采前采后处理对葡萄柚果实贮藏品质的影响
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摘要
我国是柑橘的主要原产地之一,其中柚类生产面积及产量均位居世界前列。由于贮藏保鲜技术的相对落后,柚果实采后质地品质劣变,导致果实易感染腐烂、发生生理病害,从而降低果实品质和商品价值。因此,采后果实质地品质已成为当前柚类果实贮藏保鲜研究的热点问题之一。本试验以“里约红”葡萄柚(Rio Red Grapefruit)为试材,研究贮藏期果实细胞壁物质降解、细胞壁降解酶活性及其基因表达,探讨采前矿质元素(钙、硼)、植物激素(赤霉素、水杨酸)及采后钙、热处理对贮藏期果实质地软化的调控效应及作用机理,为葡萄柚果实贮藏技术方法改进和贮藏期质地品质调控提供理论依据。论文主要研究结果如下：
(1)生长发育期喷施钙、硼肥处理能够使贮藏过程中葡萄柚果实维持较高的硬度,抑制果实失重率的上升,提高采收时果实可溶性固形物含量,但对贮藏期果实可溶性固形物含量增加没有显著影响。采前喷施钙、硼肥处理可不同程度减小果实AIR的损失,减缓果实WSP.ESP和4KSF含量上升,并使果实SCSP.24KSF含量保持在较高水平。此外,采前钙、硼处理可有效抑制贮藏果实PG.PME.Cx.β-Gal和α-L-Af细胞壁水解酶活性上升；其中,采前钙、硼处理对α-L-Af调控效果在贮藏前期明显,对PME的调控效果在中期明显,对PG.Cx的调控效果在中、后期明显,而对p-Gal的调控效果在整个贮藏阶段均明显。同时,葡萄柚果实细胞壁降解酶基因表达明显受采前钙、硼处理调控抑制,果实PG、PME和Cx基因表达趋势与酶活性变化基本一致,而α-L-Af、β-Gαl基因表达先于酶活；其中,PG.α-L-Af主要于贮藏前期受明显抑制,Cx在中、后期抑制,PME在后期受Ca调控基因表达水平最低,而β-Gal在整个贮藏时期受抑制。综上所述,采前喷施钙、硼肥对贮藏葡萄柚果实软化具有重要的调控作用,其调控作用在贮藏中、后期效果显著；三种采前钙、硼处理方法中,Ca+B处理调控效果最突出,B处理调控效果次之,Ca处理调控效果最弱。
(2)生长期喷施赤霉素、水杨酸处理有效减缓贮藏期葡萄柚果实硬度下降,抑制果实失重率上升,但对果实可溶性固形物含量变化没有明显的影响。赤霉素、水杨酸处理可有效地减缓贮藏期果实SCSP的降解,降低果实WSP.ESP含量,减小果实24KSF的损失,降低4KSF含量,抑制果实细胞壁物质果胶、半纤维素降解；赤霉素、水杨酸处理对细胞壁物降解抑制效果在贮藏前期较明显。同时,赤霉素、水杨酸处理还抑制果实细胞壁水解酶活性上升；其中,果实PG活性在贮藏前期受调控作用显著,PME活性在贮藏中期受调控效果明显,Cx.α-L-Af和p-Gal活性则在贮藏中、后期受调控作用明显。此外,不同处理对细胞壁水解酶调控效果差异明显,GA3+SA对PG.Cx.α-L-Af和β-Gal调控效果显著,GA3对PME调控效果明显。葡萄柚果实贮藏期间PG、Gx、α-L-Af和β-Gal基因表达趋势与酶活性变化基本一致,而PME基因表达先于酶活性上升；赤霉素、水杨酸处理对PG基因表达的抑制主要于贮藏前期,对PME基因表达抑制主要在贮藏前、中期；而对Cx.α-L-Af和β-Gαl基因表达抑制则在后期受明显。其中,PG.α-L-Af和β-Gal基因表达受GA3+SA调控作用较明显,GA3对PME.Cx基因表达的调控效果强于其它处理。对外源激素处理后果实特性指标的研究表明,采前喷施赤霉素、水杨酸能够通过抑制细胞壁水解酶基因表达及酶活性,减缓细胞壁物质降解代谢,从而保持贮藏期葡萄柚果实的质地品质；赤霉素、水杨酸处理在贮藏中、后期对葡萄柚果实品质的调控效果显著；GA3+SA复合处理对葡萄柚果实品质的调控效果最突出,GA3处理调控效果次之,SA处理调控效果最弱。
(3)采后浸钙、热处理均可以显著抑制贮藏期葡萄柚果实硬度下降,减缓果实失重率上升,但对可溶性固形物含量无明显影响。采后钙、热处理对葡萄柚果实细胞壁组分WSP、ESP的调控作用主要表现在贮藏中、后期,而对SCSP.4KSF.24KSF的调控作用贯穿于整个贮藏阶段；钙、热处理对各种细胞壁降解酶活性的调控抑制时期不相同,PME.Cx和α-L-Af在贮藏前、中期受调控作用明显,PG.β-Gal在贮藏中、后期受调控作用显著。贮藏期果实PG.α-L-Af和β-Gαl基因表达趋势与酶活性变化基本一致,而PME、Cx基因表达与酶活性变化存在时空差异；PG、PME和β-Gal基因表达主要于贮藏中、后期受明显抑制,而Cx和α-L-Af基因表达则在贮藏初、末期受有效抑制。综合研究认为,采后钙、热处理均能够明显减缓贮藏期葡萄柚果实细胞壁降解酶活性增加及其基因表达上调,从而降低果实细胞壁果胶、半纤维素降解,维持果实硬度,提高果实贮藏质地品质；在贮藏中、后期,钙、热处理对葡萄柚果实调控作用显著；三种采后钙、热处理方式中,Ca+50℃复合处理对果实调控效果效果最突出,Ca处理效果次之,50℃处理效果最弱。
China is one of the important origin area of citrus. The planting acreage and yield of pomelo fruits are top the list of the world. However, the study of the storage and fresh-keeping techniques are limited, and this backward make pomelo fruits vulnerable to decay infection, physiological disease, and thus, the fruits quality was easy to decrease after harvest. Therefore, more and