库尔勒香梨采后贮藏中果皮锈斑产生机制的研究
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摘要
库尔勒香梨是新疆名优特水果之一，以其特有的色、香、味等品质而深受国内外广大消费者青睐。库尔勒香梨果皮锈斑在采前很少发生，而在采后贮藏期间极易发生，严重影响其外观，大大降低了商品价值。目前，国内外对梨果实采前锈斑形成原因及其防治技术研究较多，而对其采后贮藏锈斑发生原因及其机制的研究未见报道。本课题在对库尔勒香梨主产区锈斑发病规律统计分析的基础上，分析了引起果实锈斑病发生的因素，进行了诱发因素与锈斑病形成的相关性研究，并对锈斑特征成分进行了分析鉴定，提出了库尔勒香梨采后贮藏过程中果皮锈斑的产生机制。主要结果如下：
1．从2008年至2010年，以规模化贮藏的库尔勒香梨为研究对象，对其锈斑发病抽样调查研究发现，在库尔勒地区，气调库贮藏的库尔勒香梨锈斑发病率比普通冷库高0.23%～1.32%，气调库中的发病时间比普通冷库高早5d；在阿克苏地区，气调库贮藏的库尔勒香梨锈斑发病率比普通冷库高0.34%～1.55%，气调库中的发病时间比普通冷库高早10d；在普通冷藏中，阿克苏地区库尔勒香梨锈斑发病率比库尔勒地区高0.13%～0.57%；在气调贮藏中，阿克苏地区库尔勒香梨锈斑发病率比库尔勒地区高0.24%～1.11%。阿克苏地区锈斑发病时间比库尔勒地区提前5d发病。在普通冷库及气调库贮藏中，不同位置的库尔勒香梨锈斑发病率由高到低依次为：加湿口处＞冷库中部＞离加湿口最远处。这两种贮藏条件下库尔勒香梨锈斑发病统计分析表明，气调库贮藏的库尔勒香梨锈斑发病率显著高于普通冷库(p<0.01)；阿克苏地区库尔勒香梨锈斑发病率显著高于库尔勒地区(p<0.01)。普通冷库贮藏条件下，贮藏180d后需要防止锈斑病的发生；气调贮藏条件下，174d后锈斑发病率急剧上升。统计调查结果显示，库尔勒香梨锈斑病首先出现在果实萼部，逐渐向胴部、梗部延伸。实验室贮藏研究表明，库尔勒香梨采后贮藏环境的温度、湿度、气体成分与其果皮锈斑形成密切相关。
2．库尔勒香梨在采后贮藏期间，随着锈斑面积的不断扩大，库尔勒香梨锈斑部位和完好部位的呼吸强度逐渐降低，前者降低速度大于后者；乙烯释放量逐渐升高，锈斑部位升高幅度大，完好部位平缓上升；细胞膜透性和丙二醛含量不断升高，锈斑部位升高幅度大于完好部位；酚类物质的含量不断减少，锈斑部位的减少速度大于完好部位；多酚氧化酶活性和不断增强，锈斑部位的酶活性大于完好部位；冷藏条件下的库尔勒香梨的苯丙氨酸解氨酶活性不断增强，气调贮藏条件下的苯丙氨酸解氨酶活性先上升后下降，锈斑部位的酶活性大于完好部位；木质素含量不断增加，锈斑部位含量高于完好部位；锈斑部位纤维素含量不断上升，完好部位略有下降。提出库尔勒香梨采后贮藏中果皮锈斑形成的生理生化机制：库尔勒香梨受不良贮藏环境的刺激，表皮蜡质和角质破裂，表皮细胞结构破坏，细胞内酚类物质和多酚氧化酶的作用发生酶促褐变，同时木栓层形成，锈斑物质逐渐积累，导致果实发生锈斑病。
3．库尔勒香梨表皮细胞显微结构研究表明，锈斑部位的木质素含量高于完好部位；木质素首先沿细胞壁方向延伸，然后垂直于细胞壁面伸展，细胞木质化程度越高，锈斑面积越大。同时，锈斑部位的细胞膜和细胞壁逐渐消失，线粒体、内质网、高尔基体等细胞器逐渐解体，黑色物质沉积增多，直到填满整个细胞；完好部位的细胞膜和细胞壁完整，细胞内没有物质流失。随着库尔勒香梨表皮细胞结构破坏，胞间层木质素增多，促进了愈伤组织的形成，愈伤组织进一步木栓化，形成了锈斑。
4．通过HPLC、UV光谱扫描及LC-MS对库尔勒香梨表皮锈斑特征成分分析得出：锈斑物质的主要化学成分为大黄酸、芦荟大黄素、大黄酚和大黄素等蒽醌类物质，进一步确定了库尔勒香梨锈斑形成过程中发生了酶促褐变反应。
通过本课题研究，提出了库尔勒香梨锈斑的产生机制：库尔勒香梨在采后贮藏过程中，受贮藏环境温度、湿度、CO2浓度的不良刺激，加速了果实衰老，使果实表皮细胞的蜡质和角质破裂，液泡膜内陷并降解，失去了分室作用，液泡内的一系列水解酶外渗并分散到整个细胞中，细胞产生自溶作用，进而使细胞解体和死亡，细胞内的多酚氧化酶与酚类物质被释放，并发生酶促褐变反应，最终生成了深褐色的蒽醌类物质；同时，库尔勒香梨受到贮藏环境的逆境胁迫时，细胞壁内会发生一系列的抗性反应，表皮下薄壁细胞的细胞壁不断增厚，木栓化后形成木栓层，并不断分生出大量的木栓细胞，其逐渐堆积并突出于果实表皮，从而形成了锈斑。
Korla fragrant pear is the famous special fruit in Xinjiang, it was favored with its uniquecolor, aroma, taste, quality. Korla fragrant pear peel epidermis rust in pre-harvest occurredrarely, however, it was easy to occur during postharvest storage, and its appearance wasseriously influenced, and the value of the goods were greatly reduced. At present, moreresearch on the rust formation reasons before pear harvest and prevention technologies. It hasnot been reported on rust occurrence reason and its mechanism during postharvest storage.The purpose of this work was to analyze factors causing fruit rust spot disease and researchedthe correlation between factors and formation of rust on the basis of extensive investigationand statistic analysis of discipline on pear fruit rust incidence in the main producing areas ofKorla fragrant pear. Then characteristic components of rust were analyzed and identified. TheKorla fragrant pear fruit rust generation mechanism during postharvest storage was alsopreliminarily explored. The main conclusions as follows;
