黄瓜果实成熟度与耐冷性的关系及其生理机制研究
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摘要
果蔬采收成熟度影响其采后品质和对逆境的响应,所以是农业生产和农产品开发的焦点。本文以黄瓜为材料,研究其采收成熟度对采后耐冷性的影响,其对黄瓜作物农业生产指导和果实利用开发有极大的意义。
黄瓜果实呈单S型生长曲线,根据果实大小和颜色可将黄瓜果实分为未熟(3-8DAA)、成熟(9-16DAA)、转色(17-22DAA)和黄色(35-40DAA)四个成熟度。黄瓜果实成熟度可以明显影响其采后耐冷性。处于早期发育阶段的果实更易发生冷害,而成熟度高的果实有更高的耐冷性。因此,在商品允许范围内,收获成熟度较高的黄瓜可以减轻采收冷害,延长保鲜期。
氧化胁迫随着黄瓜果实的发育进程不断加剧。生成H2O2的SOD活性显著升高,而清除H2O2的CAT和APX酶活性下降,导致黄瓜果实发育过程中H2O2积累。黄瓜果实成熟过程中GSH含量下降和ASC含量升高说明GSH是促使黄瓜果实形成过程中细胞分裂的重要因素,而ASC则促进后期果实发育过程中细胞膨大。POD基因表达水平和酶活性的显著升高以及ASC含量的升高是黄瓜果实发育和衰老过程中的显著生理变化。
黄瓜果实冷害发生与低温导致的氧化胁迫增加直接相关。SOD可迅速响应低温胁迫,在冷害早期有效清除氧化胁迫,但是在冷害后期其作用逐渐下降。CAT和APX活性在冷害中下降,说明其在冷胁迫诱导的H2O2清除过程中的作用效果减弱。ASC再生循环对维持黄瓜果实冷害下ASC/DHA比值十分关键,成熟度高的黄瓜果实中较高的ASC含量和其还原状态可以阻止活性氧伤害。成熟度高的黄瓜果实中较高POD活性和ASC含量使得其抗氧化能力增强,从而具有较强的耐冷性。
黄瓜果实生长发育过程中总糖和还原糖含量变化不明显,直到黄色期其含量都显著下降。蔗糖、葡萄糖、果糖含量随着果实生长而降低。随着果实生长发育,SS和SPS活性上升,但是SS基因表达水平降低。在黄瓜果实发育期间,有机酸积累,绿原酸和总草酸随着果实的发育和衰老而积累,而单宁酸、没食子酸和水溶性草酸降解。该糖酸结构形成了黄瓜果实特殊风味。
在冷藏过程中所有黄瓜果实中可溶性总糖和还原糖含量稍有上升,但是冷藏结束常温放置后果实中可溶性总糖和还原糖含量明显下降。冷藏期间蔗糖含量在幼嫩的未熟和成熟果实中升高,而在衰老的转色和黄色果实中降低。冷藏结束常温放置后,幼嫩果实蔗糖含量下降,而衰老果实蔗糖含量上升。葡萄糖和果糖含量在果实冷害过程中稍微降低或变化不明显,而冷藏结束常温放置后其在黄色果实中下降,而在其他果实中上升。冷害胁迫下ss和sPs活性在未熟和黄色果实中呈互补关系,而在成熟和转色果实中,ss和sPs活性变化不明显,且都呈上升趋势。ss基因表达水平和其活性在冷害中变化相似。冷害过程中有机酸含量上升。在冷藏中各有机酸含量变化不甚明显,唯有没食子酸含量在未熟果实中明显降低。冷藏结束常温放置后,除了单宁酸含量明显升高,其它各有机酸含量缓慢上升。
黄瓜果实生长及衰老过程中,PG和果胶酶活性变化不明显,只有转色果实中PG和果胶酶活性显著高于其他果实,而黄色果实中果胶酶活性较低。黄瓜果实生长过程中纤维素酶和β-葡萄糖苷酶都是在前期较高,而后期下降。PG、纤维素酶和β-葡萄糖苷酶活性受其基因表达直接影响。黄瓜果实生长过程中几丁质酶活性升高,而β-1,3-葡聚糖酶在成熟期果实中含量较高。几丁质酶和β-1,3-葡聚糖酶基因表达水平都随果实生长而显著提高。黄瓜果实成熟过程中水溶性果胶变化不明显,只有在成熟果实中急剧上升。原果胶在所有果实中变化不明显。
冷藏过程中PG和果胶酶活性呈先下降后上升趋势,黄色果实中果胶酶活性一直呈上升趋势。冷害结束常温放置后,PG活性下降而果胶酶活性稍有上升。在冷藏过程中纤维素酶和β-葡萄糖苷酶活性在未熟和黄色果实中都稍有下降,而在成熟和转色果实中下降而后上升。幼嫩果实中纤维素酶和β-葡萄糖苷酶活性较高。黄瓜果实采后冷害过程中几丁质酶活性增加,同时冷害严重的幼嫩果实中几丁质酶活性较低,而冷害较轻的转色果实几丁质酶活性则较高。黄色果实中几丁质酶变化与其他果实不同,在冷藏中几丁质酶活性降低。冷胁迫下β-1,3-葡聚糖酶活性在幼嫩的未熟和成熟果实中上升,而在衰老的转色和黄色果实中下降,冷藏结束及其后常温放置后幼嫩果实中β-1,3-葡聚糖酶活性高于成熟度较高的果实。黄瓜果实中水溶性果胶含量随冷害的发生而上升。原果胶含量在冷藏过程中降低而随后常温放置后含量上升。
The harvest maturity stage of horticultural product can influence its postharvest quality and stress response, so it is always be the focus of agricultural practice and utilization of product. This research study the effect of fruit maturity on the chilling tolerance of postharvest cucumber and the physiological mechanism involved, the result would greatly benefit the instruction of agricultural cultivation and postharvest fruit utilization of cucumber.