more attentions were put into developing the storage and fresh-keeping technologies for pomelo fruits. In this study, the effects and mechanisms of pre-harvest foliar application of mineral element (Ca, B), plant hormones (GA3, SA), and postharvest Ca, hot treatments to control fruit texture quality during storage were investigated. Meanwhile, the cell wall degradation, cell wall enzyme activities, and cell wall degradation enzymes gene expression of the treated and untreated'Rio Red'grapefruit fruits were comparatively analyzed after harvest. These technologies and their mechanisms resulted from the study may provide theoretical foundation in grapefruit fruit postharvest conservation. The main results obtained from the study are listed as follows:
(1) Pre-harvest applications of calcium, boron alone or in combination could maintain fruits firmness, restrain fruits weight loss, and improve fruits SSC content during postharvest storage period. The treatments retarded the increase of WSP, ESP, and4KSF in cell wall of grapefruit fruits, and restained the degradation of cell wall materials, including AIR, SCSP, and24KSF. In addition, calcium and/or boron treatments inhibited activities cell wall enzymes effectively in different storage period (a-L-Af, early period; PME, middle period; PG and Cx, middle-later period;(3-Gal, all storage stage). Meanwhile, gene expression levels were significantly regulated by pre-harvest application of calcium and/or boron treatments in grapefruit fruits. The change tendencies of PG, PME, and Cx enzyme expression accorded with their corresponding enzyme activity, while expression levels of a-L-Af and β-Gal enzymes preceded their enzyme activity changes. The expression levels of PG、 a-L-Af enzymes were regulated significantly in earlier storage stage and the expression levels of Cx were regulated in middle-later storage stage. The PME exhibited lowest transcript levels in later storage stage with Ca treatment, while the expression level of β-Gal was restrained by calcium and/or boron treatment during whole storage period. In conclusion, applications of calcium, boron alone or in combination provided significant regulation effect in grapefruit fruit texture quality, and the most desirable effect were detected in middle-later period of postharvest storage. Calcium and boron combined treatment showed the best regulating effect on grapefruit quality, followed by boron treatment, and calcium application showed the weakest effect than the others.