1. From2008to2010, the Korla Pear Rust incidence of sampling was surveyed on massstorage of Korla fragrant pear in enterprises. It was found that the Korla Fragrant Pear rustincidence in controlled atmosphere storage (0.23%) was higher than that of in the commoncold storage(1.32%) in Korla area. The incidence time was5days before the latter; In Akesuarea, the Korla Fragrant Pear rust incidence in controlled atmosphere storage (0.34%)washigher than that of in the common cold storage(1.55%) The incidence time was10daysbefore the latter. In common cold storage, pear rust incidence ratio of was0.13%to0.57%higher than that of in Korla area. In controlled atmosphere storage area, Akesu Korla PearRust incidence ratio of0.24%to1.11%higher that of in Korla area. In Akesu area, rustdisease incidence was ahead of5days earlier than in Korla area. In common clod storage andcontrolled atmosphere storage, the fruit rust incidence rate was not same with the differentposition. It was highest in humidifying mouth, then the central section of cold room, finallythe most distant from the humidifying export. The two storage conditions of Korla FragrantPear shows from rust incidence statistics analysis, it was shown Korla Fragrant Pear Rustincidence was significantly higher(p<0.01) in controlled atmosphere storage than that of incommon storage; Akesu area of Korla Fragrant Pear rust incidence rates were significantlyhigher(p<0.01) than that of Korla area. In common cold storage conditions, the rust diseasemust be prevented after storage for180d. In controlled atmosphere storage conditions, the rustdisease sharply raised after storage for174d. It was shown from the statistical survey, KorlaFragrant Pear rust disease first appeared in fruit calyx, then gradually extended to the carcassand the stem of the pear fruit. The laboratory studies showed that the fruit peel rust formationwas closely related to environment temperature, humidity, gas composition during storage.
2. Korla fragrant pear rust and respiratory strength were gradually reduced with rust areaexpanding during the storage of Korla Fragrant Pear, and the former speed was lower than thelatter. Meanwhile, the ethylene release rate gradually increased, Rust spot range was higher,the sound parts was gentle rising; cell membrane permeability and MDA content increased,the elevated range of rust was lager than the sound parts parts; the content of phenolic substances content decrease gradually, rusty part decrease faster than the sound parts;Polyphenol oxidase activity strengthened gradually, rust site of the enzyme activity wasgreater than the sound parts. Under common cold storage, phenylalanine ammonia-lyaseactivity of Korla pear strengthened. In controlled atmosphere storage, phenylalanineammonia-lyase activity increased then decreased, the enzyme activity in rust site was greaterthan that of in the sound parts; Lignin content increased, it was higher in rust site than in thesound parts; cellulose content raised in rust spot, and dropped in the sound parts. Therefore,after postharvest, physiological and biochemical mechanism of Korla pear fruit rust formationmay be Korla fragrant pear were stimulated by adverse storage environment, and epicuticularwax and cuticle ruptured, and the epidermal cell structure damage, and intracellular phenolsand the polyphenol oxidase enzymatic browned, and at the same time, the cork layed, rustsubstances accumulated gradually, then led to the occurrence of fruit rust disease.
