中国红色砂梨着色过程及其生理变化的研究
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摘要
目前我国南方主栽的梨品种大多属于砂梨(Pyrus Pyrifolia Nakai),果实成熟时常为黄绿色(或绿色、黄色)或褐色。中国红色砂梨果皮红色,为我国独有,主要分布于四川和云南等省。目前,以著名的红色砂梨‘火把梨'为亲本育成的新品种在国内各地被广泛引种栽培。但是,在生产中发现红色砂梨的着色表现不稳定,直接导致了经济效益的下降。本研究选择两个红色砂梨主栽品种‘美人酥'和‘云红梨1号'系统研究了红色砂梨的着色过程及与着色相关的生理变化规律,以期为优质红色砂梨的生产提供理论指导。
自然生长条件下,红色砂梨果实生长曲线呈“S”型。大约在盛花后13周果实中叶绿素含量达到最高值,类胡萝卜素、类黄酮和总酚含量都处于一个较高的水平,PAL和UFGT处于活跃状态,花青苷开始在果皮中快速合成。之后,随着果实的成熟,果皮中叶绿素持续下降,花青苷含量不断积累并在采收时达到最大值,果面色泽由绿色渐变为红色。花青苷积累过程中,UFGT活性一直很高,而PAL活性持续下降。整个发育过程中果实总糖含量持续上升,蔗糖含量在果实着色期上升比较明显。果实中总酸含量呈现“低-高-低”的变化趋势,含量最大的苹果酸变化趋势与总酸一致。
红色砂梨套不透光的果袋,果面光亮度提高(‘美人酥'的L~*=72.61;‘云红梨1号'的L~*=71.34),但果皮中没有叶绿素和花青苷的合成,果面呈现黄白色。采前3周、2周和1周分别对套袋果进行去袋处理,使果实曝光或留在单层白袋(透光率为80.31%)或红袋(透光率为34.71%)中,观测不同果袋类型、去袋方式和去袋时间对果实着色面积、果皮中花青苷、叶绿素、类胡萝卜素、类黄酮、总酚含量、酶活性和果肉中糖酸含量等的影响。结果表明,不同套袋处理不仅影响了果实的外观品质,而且影响了果实的内在品质。曝光处理的果实外观品质最佳,红色面积最大,果皮中花青苷的含量最高,尤其是采前2周或3周处理时‘美人酥'花青苷含量显著高于对照,L~*、C、h~o值优于对照果实。考虑到随着时间的延长叶绿素和类胡萝卜素的不断积累和L~*值的下降,曝光处理以采前2周效果最好。红袋处理的果实,花青苷合成最少,即使采前3周去袋,果面仅有淡淡的红色。白袋处理的‘美人酥'合成花青苷的量有限,但相同处理的‘云红梨1号'能合成较多的花青苷,且随着去袋时间的提前,含量逐渐增多,果实着色面积也扩大。尽管套袋处理的果实总糖含量与对照相比没有显著差异,但果实中果糖、山梨糖醇、葡萄糖和蔗糖占总糖的比例有所不同,最突出的是果实中蔗糖含量增多。套袋处理降低了果实中总酸的含量,使得糖酸比值上升,果实风味得以改善。
为了探讨红色砂梨着色的生理机制,跟踪观测了果实去袋曝光15d内花青苷在果皮中的分布和果皮中各色素含量、相关酶活性以及果肉中糖酸含量的变化,同时还进行了花青苷含量与酶活性、糖酸含量之间的相关性分析。通过切片观察得出红色砂梨着色的解剖学原因是表皮下存在3-4层积聚花青苷的细胞,在整个着色过程中这些细胞数目逐渐增多,细胞红色浓度不断增大。尽管去袋后果皮中叶绿素、类胡萝卜素、类黄酮和总酚的含量都在上升,但花青苷含量上升速度最快,在去袋后10天左右达到高峰。曝光诱导了PAL和UFGT的活性,使得花青苷能够快速合成,相关性分析得出‘美人酥'中花青苷含量分别与PAL(r=0.90919*)和UFGT(r=0.89598**)活性呈显著正相关和极显著正相关;‘云红梨1号'中花青苷含量则分别与PAL(r=0.93841**)和UFGT(r=0.97245**)活性呈极显著正相关。去袋后果实中总糖含量上升,总酸含量下降。各组份的糖或酸与花青苷的相关性分析得到蔗糖含量与花青苷含量呈极显著正相关,相关系数在‘美人酥'和‘云红梨1号'中分别为0.92007和0.99594,苹果酸含量与花青苷含量却呈极显著负相关,相关系数分别为-0.97993和-0.91827。
总之,红色砂梨花青苷的积累伴随着果实的成熟,花青苷的合成需要高强度的光照。去袋后曝光诱导了UFGT等与花青苷合成相关酶的活性,促进花青苷快速积累,在去袋后10天左右达到高峰。着色的解剖学原因是表皮下存在3-4层积聚花青苷的细胞。生产上可以通过在生长期对果实套不透光的果袋,采前10天左右去袋曝光,获得着色最艳和品质最佳的果实。
Chinese sand pear (P. pyrifolia Nakai) is the major cultivated group in south China and the ripen fruit usually appear green, yellow or brown (russet). Recently, some cultivars with red skinned fruit have been discovered in Yunnan Province and southern part of Sichuan Province. As red color meets the aesthetic values and consumer demands, red skinned pears distributed widely throughout China in a short time. Nevertheless, it appeared that red color of pear skin was not uniform in different regions and under different cultivated conditions. Thus, two red Chinese sand pear cultivars 'Meirensu' and 'Yunhongli No.1' were selected to investigate the characteristics of fruit development under natural conditions and effects of fruit bagging treatment on fruit development, especially with a view to fruit coloration.