Cucumber fruits exhibit a single-sigmoidal growth curve. Based on fruit size and color, four distinct developmental stages were chosen:Immature (3-8DAA), Mature (9-16DAA), Breaker (17-22DAA) and Yellow (35-40DAA). The chilling tolerance of cucumber fruits was remarkably affected by the fruit maturity. Fruits at earlier developmental stages are more susceptible to chilling stress. So storing cucumber fruits with advanced maturity would alleviate chilling injury and extend post-harvest life at low temperate.
Increased oxidative stress may be an intrinsic feature of the development and senescence of cucumber fruits. The increase in H2O2-generating enzyme (SOD) activity and the decrease in the activities of the two main H2O2-scavenging enzymes (CAT and APX) may cause the accumulation of H2O2during cucumber development and senescence. The decline in GSH content and increase in ASC content during fruit maturation suggest that GSH is important for cell division during the formation of cucumber fruits, whereas ASC is critical for cell expansion during their development and senescence. Significant increases in POD mRNA level, activity, and ASC content are conspicuous features that characterize the development and senescence of cucumber fruits, reflecting the close correlation between POD, ASC, and cucumber fruit maturation.
The development of chilling injury is related to increased oxidative stress under chilling conditions. SOD showed an early response to chilling stress but became inefficient with the development of chilling injury in all fruits. CAT and APX appear less effective in scavenging the H2O2produced under chilling stress. The recycling activity of ASC may be essential to the maintenance of the ASC/DHA ratio, and the higher ASC redox state might prevent oxidative damage in more mature cucumber fruits. Significant higher POD activity and ASC content may provide a higher antioxidant capacity and achieve stronger chilling tolerance for fruits at later developmental stages.
Total sugar content and reduced sugar content showed little change during the development of cucumber fruit, and both significantly declined in Yellow fruit. Sucrose, glucose and fructose content all decreased during the development and senescence of cucumber fruit. The activity of SS and SPS increased during fruit maturation, but the mRNA level of SS declined. The organic acid accumulated during the growth of cucumber fruit, the content of chlorogenic acid and total oxalic acid increased, but the content of tannic acid, gallic acid and water soluble oxalic acid declince. This sugar-acid composition constitutes the special flavor of cucumber fruit.