(2) Pre-harvest applications of gibberellin, salicylic acid alone or in combination could maintain fruits firmness and restrain fruits weight loss during postharvest storage period. However, the treatments did not showed an obvious influence on SSC of grapefruit fruits. The degradation process of cell wall material, including pectin and hemicelluloses, was restained by gibberellin and/or salicylic acid treatment, and so the increase speeds of WSP, ESP, and4KSF contents, and the decrease speeds of SCSP and24KSF contents were reduced by the technical treatments. The activities of cell wall hydrolytic enzymes were inhibited by gibberellin and/or salicylic acid applications in different postharvest storage stages (PG, early period; PME, middle period; Cx, a-L-Af, and β-Gal, middle-later period). In addition, notable regulation differences were observed between different treatment techniques, and the activities of PG, Cx, a-L-Af, and β-Gal enzymes were regulated notably by gibberellin and salicylic acid combined treatment, while the activity of PME was obviously controlled by GA3treatment. The expression levels of PG, Cx, a-L-Af and β-Gal accorded with their corresponding enzyme activity changes, but the expression levels of PME preceded its'enzyme activity change. The expression levels of PG were significantly regulated in early storage stage, PME was regulated in earlier-middle storage stage, and Cx, a-L-Af, and β-Gal were regulated in later storage stage. Furthermore, the gene expression of PG, a-L-Af and β-Gal were controlled effectively by gibberellin and salicylic acid combined treatment, and the gene expression of PME, Cx were obviously regulated by gibberellin treatment. The above results indicated that pre-harvest application of gibberellin, salicylic acid alone or in combination inhibited cell wall gene expressions and cell wall degradation enzyme activities, and the treatments retarded cell wall material degradation. The notable inhibition effect of the treatments was observed in middle-later storage stage. Gibberellin and salicylic acid combined treatment showed the best regulating effect, followed by gibberellin treatment, and the salicylic acid application showed inferior effect than the others. These effects are beneficial to improve grapefruit fruit texture quality.
(3) Postharvest applications calcium, hot alone or in combination restrained firmness decrease, and retarded weight loss increase, but showed no desired effect in regulation SSC content during storage. The cell wall material WSP, ESP contents of grapefruit fruits were significantly regulated in middle-later storage stage, and changes of SCSP,4KSP,24KSP contents were inhibited during whole storage period by postharvest calcium and/or hot treatments. The activities of cell wall hydrolytic enzymes were regulated by calcium and/or hot treatments in different postharvest storage stages (PME, Cx, and a-L-Af, early-middle period; PME, middle period; PG and β-Gal, middle-later period). In addition, the tendencies of gene expression levels of PG, a-L-Af, β-Gal in grapefruit fruits accorded with their corresponding enzyme activist changes, while PME, Cx gene expressions exhibited time-differences with the their enzyme activist changes. The expressions of PG, PME, β-Gal genes were suppressed obviously by calcium and/or hot treatments in middle-later stage, while Cx and a-L-Af gene expressions were regulated notably in early and later stages during fruit storage. Based on the above results, it could be concluded that the calcium and/or hot treatments significantly inhibited fruit cell wall hydrolytic enzyme activities and gene expressions increase, and retarded pectin and hemicelluloses degradation. The remarkable regulation effects of the technical treatments were observed in middle-later storage stage, and it is helpful to maintain an ideal texture quality of grapefruit fruits in postharvest storage. Calcium and hot combined treatment showed the best regulating effect on grapefruit quality, followed by calcium treatment, and hot application showed inferior effect than the others.
