid residues and the loss of arabinosyl and galactosyl residues from pectic side chains. The loss of arabinosyl and galactosyl residues also showed in hemicellulosic polysaccharides. The difference between the two cultivars was the initial time of degradation which means that the melting-flesh peach fruit'Yuhuasanhao'was in the front of the ripening while the non-melting-flesh fruit'Jianayan'was in the later of the ripening. And branched-chain galactose degraded mainly in the period of pre-softening, while branched-chain arabinose degraded mainly in the period of rapid softening
Changes of correlative glycosidase for the degradation of cell wall polysaccharides were analyzed during the ripening of peach fruit. The results showed that P-galactosidase gene expression and activity had a higher level in the early stage of softening and it may contribute to softening initiation. The gene expression and activity of a-L-arabinofuranosidase were higher in the late ripening, which were closely related with fruit fast softening. The change of a-L-arabinofuranosidase slightly lagged behind the rapid changes of ethylene and ethylene synthetic enzymes, indicating that the enzyme activation was closely related with accumulation of the endogenous ethylene and had more remarkably effect on peach mid to later fast-softening. At the same time, a-L-arabinofuranosidase protein accumulation and activity as well as gene expression had a good correlation with fruit softening. This is the further evidence for the effect of a-L-arabinofuranosidase on peach fruit softening.
1-MCP effectively delayed the decline of peach fruit firmness and also delayed the increase of endogenous ethylene production and respiration rate, which made the cell wall of pulp maintain its integrity for a longer time. But the time of1-MCP effect during storage differed to some extent between the two different cultivar. The inhibitory effect of1-MCP on'Yuhuasanhao' was in the early to mid-term storage, while the effect on 'Jianayan'was in the late storage. a-L-Arabinofuranosidase translation, protein accumulation and enzyme activity were inhibited accompanied with the postpone of peach fruit softening. That further proved that the role of a-L-arabinofuranosidase on the degradation of cell wall was regulated by ethylene.
1.对两种溶质型桃果实成熟软化过程中果实硬度、内源乙烯释放量及果肉细胞壁超微结构进行了分析。结果表明：‘雨花三号’桃果实树体成熟及采后贮藏过程中随着果实硬度下降,呼吸速率和乙烯释放量明显增加并出现峰值；但‘加纳岩’桃果实在树体成熟过程中一直保持相对较高的硬度,采后贮藏过程中‘加纳岩’在贮藏末期硬度有所下降。‘加纳岩’桃果实在树体成熟及采后贮藏过程中乙烯释放量很低,其最高值不及‘雨花三号’桃果实乙烯释放量的1%。桃果实成熟软化过程中对果肉细胞壁超微结构观察的结果表明,与‘加纳岩’桃果实相比,‘雨花三号’桃果实树体成熟过程中果肉细胞壁降解较早被启动；‘雨花三号’在采后贮藏过程前期细胞壁结构发生显著变化。
2.对细胞壁多糖分析结果表明：两种不同溶质型桃果实在树体上的成熟软化过程中,CDTA-1溶性果胶含量都呈现上升趋势,而Na2CO3-1溶性果胶、Na2CO3-2溶性果胶、KOH.1溶性半纤维素多糖、KOH-2溶性半纤维素多糖和CWM-残渣纤维素多糖含量在两种品种中总体都呈现下降趋势,但在软溶质型‘雨花三号’桃果实成熟软化过程中的下降速度相对‘加纳岩’桃果实中更快。说明桃果实成熟过程中果胶分子发生了降解,共价紧密结合型果胶转变为离子可溶性果胶,半纤维素和纤维素多糖含量也出现下降,细胞壁多糖的降解参与了果实成熟软化进程,但细胞壁多糖的降解速度在不同溶质型桃果实中有明显差异。
3.采用高效液相色谱法测定细胞壁多糖组份分子量分布变化结果表明：CDTA、 Na2CO3和KOH组份多糖的分子量在两种桃果实成熟软化过程中都发生下降,说明细胞壁多糖在桃果实成熟过程中发生了解聚,高分子量物质向低分子量物质转变。但两种溶质型桃在多糖组份分子量的大小,各分子量组份含量及出现解聚的时间都明显不同,说明多糖组份的解聚方式对桃果实软化过程有一定影响。
4.采用气相色谱法对两种溶质型桃果实中细胞壁多糖的单糖组成分析结果表明：桃果实成熟软化过程中,果胶多糖解聚,细胞壁富含半乳糖醛酸的果胶主链断裂,多聚半乳糖醛酸降解,支链阿拉伯糖、半乳糖解离,半纤维素多糖中也伴随着阿拉伯糖、半乳糖等中性糖的不同程度的解离。不同的是软溶质型桃果实‘雨花三号’果胶主链的降解主要在成熟前期,而硬溶质型桃果实‘加纳岩’的降解主要发生在成熟后期,且支链半乳糖的解离主要在软化前期,而支链阿拉伯糖的解离主要在快速软化期。
5.分析了桃果实成熟软化过程中,细胞壁多糖降解相关糖苷酶的基因表达和活性变化。结果表明,β-半乳糖苷酶基因表达量和活性在桃果实成熟软化前期较高,表明其与桃果实成熟软化启动密切相关。α-L-阿拉伯呋哺糖苷酶基因表达和活性在成熟软化后期较高,与桃果实的快速软化期密切相关,且α-L-阿拉伯呋哺糖苷酶的快速变化期稍滞后于乙烯及其合成相关酶的变化,表明该酶的激活与乙烯释放有一定的相关性,且其对果实成熟中后期快速软化影响更为显著。桃果实成熟过程中α-L-阿拉伯呋哺糖苷酶蛋白的积累和活性与a-L-阿拉伯呋哺糖苷酶基因的表达以及果实成熟软化进程具有一定的相关性。进一步证明α-L-阿拉伯呋哺糖苷酶对桃果实软化过程的必要性。
6.1-MCP有效延缓了‘雨花三号’和‘加纳岩’桃果实硬度的下降,抑制了内源乙烯的产生及呼吸速率的增加,使得采后贮藏过程中的细胞壁降解速度延缓。但1-MCP对两个不同品种的显著抑制期有一定差异,对软溶质型桃‘雨花三号’的显著抑制效应主要表现在贮藏前期和中期,而对硬溶质型桃‘加纳岩’的抑制效应主要表现在贮藏后期。1-MCP处理后α-L-阿拉伯呋哺糖苷酶基因、蛋白表达和酶活性受到抑制,果实软化延迟,进一步证明α-L-阿拉伯呋哺糖苷酶作用于细胞壁的降解受乙烯调控。
Fruit softening is a significant event during ripening and has been widely accepted to be the result of the alteration of cell wall components. Studies on two peach cultivars:'Yuhuasanhao'and'Jianayan'showed differences in softening behavior. Each cultivar showed a unique relationship between softening, ethylene production and respiration rate. The changes of physiological, the cell wall ultrastructure in the flesh cells, the degradation characteristics of cell wall polysaccharides as well as correlative glycosidaes for cell wall polysaccharides degration during peach fruit ripening and softening were studied, providing some theoretical basis for further study the role of cell wall changes in stone hard fruit ripening and softening. The main results were as follows:
