扁桃内果皮木质化过程中相关酶活性的变化
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摘要
【目的】探讨扁桃内果皮的发育与木质素积累及相关酶活性的关系。【方法】以薄壳‘纸皮’和厚壳‘长石头’两个不同壳厚类型扁桃品种为试材,分析内果皮不同发育时期木质素、纤维素和酚类物质含量及其相关酶活性的变化规律。【结果】两个扁桃木质化过程中木质素和纤维素含量均在积累,酚类物质含量呈现下降趋势,PAL、CAD、C4H、PPO、POD活性呈现先上升后下降趋势;‘长石头’扁桃内果皮木质素和纤维素含量分别在花后55 d和40 d起始终高于‘纸皮’,但内果皮的酚类物质含量始终低于‘纸皮’,‘长石头’扁桃内果皮PAL、CAD、C4H、PPO、POD活性高峰值显著高于‘纸皮’;扁桃内果皮木质素与纤维素含量呈正相关,而与酚类物质含量呈现负相关;PAL、CAD、C4H、PPO、POD活性与木质素快速积累后期呈现负相关。【结论】较高的PAL、CAD、C4H、PPO、POD活性和大量的木质素所需要的前提物质促进了木质素的积累,从而使内果皮木质化加厚。
【Objective】Analysis of the relationship between lignin content in almond endocarp and related enzyme activity in the process of development reveals endocarp lignification and its mechanism.【Methods】This study used thin shelled'Zhipi'and thick shelled'Changshitou'as the experimental materials. Lignin content, cellulose content, polyphenol content and related enzyme activities in almond endocarp were determined in different development periods and the correlations between various parameters were analyzed.【Results】The endocarp development in the thin shelled'Zhipi'and thick shelled'Changshitou'was consistent. Lignin content showed a trend of rising. From 25 to 40 d after full blossom,'Zhipi'and'Changshitou'accumulated lignin slowly. The lignin content in'Zhipi'was higher than in'Changshitou'during this period. Rapid lignin accumulation started at different times. The rapid increase in lignin content in'Changshitou'started around 40 days after full blossom, and it started from 55 d after full blossom in'Zhipi'.'Changshitou'was thus 15 days earlier than'Zhipi'. At 55 d after full blossom, the lignin content in'Zhipi'and'Changshitou'rose rapidly, and in'Changshitou',lignin content was higher than in'Zhipi'. During the endocarp development, cellulose content tended to rise in both varieties. At first, the cellulose content in'Zhipi'was higher than in'Changshitou'.Starting around 35 d after full blossom,'Changshitou'had a higher lignin content than'Zhipi'. The polyphenol content declined constantly. At first, it was higher in'Changshitou', but from 45 d after fullblossom, it declined faster in'Changshitou'than in 'Zhipi'. The change patterns of C4H, CAD and PAL activities were similar. They displayed a downward trend after an initial rise. The enzyme activities in'Zhipi'reached a peak at 55 d after full blossom, while those in'Changshitou'at 70 d after full blossom. From 25 to 70 d after full blossom, there were no significant differences between varieties. At 70 d after full blossom, the enzyme activities in'Changshitou'were higher than in'Zhipi'. The occurrence the enzyme activity peak was coincide with the rapid accumulation of lignin. The study also found that the enzyme activity dropped to certain level, but rapid accumulation of lignin continued. The endocarp of the thick shelled'Changshitou'had constantly significantly higher activities of POD and PPO than that of the thin shelled'Zhipi', indicating that thicker endocarp needs to have higher the POD and PPO activities. In the whole process of endocarp development'Changshitou'always had a higher PPO activity than'Zhipi'. In,'Zhipi'endocarp, lignin content and activity of CAD were significantly correlated,and cellulose content and PPO activity had a significant positive correlation during the early stage. During late period of rapid endocarp growth, lignin content was significantly and negatively correlated to C4H and CAD activities, and cellulose content had significant negative correlations with PPO, PAL,C4H and CAD activities, while polyphenol content and POD activity had a significant positive correlation. In'Changshitou', during the early stage of endocarp development, a significant negative correlation was found between polyphenol and lignin contents; cellulose content was significantly positively correlated to PAL, C4H and CAD activities; and polyphenol content was significantly and negatively correlated to PAL and CAD activities. During the late period of rapid endocarp development, significant negative correlations were found between lignin content and PPO activity and between cellulose content and PPO activity.【Conclusion】Higher activities of PAL, CAD, C4H, PPO and POD as well abundant lignin precursors enables greater accumulation of lignin, which generates a thicker lignified endocarp as shown in the thick shelled'Changshitou'.