(1)生长发育期喷施钙、硼肥处理能够使贮藏过程中葡萄柚果实维持较高的硬度,抑制果实失重率的上升,提高采收时果实可溶性固形物含量,但对贮藏期果实可溶性固形物含量增加没有显著影响。采前喷施钙、硼肥处理可不同程度减小果实AIR的损失,减缓果实WSP.ESP和4KSF含量上升,并使果实SCSP.24KSF含量保持在较高水平。此外,采前钙、硼处理可有效抑制贮藏果实PG.PME.Cx.β-Gal和α-L-Af细胞壁水解酶活性上升；其中,采前钙、硼处理对α-L-Af调控效果在贮藏前期明显,对PME的调控效果在中期明显,对PG.Cx的调控效果在中、后期明显,而对p-Gal的调控效果在整个贮藏阶段均明显。同时,葡萄柚果实细胞壁降解酶基因表达明显受采前钙、硼处理调控抑制,果实PG、PME和Cx基因表达趋势与酶活性变化基本一致,而α-L-Af、β-Gαl基因表达先于酶活；其中,PG.α-L-Af主要于贮藏前期受明显抑制,Cx在中、后期抑制,PME在后期受Ca调控基因表达水平最低,而β-Gal在整个贮藏时期受抑制。综上所述,采前喷施钙、硼肥对贮藏葡萄柚果实软化具有重要的调控作用,其调控作用在贮藏中、后期效果显著；三种采前钙、硼处理方法中,Ca+B处理调控效果最突出,B处理调控效果次之,Ca处理调控效果最弱。
(2)生长期喷施赤霉素、水杨酸处理有效减缓贮藏期葡萄柚果实硬度下降,抑制果实失重率上升,但对果实可溶性固形物含量变化没有明显的影响。赤霉素、水杨酸处理可有效地减缓贮藏期果实SCSP的降解,降低果实WSP.ESP含量,减小果实24KSF的损失,降低4KSF含量,抑制果实细胞壁物质果胶、半纤维素降解；赤霉素、水杨酸处理对细胞壁物降解抑制效果在贮藏前期较明显。同时,赤霉素、水杨酸处理还抑制果实细胞壁水解酶活性上升；其中,果实PG活性在贮藏前期受调控作用显著,PME活性在贮藏中期受调控效果明显,Cx.α-L-Af和p-Gal活性则在贮藏中、后期受调控作用明显。此外,不同处理对细胞壁水解酶调控效果差异明显,GA3+SA对PG.Cx.α-L-Af和β-Gal调控效果显著,GA3对PME调控效果明显。葡萄柚果实贮藏期间PG、Gx、α-L-Af和β-Gal基因表达趋势与酶活性变化基本一致,而PME基因表达先于酶活性上升；赤霉素、水杨酸处理对PG基因表达的抑制主要于贮藏前期,对PME基因表达抑制主要在贮藏前、中期；而对Cx.α-L-Af和β-Gαl基因表达抑制则在后期受明显。其中,PG.α-L-Af和β-Gal基因表达受GA3+SA调控作用较明显,GA3对PME.Cx基因表达的调控效果强于其它处理。对外源激素处理后果实特性指标的研究表明,采前喷施赤霉素、水杨酸能够通过抑制细胞壁水解酶基因表达及酶活性,减缓细胞壁物质降解代谢,从而保持贮藏期葡萄柚果实的质地品质；赤霉素、水杨酸处理在贮藏中、后期对葡萄柚果实品质的调控效果显著；GA3+SA复合处理对葡萄柚果实品质的调控效果最突出,GA3处理调控效果次之,SA处理调控效果最弱。
(3)采后浸钙、热处理均可以显著抑制贮藏期葡萄柚果实硬度下降,减缓果实失重率上升,但对可溶性固形物含量无明显影响。采后钙、热处理对葡萄柚果实细胞壁组分WSP、ESP的调控作用主要表现在贮藏中、后期,而对SCSP.4KSF.24KSF的调控作用贯穿于整个贮藏阶段；钙、热处理对各种细胞壁降解酶活性的调控抑制时期不相同,PME.Cx和α-L-Af在贮藏前、中期受调控作用明显,PG.β-Gal在贮藏中、后期受调控作用显著。贮藏期果实PG.α-L-Af和β-Gαl基因表达趋势与酶活性变化基本一致,而PME、Cx基因表达与酶活性变化存在时空差异；PG、PME和β-Gal基因表达主要于贮藏中、后期受明显抑制,而Cx和α-L-Af基因表达则在贮藏初、末期受有效抑制。综合研究认为,采后钙、热处理均能够明显减缓贮藏期葡萄柚果实细胞壁降解酶活性增加及其基因表达上调,从而降低果实细胞壁果胶、半纤维素降解,维持果实硬度,提高果实贮藏质地品质；在贮藏中、后期,钙、热处理对葡萄柚果实调控作用显著；三种采后钙、热处理方式中,Ca+50℃复合处理对果实调控效果效果最突出,Ca处理效果次之,50℃处理效果最弱。
China is one of the important origin area of citrus. The planting acreage and yield of pomelo fruits are top the list of the world. However, the study of the storage and fresh-keeping techniques are limited, and this backward make pomelo fruits vulnerable to decay infection, physiological disease, and thus, the fruits quality was easy to decrease after harvest. Therefore, more and more attentions were put into developing the storage and fresh-keeping technologies for pomelo fruits. In this study, the effects and mechanisms of pre-harvest foliar application of mineral element (Ca, B), plant hormones (GA3, SA), and postharvest Ca, hot treatments to control fruit texture quality during storage were investigated. Meanwhile, the cell wall degradation, cell wall enzyme activities, and cell wall degradation enzymes gene expression of the treated and untreated'Rio Red'grapefruit fruits were comparatively analyzed after harvest. These technologies and their mechanisms resulted from the study may provide theoretical foundation in grapefruit fruit postharvest conservation. The main results obtained from the study are listed as follows:
(1) Pre-harvest applications of calcium, boron alone or in combination could maintain fruits firmness, restrain fruits weight loss, and improve fruits SSC content during postharvest storage period. The treatments retarded the increase of WSP, ESP, and4KSF in cell wall of grapefruit fruits, and restained the degradation of cell wall materials, including AIR, SCSP, and24KSF. In addition, calcium and/or boron treatments inhibited activities cell wall enzymes effectively in different storage period (a-L-Af, early period; PME, middle period; PG and Cx, middle-later period;(3-Gal, all storage stage). Meanwhile, gene expression levels were significantly regulated by pre-harvest application of calcium and/or boron treatments in grapefruit fruits. The change tendencies of PG, PME, and Cx enzyme expression accorded with their corresponding enzyme activity, while expression levels of a-L-Af and β-Gal enzymes preceded their enzyme activity changes. The expression levels of PG、 a-L-Af enzymes were regulated significantly in earlier storage stage and the expression levels of Cx were regulated in middle-later storage stage. The PME exhibited lowest transcript levels in later storage stage with Ca treatment, while the expression level of β-Gal was restrained by calcium and/or boron treatment during whole storage period. In conclusion, applications of calcium, boron alone or in combination provided significant regulation effect in grapefruit fruit texture quality, and the most desirable effect were detected in middle-later period of postharvest storage. Calcium and boron combined treatment showed the best regulating effect on grapefruit quality, followed by boron treatment, and calcium application showed the weakest effect than the others.