The modification of firmness, ethylene production and the ultrastructural change in the flesh cells were determined in present study. The results preliminary showed that, respiration rate and ethylene production increased significantly in'Yuhuasanhao'accompanied by the decline of firmness, while relatively higher firmness had been remained during'Jianayan'peach fruit ripening and the firmness decreased little in the end of postharvest storage. The ethylene production in'Jianayan'peach fruit was less than1%of that in'Yuhuasanhao'peach fruit. The results of cell wall ultrastructure showed that the cell wall degradation in 'Yuhuasanhao'peach fruit launched earlier than'Jianayan'peach fruit. A significant change in 'Yuhuasanhao'cell wall was observed in the early stage of postharvest storage.
The results for cell wall polysaccharides showed that the content of CDTA-1pectin polysaccharide increased while the content of Na2CO3-1, Na2CO3-2pectin polysaccharide, KOH-1and KOH-2hemicellulosic polysaccharides and CWM-residue polysaccharide decreased during peach ripening and softening. However, the rate of decline in'Yuhuasanhao'was faster than in'Jianayan'. That may contribute to the process of fruit softening, indicating that pectin experienced degradation and tightly covalently bound pectin transformed into ion bound pectin. However, the degradation rates of cell wall polysaccharides differed significantly in the two different types of peach fruit.
High performance liquid chromatography was used to determine molecular weight distribution in cell wall polysaccharides. The results showed that the molecular mass distributions of CDTA, Na2CCO3and KOH fractions exhibited downshift tendency in both of two types of peach fruit, indicating that cell wall polysaccharides occurred depolymerization and high molecular weight materials changed to the low molecular weight substances during peach fruit ripening. But the molecular weight, the content and the time of depolymerization in polysaccharide components between the two cultivars showed a significant difference, indicating that the pattern of depolymerization had some influence on the peach fruit softening.
Monosaccharide compositions of cell wall polysaccharides were determined by gas chromatography. The results showed that pectic polysaccharides depolymerized, the pectic main chains that riched in galacturonic acid took rupture along with the degradation of galacturonic acid residues and the loss of arabinosyl and galactosyl residues from pectic side chains. The loss of arabinosyl and galactosyl residues also showed in hemicellulosic polysaccharides. The difference between the two cultivars was the initial time of degradation which means that the melting-flesh peach fruit'Yuhuasanhao'was in the front of the ripening while the non-melting-flesh fruit'Jianayan'was in the later of the ripening. And branched-chain galactose degraded mainly in the period of pre-softening, while branched-chain arabinose degraded mainly in the period of rapid softening
Changes of correlative glycosidase for the degradation of cell wall polysaccharides were analyzed during the ripening of peach fruit. The results showed that P-galactosidase gene expression and activity had a higher level in the early stage of softening and it may contribute to softening initiation. The gene expression and activity of a-L-arabinofuranosidase were higher in the late ripening, which were closely related with fruit fast softening. The change of a-L-arabinofuranosidase slightly lagged behind the rapid changes of ethylene and ethylene synthetic enzymes, indicating that the enzyme activation was closely related with accumulation of the endogenous ethylene and had more remarkably effect on peach mid to later fast-softening. At the same time, a-L-arabinofuranosidase protein accumulation and activity as well as gene expression had a good correlation with fruit softening. This is the further evidence for the effect of a-L-arabinofuranosidase on peach fruit softening.
1-MCP effectively delayed the decline of peach fruit firmness and also delayed the increase of endogenous ethylene production and respiration rate, which made the cell wall of pulp maintain its integrity for a longer time. But the time of1-MCP effect during storage differed to some extent between the two different cultivar. The inhibitory effect of1-MCP on'Yuhuasanhao' was in the early to mid-term storage, while the effect on 'Jianayan'was in the late storage. a-L-Arabinofuranosidase translation, protein accumulation and enzyme activity were inhibited accompanied with the postpone of peach fruit softening. That further proved that the role of a-L-arabinofuranosidase on the degradation of cell wall was regulated by ethylene.
引文
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