Total sugar content and reduced sugar content in all fruit slightly increased during cold storage, but both notablely declined after rewarm. Sucrose content increased in Immature and Mature fruit during cold storage and declined after rewarm, but decreased in Breaker and Yellow fruit during cold storage and increased after rewarm. Glucose and fructose content showed little change during cold storage, but increased after rewarm except declined in Yellow fruit. SS and SPS activity both slightly increased in Maturee and Breaker fruit during chilling stress, whereas complement to each other in Immature and Yellow fruit. SS activity may directly influenced by SS transcription level. Organic acid content increased in cucumber fruit under chilling stress. During cold storage analysed organic acid in cucumber fruit showed little change, only gallic acid content declined in Immature fruit. Organic acid content in cucumber fruit increased after rewarm.
During the development and senescence of cucumber fruit PG and pectinase activity showed little change, while both significantly increased in Breaker fruit, and pectinase activity declined remarkablely in Yellow fruit. Both cellulase and β glucosidase activity declined during the growth of cucumber fruit. All PG. cellulase and β-glucosidase activity may directly influenced by gene expression of these enzymes. Chitinase activity increased during fruit maturation,β-1,3-glucanase only had high activity in Mature fruit, and the gene expression level of both enzymes increased during growth. The water soluble pectin and protopectin content showed little change during the development and senescence of cucumber fruit, except Mature fruit had significant high level of water soluble pectin.
PG and pectinase activity declined during cold storage and also increased before the end of cold storage, pectinase activity consistently increased during cold storage. PG activity decreased and pectinase activity slightly increased after rewarm. During cold storage cellulase and β-glucosidase activity slightly decreased in Immature and Yellow fruit, while in Mature and Breaker fruit both declined and then increased till end of cold storage. Fruit at earlier developmental stage have higher cellulase and β-glucosidase activity under chilling stress. Chitinase activity increased in all fruit under chilling stress, and higher chitinase activity appeared in more mature fruit which suffered less chilling injury. Change of chitinase activity inYellow fruit was different with other fruit, which showed declined trend. Under chilling stress,β-1,3glucanase activity increased in Immature and Mature fruit, but declined in Breaker and Yellow fruit, fruit at earlier developmental stage had higher β-1,3-glucanase activity than more mature fruit after rewarm. The water soluble pectin content increased, but protopectin content decreased in all fruit under chilling stress, protopectin content also increased after rewarm.