Under natural conditions, the fruit development showed in an "S" curve. Thirteen weeks after full bloom, limited amount of anthocyanin could be detected in the fruit peel, but the contents of chlorophyll, carotenoid, flavonoid, and total phenolics were high as well as the activities of PAL and UFGT. Besides, the content of malic acid and total organic acids were the highest and the content of soluble sugars increased rapidly. Then, with the fruit growth the content of chlorophyll decreased drastically and the content of carotenoid, flavonoid, and total phenolics and the enzymes activities also decreased but in a limited scale. However, the concentration of anthoycanin increased gradually and thereafter the skin became red from green. At the same time, the ratio of total soluble sugars to organic acids also increased, as the content of fructose and sucrose increased and the content of total organic acids decreased.
Fruit covered with light-impermeable bags showed higher lightness (L* = 72.61 in 'Meirensu' and L* = 71.34 in 'Yunhongli No.1') indicating brighter skin than control. Besides, Fruit bagging promoted degradation of existing chlorophyll and inhibited anthocyanin synthesis entirely in fruit peel and thus the skin looked yellowy. Aiming at effects of fruit bagging treatment on color development and inner qualities of fruit, fruit bags were removed 3, 2 and 1 weeks before harvest and then the fruit were exposed or kept in white bags (80.31% Photo-permeability) or red bags (34.71% Photo-permeability). Results showed that both the visual qualities and the inner qualities were influenced by fruit bagging treatments. Fruit with re-exposed treatment had high anthocyanin and low chlorophyll contents, as well as the high lightness and chroma, which were responsible for the better visual quality than control. Considering the accumulation of chlorophyll and carotenoid, the best date for bag removal was 2 weeks before harvest. Fruit kept in red bag formed few anthocyanin even though the outer layer bag was removed 3 weeks before harvest. Limited anthocyanin production was induced by white bag treatment and was enhanced when treated early. With the same treatment, 'Yunhongli No.1' showed greater potential in anthocyanin formation than that in 'Meirensu'. Although there were no significant differences in soluble sugars between control and treatments, the composition of soluble sugars was changed and the concentration of sucrose was enhanced in both cultivars. Fruit bagging decreased the content of total organic acids in both cultivars.
In order to study the mechanism of coloring, fruit re-exposed were collected at different times of 0, 1, 3, 5, 10, and 15 days after the bag removal and the anatomical structure of skin, the concentrations of pigments, the activities of enzymes and the contents of soluble sugars and organic acids were examined. The correlation analysis was conducted between anthocyanin concentration and enzyme activity, content of sugars and acids. Results of anatomical structure observation showed that the pattern of anthocyanin distribution in red Chinese sand pear involved three to four layers containing red cells where anthocyanin accumulated. Then, both the numbers of red cells and cell pigment concentrations increased and the fruit skin became red. After the bag removal, the contents of chlorophyll, carotenoid, flavonoid, and total phenolics were all accumulated, but the speed of anthocyanin accumulation was faster and thus the skin became redder and redder. With the increasing of anthocyanin content, PAL and UFGT activity also increased and the correlation analysis showed that there were close positive relationship. The increasing of anthocyanin concentration had the closest correlation with the change of sucrose (r = 0.92007** in 'Meirensu' and r = 0.99594** in 'Yunhongli No.1') and malic acid (r = -0.97993** in 'Meirensu' and r = -0.91827** in 'Yunhongli No.1').