3. It was also shown in Korla fragrant pear epidermal microstructure studies, rust spotlignin content was higher than the sound parts, and lignin directed along the cell wall, andthen was perpendicular to the cell wall extension. The higher the Cell lignification degree andthe larger rust area. Meanwhile rust site on the cell membrane and cell wall graduallydisappeared,,and the mitochondria, endoplasmic reticulum, golgi apparatus and otherorganelles disintegrated, deposition of black material increased, and until it filled the entirecell; the cell membrane and cell wall of intact part was completed, there was no material lossin intracellular. With the Korla fragrant pear epidermal structure damaged, the middle lamellalignin increased, the callus formation was promoted, callus further corky, finally the rust wasformed.
4. Korla fragrant pear skin rust characteristic components were analysed by HPLC (Highpressure liquid chromatography), UV(Ultraviolet spectrophotometer), LC-MS(Liquidchromatography and mass spectrometry) methods. The results were rust material mainchemical compositions were Rhein, Aloe Emodin, Emodin, chrysophanol and anthraquinones.It was confirmed deeply that the enzymatic browning reaction occurred during the Korla pearrust formation.
Through this research, Korla pear rust generation mechanism was put forward, KorlaFragrant Pear were influenced by storage temperature, humidity, concentration of CO2adverse stimulus during postharvest storage. It accelerated senescence of fruit, the fruitepidermis wax and cuticle of ruptured, vacuole membrane invaginated and degradated, roomfunction was lost, a series of hydrolase extravasated within vacuoles and dispersed throughoutthe cell. Cell autolysis developed, the cell was disintegration and death, Intracellular phenolicsand polyphenol oxidase interacted, enzymatic browning was produced, dark brownanthraquinones generated; at the same time, the Korla Pear were Storage stressed by thestorage environment, cell wall would produce a series of resistance, and the epidermis thincell wall of continuous was thickening, corky was happened after the formation of cork layer,and continued to produce large quantities of cork cell division, the gradual accumulation andprominent on the fruit epidermis, thus Korla fragrant pear rust was formed.
1．从2008年至2010年，以规模化贮藏的库尔勒香梨为研究对象，对其锈斑发病抽样调查研究发现，在库尔勒地区，气调库贮藏的库尔勒香梨锈斑发病率比普通冷库高0.23%～1.32%，气调库中的发病时间比普通冷库高早5d；在阿克苏地区，气调库贮藏的库尔勒香梨锈斑发病率比普通冷库高0.34%～1.55%，气调库中的发病时间比普通冷库高早10d；在普通冷藏中，阿克苏地区库尔勒香梨锈斑发病率比库尔勒地区高0.13%～0.57%；在气调贮藏中，阿克苏地区库尔勒香梨锈斑发病率比库尔勒地区高0.24%～1.11%。阿克苏地区锈斑发病时间比库尔勒地区提前5d发病。在普通冷库及气调库贮藏中，不同位置的库尔勒香梨锈斑发病率由高到低依次为：加湿口处＞冷库中部＞离加湿口最远处。这两种贮藏条件下库尔勒香梨锈斑发病统计分析表明，气调库贮藏的库尔勒香梨锈斑发病率显著高于普通冷库(p<0.01)；阿克苏地区库尔勒香梨锈斑发病率显著高于库尔勒地区(p<0.01)。普通冷库贮藏条件下，贮藏180d后需要防止锈斑病的发生；气调贮藏条件下，174d后锈斑发病率急剧上升。统计调查结果显示，库尔勒香梨锈斑病首先出现在果实萼部，逐渐向胴部、梗部延伸。实验室贮藏研究表明，库尔勒香梨采后贮藏环境的温度、湿度、气体成分与其果皮锈斑形成密切相关。
2．库尔勒香梨在采后贮藏期间，随着锈斑面积的不断扩大，库尔勒香梨锈斑部位和完好部位的呼吸强度逐渐降低，前者降低速度大于后者；乙烯释放量逐渐升高，锈斑部位升高幅度大，完好部位平缓上升；细胞膜透性和丙二醛含量不断升高，锈斑部位升高幅度大于完好部位；酚类物质的含量不断减少，锈斑部位的减少速度大于完好部位；多酚氧化酶活性和不断增强，锈斑部位的酶活性大于完好部位；冷藏条件下的库尔勒香梨的苯丙氨酸解氨酶活性不断增强，气调贮藏条件下的苯丙氨酸解氨酶活性先上升后下降，锈斑部位的酶活性大于完好部位；木质素含量不断增加，锈斑部位含量高于完好部位；锈斑部位纤维素含量不断上升，完好部位略有下降。提出库尔勒香梨采后贮藏中果皮锈斑形成的生理生化机制：库尔勒香梨受不良贮藏环境的刺激，表皮蜡质和角质破裂，表皮细胞结构破坏，细胞内酚类物质和多酚氧化酶的作用发生酶促褐变，同时木栓层形成，锈斑物质逐渐积累，导致果实发生锈斑病。
3．库尔勒香梨表皮细胞显微结构研究表明，锈斑部位的木质素含量高于完好部位；木质素首先沿细胞壁方向延伸，然后垂直于细胞壁面伸展，细胞木质化程度越高，锈斑面积越大。同时，锈斑部位的细胞膜和细胞壁逐渐消失，线粒体、内质网、高尔基体等细胞器逐渐解体，黑色物质沉积增多，直到填满整个细胞；完好部位的细胞膜和细胞壁完整，细胞内没有物质流失。随着库尔勒香梨表皮细胞结构破坏，胞间层木质素增多，促进了愈伤组织的形成，愈伤组织进一步木栓化，形成了锈斑。