黄瓜果实呈单S型生长曲线,根据果实大小和颜色可将黄瓜果实分为未熟(3-8DAA)、成熟(9-16DAA)、转色(17-22DAA)和黄色(35-40DAA)四个成熟度。黄瓜果实成熟度可以明显影响其采后耐冷性。处于早期发育阶段的果实更易发生冷害,而成熟度高的果实有更高的耐冷性。因此,在商品允许范围内,收获成熟度较高的黄瓜可以减轻采收冷害,延长保鲜期。
氧化胁迫随着黄瓜果实的发育进程不断加剧。生成H2O2的SOD活性显著升高,而清除H2O2的CAT和APX酶活性下降,导致黄瓜果实发育过程中H2O2积累。黄瓜果实成熟过程中GSH含量下降和ASC含量升高说明GSH是促使黄瓜果实形成过程中细胞分裂的重要因素,而ASC则促进后期果实发育过程中细胞膨大。POD基因表达水平和酶活性的显著升高以及ASC含量的升高是黄瓜果实发育和衰老过程中的显著生理变化。
黄瓜果实冷害发生与低温导致的氧化胁迫增加直接相关。SOD可迅速响应低温胁迫,在冷害早期有效清除氧化胁迫,但是在冷害后期其作用逐渐下降。CAT和APX活性在冷害中下降,说明其在冷胁迫诱导的H2O2清除过程中的作用效果减弱。ASC再生循环对维持黄瓜果实冷害下ASC/DHA比值十分关键,成熟度高的黄瓜果实中较高的ASC含量和其还原状态可以阻止活性氧伤害。成熟度高的黄瓜果实中较高POD活性和ASC含量使得其抗氧化能力增强,从而具有较强的耐冷性。
黄瓜果实生长发育过程中总糖和还原糖含量变化不明显,直到黄色期其含量都显著下降。蔗糖、葡萄糖、果糖含量随着果实生长而降低。随着果实生长发育,SS和SPS活性上升,但是SS基因表达水平降低。在黄瓜果实发育期间,有机酸积累,绿原酸和总草酸随着果实的发育和衰老而积累,而单宁酸、没食子酸和水溶性草酸降解。该糖酸结构形成了黄瓜果实特殊风味。
在冷藏过程中所有黄瓜果实中可溶性总糖和还原糖含量稍有上升,但是冷藏结束常温放置后果实中可溶性总糖和还原糖含量明显下降。冷藏期间蔗糖含量在幼嫩的未熟和成熟果实中升高,而在衰老的转色和黄色果实中降低。冷藏结束常温放置后,幼嫩果实蔗糖含量下降,而衰老果实蔗糖含量上升。葡萄糖和果糖含量在果实冷害过程中稍微降低或变化不明显,而冷藏结束常温放置后其在黄色果实中下降,而在其他果实中上升。冷害胁迫下ss和sPs活性在未熟和黄色果实中呈互补关系,而在成熟和转色果实中,ss和sPs活性变化不明显,且都呈上升趋势。ss基因表达水平和其活性在冷害中变化相似。冷害过程中有机酸含量上升。在冷藏中各有机酸含量变化不甚明显,唯有没食子酸含量在未熟果实中明显降低。冷藏结束常温放置后,除了单宁酸含量明显升高,其它各有机酸含量缓慢上升。
黄瓜果实生长及衰老过程中,PG和果胶酶活性变化不明显,只有转色果实中PG和果胶酶活性显著高于其他果实,而黄色果实中果胶酶活性较低。黄瓜果实生长过程中纤维素酶和β-葡萄糖苷酶都是在前期较高,而后期下降。PG、纤维素酶和β-葡萄糖苷酶活性受其基因表达直接影响。黄瓜果实生长过程中几丁质酶活性升高,而β-1,3-葡聚糖酶在成熟期果实中含量较高。几丁质酶和β-1,3-葡聚糖酶基因表达水平都随果实生长而显著提高。黄瓜果实成熟过程中水溶性果胶变化不明显,只有在成熟果实中急剧上升。原果胶在所有果实中变化不明显。
冷藏过程中PG和果胶酶活性呈先下降后上升趋势,黄色果实中果胶酶活性一直呈上升趋势。冷害结束常温放置后,PG活性下降而果胶酶活性稍有上升。在冷藏过程中纤维素酶和β-葡萄糖苷酶活性在未熟和黄色果实中都稍有下降,而在成熟和转色果实中下降而后上升。幼嫩果实中纤维素酶和β-葡萄糖苷酶活性较高。黄瓜果实采后冷害过程中几丁质酶活性增加,同时冷害严重的幼嫩果实中几丁质酶活性较低,而冷害较轻的转色果实几丁质酶活性则较高。黄色果实中几丁质酶变化与其他果实不同,在冷藏中几丁质酶活性降低。冷胁迫下β-1,3-葡聚糖酶活性在幼嫩的未熟和成熟果实中上升,而在衰老的转色和黄色果实中下降,冷藏结束及其后常温放置后幼嫩果实中β-1,3-葡聚糖酶活性高于成熟度较高的果实。黄瓜果实中水溶性果胶含量随冷害的发生而上升。原果胶含量在冷藏过程中降低而随后常温放置后含量上升。
The harvest maturity stage of horticultural product can influence its postharvest quality and stress response, so it is always be the focus of agricultural practice and utilization of product. This research study the effect of fruit maturity on the chilling tolerance of postharvest cucumber and the physiological mechanism involved, the result would greatly benefit the instruction of agricultural cultivation and postharvest fruit utilization of cucumber.
Cucumber fruits exhibit a single-sigmoidal growth curve. Based on fruit size and color, four distinct developmental stages were chosen:Immature (3-8DAA), Mature (9-16DAA), Breaker (17-22DAA) and Yellow (35-40DAA). The chilling tolerance of cucumber fruits was remarkably affected by the fruit maturity. Fruits at earlier developmental stages are more susceptible to chilling stress. So storing cucumber fruits with advanced maturity would alleviate chilling injury and extend post-harvest life at low temperate.