Based on the current results, we can conclude that anthocyanin biosynthesis in red Chinese sand pear commences with the fruit maturation and higher light intensity was required in red color development. The pattern of anthocyanin distribution in red Chinese sand pear involved three to four layers containing red cells where anthocyanin accumulated. In order to obtain red Chinese sand pear fruit with attractive appearance, fruit must be covered with light-impermeable bags at the early stage of fruit development and the bag should be removed totally at least 10 days before harvest.
自然生长条件下,红色砂梨果实生长曲线呈“S”型。大约在盛花后13周果实中叶绿素含量达到最高值,类胡萝卜素、类黄酮和总酚含量都处于一个较高的水平,PAL和UFGT处于活跃状态,花青苷开始在果皮中快速合成。之后,随着果实的成熟,果皮中叶绿素持续下降,花青苷含量不断积累并在采收时达到最大值,果面色泽由绿色渐变为红色。花青苷积累过程中,UFGT活性一直很高,而PAL活性持续下降。整个发育过程中果实总糖含量持续上升,蔗糖含量在果实着色期上升比较明显。果实中总酸含量呈现“低-高-低”的变化趋势,含量最大的苹果酸变化趋势与总酸一致。
红色砂梨套不透光的果袋,果面光亮度提高(‘美人酥'的L~*=72.61;‘云红梨1号'的L~*=71.34),但果皮中没有叶绿素和花青苷的合成,果面呈现黄白色。采前3周、2周和1周分别对套袋果进行去袋处理,使果实曝光或留在单层白袋(透光率为80.31%)或红袋(透光率为34.71%)中,观测不同果袋类型、去袋方式和去袋时间对果实着色面积、果皮中花青苷、叶绿素、类胡萝卜素、类黄酮、总酚含量、酶活性和果肉中糖酸含量等的影响。结果表明,不同套袋处理不仅影响了果实的外观品质,而且影响了果实的内在品质。曝光处理的果实外观品质最佳,红色面积最大,果皮中花青苷的含量最高,尤其是采前2周或3周处理时‘美人酥'花青苷含量显著高于对照,L~*、C、h~o值优于对照果实。考虑到随着时间的延长叶绿素和类胡萝卜素的不断积累和L~*值的下降,曝光处理以采前2周效果最好。红袋处理的果实,花青苷合成最少,即使采前3周去袋,果面仅有淡淡的红色。白袋处理的‘美人酥'合成花青苷的量有限,但相同处理的‘云红梨1号'能合成较多的花青苷,且随着去袋时间的提前,含量逐渐增多,果实着色面积也扩大。尽管套袋处理的果实总糖含量与对照相比没有显著差异,但果实中果糖、山梨糖醇、葡萄糖和蔗糖占总糖的比例有所不同,最突出的是果实中蔗糖含量增多。套袋处理降低了果实中总酸的含量,使得糖酸比值上升,果实风味得以改善。
为了探讨红色砂梨着色的生理机制,跟踪观测了果实去袋曝光15d内花青苷在果皮中的分布和果皮中各色素含量、相关酶活性以及果肉中糖酸含量的变化,同时还进行了花青苷含量与酶活性、糖酸含量之间的相关性分析。通过切片观察得出红色砂梨着色的解剖学原因是表皮下存在3-4层积聚花青苷的细胞,在整个着色过程中这些细胞数目逐渐增多,细胞红色浓度不断增大。尽管去袋后果皮中叶绿素、类胡萝卜素、类黄酮和总酚的含量都在上升,但花青苷含量上升速度最快,在去袋后10天左右达到高峰。曝光诱导了PAL和UFGT的活性,使得花青苷能够快速合成,相关性分析得出‘美人酥'中花青苷含量分别与PAL(r=0.90919*)和UFGT(r=0.89598**)活性呈显著正相关和极显著正相关;‘云红梨1号'中花青苷含量则分别与PAL(r=0.93841**)和UFGT(r=0.97245**)活性呈极显著正相关。去袋后果实中总糖含量上升,总酸含量下降。各组份的糖或酸与花青苷的相关性分析得到蔗糖含量与花青苷含量呈极显著正相关,相关系数在‘美人酥'和‘云红梨1号'中分别为0.92007和0.99594,苹果酸含量与花青苷含量却呈极显著负相关,相关系数分别为-0.97993和-0.91827。
总之,红色砂梨花青苷的积累伴随着果实的成熟,花青苷的合成需要高强度的光照。去袋后曝光诱导了UFGT等与花青苷合成相关酶的活性,促进花青苷快速积累,在去袋后10天左右达到高峰。着色的解剖学原因是表皮下存在3-4层积聚花青苷的细胞。生产上可以通过在生长期对果实套不透光的果袋,采前10天左右去袋曝光,获得着色最艳和品质最佳的果实。
Chinese sand pear (P. pyrifolia Nakai) is the major cultivated group in south China and the ripen fruit usually appear green, yellow or brown (russet). Recently, some cultivars with red skinned fruit have been discovered in Yunnan Province and southern part of Sichuan Province. As red color meets the aesthetic values and consumer demands, red skinned pears distributed widely throughout China in a short time. Nevertheless, it appeared that red color of pear skin was not uniform in different regions and under different cultivated conditions. Thus, two red Chinese sand pear cultivars 'Meirensu' and 'Yunhongli No.1' were selected to investigate the characteristics of fruit development under natural conditions and effects of fruit bagging treatment on fruit development, especially with a view to fruit coloration.