4．通过HPLC、UV光谱扫描及LC-MS对库尔勒香梨表皮锈斑特征成分分析得出：锈斑物质的主要化学成分为大黄酸、芦荟大黄素、大黄酚和大黄素等蒽醌类物质，进一步确定了库尔勒香梨锈斑形成过程中发生了酶促褐变反应。
通过本课题研究，提出了库尔勒香梨锈斑的产生机制：库尔勒香梨在采后贮藏过程中，受贮藏环境温度、湿度、CO2浓度的不良刺激，加速了果实衰老，使果实表皮细胞的蜡质和角质破裂，液泡膜内陷并降解，失去了分室作用，液泡内的一系列水解酶外渗并分散到整个细胞中，细胞产生自溶作用，进而使细胞解体和死亡，细胞内的多酚氧化酶与酚类物质被释放，并发生酶促褐变反应，最终生成了深褐色的蒽醌类物质；同时，库尔勒香梨受到贮藏环境的逆境胁迫时，细胞壁内会发生一系列的抗性反应，表皮下薄壁细胞的细胞壁不断增厚，木栓化后形成木栓层，并不断分生出大量的木栓细胞，其逐渐堆积并突出于果实表皮，从而形成了锈斑。
Korla fragrant pear is the famous special fruit in Xinjiang, it was favored with its uniquecolor, aroma, taste, quality. Korla fragrant pear peel epidermis rust in pre-harvest occurredrarely, however, it was easy to occur during postharvest storage, and its appearance wasseriously influenced, and the value of the goods were greatly reduced. At present, moreresearch on the rust formation reasons before pear harvest and prevention technologies. It hasnot been reported on rust occurrence reason and its mechanism during postharvest storage.The purpose of this work was to analyze factors causing fruit rust spot disease and researchedthe correlation between factors and formation of rust on the basis of extensive investigationand statistic analysis of discipline on pear fruit rust incidence in the main producing areas ofKorla fragrant pear. Then characteristic components of rust were analyzed and identified. TheKorla fragrant pear fruit rust generation mechanism during postharvest storage was alsopreliminarily explored. The main conclusions as follows;
1. From2008to2010, the Korla Pear Rust incidence of sampling was surveyed on massstorage of Korla fragrant pear in enterprises. It was found that the Korla Fragrant Pear rustincidence in controlled atmosphere storage (0.23%) was higher than that of in the commoncold storage(1.32%) in Korla area. The incidence time was5days before the latter; In Akesuarea, the Korla Fragrant Pear rust incidence in controlled atmosphere storage (0.34%)washigher than that of in the common cold storage(1.55%) The incidence time was10daysbefore the latter. In common cold storage, pear rust incidence ratio of was0.13%to0.57%higher than that of in Korla area. In controlled atmosphere storage area, Akesu Korla PearRust incidence ratio of0.24%to1.11%higher that of in Korla area. In Akesu area, rustdisease incidence was ahead of5days earlier than in Korla area. In common clod storage andcontrolled atmosphere storage, the fruit rust incidence rate was not same with the differentposition. It was highest in humidifying mouth, then the central section of cold room, finallythe most distant from the humidifying export. The two storage conditions of Korla FragrantPear shows from rust incidence statistics analysis, it was shown Korla Fragrant Pear Rustincidence was significantly higher(p<0.01) in controlled atmosphere storage than that of incommon storage; Akesu area of Korla Fragrant Pear rust incidence rates were significantlyhigher(p<0.01) than that of Korla area. In common cold storage conditions, the rust diseasemust be prevented after storage for180d. In controlled atmosphere storage conditions, the rustdisease sharply raised after storage for174d. It was shown from the statistical survey, KorlaFragrant Pear rust disease first appeared in fruit calyx, then gradually extended to the carcassand the stem of the pear fruit. The laboratory studies showed that the fruit peel rust formationwas closely related to environment temperature, humidity, gas composition during storage.