(2) Pre-harvest applications of gibberellin, salicylic acid alone or in combination could maintain fruits firmness and restrain fruits weight loss during postharvest storage period. However, the treatments did not showed an obvious influence on SSC of grapefruit fruits. The degradation process of cell wall material, including pectin and hemicelluloses, was restained by gibberellin and/or salicylic acid treatment, and so the increase speeds of WSP, ESP, and4KSF contents, and the decrease speeds of SCSP and24KSF contents were reduced by the technical treatments. The activities of cell wall hydrolytic enzymes were inhibited by gibberellin and/or salicylic acid applications in different postharvest storage stages (PG, early period; PME, middle period; Cx, a-L-Af, and β-Gal, middle-later period). In addition, notable regulation differences were observed between different treatment techniques, and the activities of PG, Cx, a-L-Af, and β-Gal enzymes were regulated notably by gibberellin and salicylic acid combined treatment, while the activity of PME was obviously controlled by GA3treatment. The expression levels of PG, Cx, a-L-Af and β-Gal accorded with their corresponding enzyme activity changes, but the expression levels of PME preceded its'enzyme activity change. The expression levels of PG were significantly regulated in early storage stage, PME was regulated in earlier-middle storage stage, and Cx, a-L-Af, and β-Gal were regulated in later storage stage. Furthermore, the gene expression of PG, a-L-Af and β-Gal were controlled effectively by gibberellin and salicylic acid combined treatment, and the gene expression of PME, Cx were obviously regulated by gibberellin treatment. The above results indicated that pre-harvest application of gibberellin, salicylic acid alone or in combination inhibited cell wall gene expressions and cell wall degradation enzyme activities, and the treatments retarded cell wall material degradation. The notable inhibition effect of the treatments was observed in middle-later storage stage. Gibberellin and salicylic acid combined treatment showed the best regulating effect, followed by gibberellin treatment, and the salicylic acid application showed inferior effect than the others. These effects are beneficial to improve grapefruit fruit texture quality.
(3) Postharvest applications calcium, hot alone or in combination restrained firmness decrease, and retarded weight loss increase, but showed no desired effect in regulation SSC content during storage. The cell wall material WSP, ESP contents of grapefruit fruits were significantly regulated in middle-later storage stage, and changes of SCSP,4KSP,24KSP contents were inhibited during whole storage period by postharvest calcium and/or hot treatments. The activities of cell wall hydrolytic enzymes were regulated by calcium and/or hot treatments in different postharvest storage stages (PME, Cx, and a-L-Af, early-middle period; PME, middle period; PG and β-Gal, middle-later period). In addition, the tendencies of gene expression levels of PG, a-L-Af, β-Gal in grapefruit fruits accorded with their corresponding enzyme activist changes, while PME, Cx gene expressions exhibited time-differences with the their enzyme activist changes. The expressions of PG, PME, β-Gal genes were suppressed obviously by calcium and/or hot treatments in middle-later stage, while Cx and a-L-Af gene expressions were regulated notably in early and later stages during fruit storage. Based on the above results, it could be concluded that the calcium and/or hot treatments significantly inhibited fruit cell wall hydrolytic enzyme activities and gene expressions increase, and retarded pectin and hemicelluloses degradation. The remarkable regulation effects of the technical treatments were observed in middle-later storage stage, and it is helpful to maintain an ideal texture quality of grapefruit fruits in postharvest storage. Calcium and hot combined treatment showed the best regulating effect on grapefruit quality, followed by calcium treatment, and hot application showed inferior effect than the others.
引文
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