Increased oxidative stress may be an intrinsic feature of the development and senescence of cucumber fruits. The increase in H2O2-generating enzyme (SOD) activity and the decrease in the activities of the two main H2O2-scavenging enzymes (CAT and APX) may cause the accumulation of H2O2during cucumber development and senescence. The decline in GSH content and increase in ASC content during fruit maturation suggest that GSH is important for cell division during the formation of cucumber fruits, whereas ASC is critical for cell expansion during their development and senescence. Significant increases in POD mRNA level, activity, and ASC content are conspicuous features that characterize the development and senescence of cucumber fruits, reflecting the close correlation between POD, ASC, and cucumber fruit maturation.
The development of chilling injury is related to increased oxidative stress under chilling conditions. SOD showed an early response to chilling stress but became inefficient with the development of chilling injury in all fruits. CAT and APX appear less effective in scavenging the H2O2produced under chilling stress. The recycling activity of ASC may be essential to the maintenance of the ASC/DHA ratio, and the higher ASC redox state might prevent oxidative damage in more mature cucumber fruits. Significant higher POD activity and ASC content may provide a higher antioxidant capacity and achieve stronger chilling tolerance for fruits at later developmental stages.
Total sugar content and reduced sugar content showed little change during the development of cucumber fruit, and both significantly declined in Yellow fruit. Sucrose, glucose and fructose content all decreased during the development and senescence of cucumber fruit. The activity of SS and SPS increased during fruit maturation, but the mRNA level of SS declined. The organic acid accumulated during the growth of cucumber fruit, the content of chlorogenic acid and total oxalic acid increased, but the content of tannic acid, gallic acid and water soluble oxalic acid declince. This sugar-acid composition constitutes the special flavor of cucumber fruit.
Total sugar content and reduced sugar content in all fruit slightly increased during cold storage, but both notablely declined after rewarm. Sucrose content increased in Immature and Mature fruit during cold storage and declined after rewarm, but decreased in Breaker and Yellow fruit during cold storage and increased after rewarm. Glucose and fructose content showed little change during cold storage, but increased after rewarm except declined in Yellow fruit. SS and SPS activity both slightly increased in Maturee and Breaker fruit during chilling stress, whereas complement to each other in Immature and Yellow fruit. SS activity may directly influenced by SS transcription level. Organic acid content increased in cucumber fruit under chilling stress. During cold storage analysed organic acid in cucumber fruit showed little change, only gallic acid content declined in Immature fruit. Organic acid content in cucumber fruit increased after rewarm.
During the development and senescence of cucumber fruit PG and pectinase activity showed little change, while both significantly increased in Breaker fruit, and pectinase activity declined remarkablely in Yellow fruit. Both cellulase and β glucosidase activity declined during the growth of cucumber fruit. All PG. cellulase and β-glucosidase activity may directly influenced by gene expression of these enzymes. Chitinase activity increased during fruit maturation,β-1,3-glucanase only had high activity in Mature fruit, and the gene expression level of both enzymes increased during growth. The water soluble pectin and protopectin content showed little change during the development and senescence of cucumber fruit, except Mature fruit had significant high level of water soluble pectin.
PG and pectinase activity declined during cold storage and also increased before the end of cold storage, pectinase activity consistently increased during cold storage. PG activity decreased and pectinase activity slightly increased after rewarm. During cold storage cellulase and β-glucosidase activity slightly decreased in Immature and Yellow fruit, while in Mature and Breaker fruit both declined and then increased till end of cold storage. Fruit at earlier developmental stage have higher cellulase and β-glucosidase activity under chilling stress. Chitinase activity increased in all fruit under chilling stress, and higher chitinase activity appeared in more mature fruit which suffered less chilling injury. Change of chitinase activity inYellow fruit was different with other fruit, which showed declined trend. Under chilling stress,β-1,3glucanase activity increased in Immature and Mature fruit, but declined in Breaker and Yellow fruit, fruit at earlier developmental stage had higher β-1,3-glucanase activity than more mature fruit after rewarm. The water soluble pectin content increased, but protopectin content decreased in all fruit under chilling stress, protopectin content also increased after rewarm.
引文
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