2. Korla fragrant pear rust and respiratory strength were gradually reduced with rust areaexpanding during the storage of Korla Fragrant Pear, and the former speed was lower than thelatter. Meanwhile, the ethylene release rate gradually increased, Rust spot range was higher,the sound parts was gentle rising; cell membrane permeability and MDA content increased,the elevated range of rust was lager than the sound parts parts; the content of phenolic substances content decrease gradually, rusty part decrease faster than the sound parts;Polyphenol oxidase activity strengthened gradually, rust site of the enzyme activity wasgreater than the sound parts. Under common cold storage, phenylalanine ammonia-lyaseactivity of Korla pear strengthened. In controlled atmosphere storage, phenylalanineammonia-lyase activity increased then decreased, the enzyme activity in rust site was greaterthan that of in the sound parts; Lignin content increased, it was higher in rust site than in thesound parts; cellulose content raised in rust spot, and dropped in the sound parts. Therefore,after postharvest, physiological and biochemical mechanism of Korla pear fruit rust formationmay be Korla fragrant pear were stimulated by adverse storage environment, and epicuticularwax and cuticle ruptured, and the epidermal cell structure damage, and intracellular phenolsand the polyphenol oxidase enzymatic browned, and at the same time, the cork layed, rustsubstances accumulated gradually, then led to the occurrence of fruit rust disease.
3. It was also shown in Korla fragrant pear epidermal microstructure studies, rust spotlignin content was higher than the sound parts, and lignin directed along the cell wall, andthen was perpendicular to the cell wall extension. The higher the Cell lignification degree andthe larger rust area. Meanwhile rust site on the cell membrane and cell wall graduallydisappeared,,and the mitochondria, endoplasmic reticulum, golgi apparatus and otherorganelles disintegrated, deposition of black material increased, and until it filled the entirecell; the cell membrane and cell wall of intact part was completed, there was no material lossin intracellular. With the Korla fragrant pear epidermal structure damaged, the middle lamellalignin increased, the callus formation was promoted, callus further corky, finally the rust wasformed.
4. Korla fragrant pear skin rust characteristic components were analysed by HPLC (Highpressure liquid chromatography), UV(Ultraviolet spectrophotometer), LC-MS(Liquidchromatography and mass spectrometry) methods. The results were rust material mainchemical compositions were Rhein, Aloe Emodin, Emodin, chrysophanol and anthraquinones.It was confirmed deeply that the enzymatic browning reaction occurred during the Korla pearrust formation.
Through this research, Korla pear rust generation mechanism was put forward, KorlaFragrant Pear were influenced by storage temperature, humidity, concentration of CO2adverse stimulus during postharvest storage. It accelerated senescence of fruit, the fruitepidermis wax and cuticle of ruptured, vacuole membrane invaginated and degradated, roomfunction was lost, a series of hydrolase extravasated within vacuoles and dispersed throughoutthe cell. Cell autolysis developed, the cell was disintegration and death, Intracellular phenolicsand polyphenol oxidase interacted, enzymatic browning was produced, dark brownanthraquinones generated; at the same time, the Korla Pear were Storage stressed by thestorage environment, cell wall would produce a series of resistance, and the epidermis thincell wall of continuous was thickening, corky was happened after the formation of cork layer,and continued to produce large quantities of cork cell division, the gradual accumulation andprominent on the fruit epidermis, thus Korla fragrant pear rust was formed.
引文
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