Under natural conditions, the fruit development showed in an "S" curve. Thirteen weeks after full bloom, limited amount of anthocyanin could be detected in the fruit peel, but the contents of chlorophyll, carotenoid, flavonoid, and total phenolics were high as well as the activities of PAL and UFGT. Besides, the content of malic acid and total organic acids were the highest and the content of soluble sugars increased rapidly. Then, with the fruit growth the content of chlorophyll decreased drastically and the content of carotenoid, flavonoid, and total phenolics and the enzymes activities also decreased but in a limited scale. However, the concentration of anthoycanin increased gradually and thereafter the skin became red from green. At the same time, the ratio of total soluble sugars to organic acids also increased, as the content of fructose and sucrose increased and the content of total organic acids decreased.
Fruit covered with light-impermeable bags showed higher lightness (L* = 72.61 in 'Meirensu' and L* = 71.34 in 'Yunhongli No.1') indicating brighter skin than control. Besides, Fruit bagging promoted degradation of existing chlorophyll and inhibited anthocyanin synthesis entirely in fruit peel and thus the skin looked yellowy. Aiming at effects of fruit bagging treatment on color development and inner qualities of fruit, fruit bags were removed 3, 2 and 1 weeks before harvest and then the fruit were exposed or kept in white bags (80.31% Photo-permeability) or red bags (34.71% Photo-permeability). Results showed that both the visual qualities and the inner qualities were influenced by fruit bagging treatments. Fruit with re-exposed treatment had high anthocyanin and low chlorophyll contents, as well as the high lightness and chroma, which were responsible for the better visual quality than control. Considering the accumulation of chlorophyll and carotenoid, the best date for bag removal was 2 weeks before harvest. Fruit kept in red bag formed few anthocyanin even though the outer layer bag was removed 3 weeks before harvest. Limited anthocyanin production was induced by white bag treatment and was enhanced when treated early. With the same treatment, 'Yunhongli No.1' showed greater potential in anthocyanin formation than that in 'Meirensu'. Although there were no significant differences in soluble sugars between control and treatments, the composition of soluble sugars was changed and the concentration of sucrose was enhanced in both cultivars. Fruit bagging decreased the content of total organic acids in both cultivars.
In order to study the mechanism of coloring, fruit re-exposed were collected at different times of 0, 1, 3, 5, 10, and 15 days after the bag removal and the anatomical structure of skin, the concentrations of pigments, the activities of enzymes and the contents of soluble sugars and organic acids were examined. The correlation analysis was conducted between anthocyanin concentration and enzyme activity, content of sugars and acids. Results of anatomical structure observation showed that the pattern of anthocyanin distribution in red Chinese sand pear involved three to four layers containing red cells where anthocyanin accumulated. Then, both the numbers of red cells and cell pigment concentrations increased and the fruit skin became red. After the bag removal, the contents of chlorophyll, carotenoid, flavonoid, and total phenolics were all accumulated, but the speed of anthocyanin accumulation was faster and thus the skin became redder and redder. With the increasing of anthocyanin content, PAL and UFGT activity also increased and the correlation analysis showed that there were close positive relationship. The increasing of anthocyanin concentration had the closest correlation with the change of sucrose (r = 0.92007** in 'Meirensu' and r = 0.99594** in 'Yunhongli No.1') and malic acid (r = -0.97993** in 'Meirensu' and r = -0.91827** in 'Yunhongli No.1').
Based on the current results, we can conclude that anthocyanin biosynthesis in red Chinese sand pear commences with the fruit maturation and higher light intensity was required in red color development. The pattern of anthocyanin distribution in red Chinese sand pear involved three to four layers containing red cells where anthocyanin accumulated. In order to obtain red Chinese sand pear fruit with attractive appearance, fruit must be covered with light-impermeable bags at the early stage of fruit development and the bag should be removed totally at least 10 days before harvest.
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