【Objective】Analysis of the relationship between lignin content in almond endocarp and related enzyme activity in the process of development reveals endocarp lignification and its mechanism.【Methods】This study used thin shelled'Zhipi'and thick shelled'Changshitou'as the experimental materials. Lignin content, cellulose content, polyphenol content and related enzyme activities in almond endocarp were determined in different development periods and the correlations between various parameters were analyzed.【Results】The endocarp development in the thin shelled'Zhipi'and thick shelled'Changshitou'was consistent. Lignin content showed a trend of rising. From 25 to 40 d after full blossom,'Zhipi'and'Changshitou'accumulated lignin slowly. The lignin content in'Zhipi'was higher than in'Changshitou'during this period. Rapid lignin accumulation started at different times. The rapid increase in lignin content in'Changshitou'started around 40 days after full blossom, and it started from 55 d after full blossom in'Zhipi'.'Changshitou'was thus 15 days earlier than'Zhipi'. At 55 d after full blossom, the lignin content in'Zhipi'and'Changshitou'rose rapidly, and in'Changshitou',lignin content was higher than in'Zhipi'. During the endocarp development, cellulose content tended to rise in both varieties. At first, the cellulose content in'Zhipi'was higher than in'Changshitou'.Starting around 35 d after full blossom,'Changshitou'had a higher lignin content than'Zhipi'. The polyphenol content declined constantly. At first, it was higher in'Changshitou', but from 45 d after fullblossom, it declined faster in'Changshitou'than in 'Zhipi'. The change patterns of C4H, CAD and PAL activities were similar. They displayed a downward trend after an initial rise. The enzyme activities in'Zhipi'reached a peak at 55 d after full blossom, while those in'Changshitou'at 70 d after full blossom. From 25 to 70 d after full blossom, there were no significant differences between varieties. At 70 d after full blossom, the enzyme activities in'Changshitou'were higher than in'Zhipi'. The occurrence the enzyme activity peak was coincide with the rapid accumulation of lignin. The study also found that the enzyme activity dropped to certain level, but rapid accumulation of lignin continued. The endocarp of the thick shelled'Changshitou'had constantly significantly higher activities of POD and PPO than that of the thin shelled'Zhipi', indicating that thicker endocarp needs to have higher the POD and PPO activities. In the whole process of endocarp development'Changshitou'always had a higher PPO activity than'Zhipi'. In,'Zhipi'endocarp, lignin content and activity of CAD were significantly correlated,and cellulose content and PPO activity had a significant positive correlation during the early stage. During late period of rapid endocarp growth, lignin content was significantly and negatively correlated to C4H and CAD activities, and cellulose content had significant negative correlations with PPO, PAL,C4H and CAD activities, while polyphenol content and POD activity had a significant positive correlation. In'Changshitou', during the early stage of endocarp development, a significant negative correlation was found between polyphenol and lignin contents; cellulose content was significantly positively correlated to PAL, C4H and CAD activities; and polyphenol content was significantly and negatively correlated to PAL and CAD activities. During the late period of rapid endocarp development, significant negative correlations were found between lignin content and PPO activity and between cellulose content and PPO activity.【Conclusion】Higher activities of PAL, CAD, C4H, PPO and POD as well abundant lignin precursors enables greater accumulation of lignin, which generates a thicker lignified endocarp as shown in the thick shelled'Changshitou'.
引文
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[3]李夕勃.核桃坚果硬壳发育与木质素代谢关系的研究[D].保定:河北农业大学,2012.LI Xibo.Studies on relationship between lignin metabolism and shell development of Juglans regia L.[D].Baoding:Agricultural University of Hebei,2012.
[4]史梦雅,李阳,张巍,余佳,刘悦萍.桃果实内果皮木质素沉积的动态变化[J].北京农学院学报,2013,28(2):25-28.SHI Mengya,LI Yang,ZHANG Wei,YU Jia,LIU Yueping.The lignin deposition process in endocarp of peach fruit[J].Journal of Beijing University of Agriculture,2013,28(2):25-28.
[5]郑志锋,邹局春,花勃,张宏健,汪蓉.核桃壳化学组分的研究[J].西南林学院学报,2006,26(2):33-36.ZHENG Zhifeng,ZOU Juchun,HUA Bo,ZHANG Hongjian,WANG Rong.Research on the chemical components of walnut shell[J].Journal of Southwest Forestry College,2006,26(2):33-36.
[6]靳丽鑫,陈梦华,王玉莲,张雪梅,李保国.核桃坚果硬壳发育研究进展[J].北方园艺,2015(5):183-187.JIN Lixin,CHEN Menghua,WANG Yulian,ZHANG Xuemei,LI Baoguo.Research progress on walnut shell development[J].Northern Horticulture,2015(5):183-187.
[7]王丹阳.木质素代谢相关酶在梨石细胞合成中的作用[D].南京:南京农业大学,2014.WANG Danyang.The role of lignin metabolism related enzymes in the synthesis of pear stone cell[D].Nanjing:Nanjing Agricultural University,2014.
[8]陶书田,张绍铃,乔勇进,盛宝龙.梨果实发育过程中石细胞团及几种相关酶活性变化的研究[J].果树学报,2004,21(6):516-520.TAO Shutian,ZHANG Shaoling,QIAO Yongjin,SHENG Baolong.Study on sclereids and activities of several related enzymes during the development of pear fruit[J].Journal of Fruit Science,2004,21(6):516-520.
[9]曹爱娟,王有年,葛水莲,杨爱珍,辛阅军,花宝光.桃内果皮木质素沉积过程中酶活性的变化[J].北京农学院学报,2009,24(1):5-8.CAO Aijuan,WANG Younian,GE Shuilian,YANG Aizhen,XIN Yuejun,HUA Baoguang.Change of enzyme activity during the depositive process of lignin in peach endocarp[J].Journal of Beijing University of Agriculture,2009,24(1):5-8.
[10]文菁,赵书岗,王红霞,张志华,李夕勃.核桃内果皮发育期内果皮木质素与相关酶活性的变化[J].园艺学报,2015,42(11):2144-2152.WEN Jing,ZHAO Shugang,WANG Hongxia,ZHANG Zhihua,LI Xibo.Changes of lignin content and its related enzyme activities in endocarp during walnut shell development period[J].Acta Horticulturae Sinica,2015,42(11):2144-2152.
[11]熊素敏,左秀凤,朱永义.稻壳中纤维素、半纤维素和木质素的测定[J].粮食与饲料工业,2005(8):40-41.XIONG Sumin,ZUO Xiufeng,ZHU Yongyi.Determination of cellulose hemi-cellulose and lignin in rice hull[J].Cereal&Feed Industry,2005(8):40-41.
[12]ZHAO S G,WEN J,WANG H X,ZHANG Z H,LI X B.Changes in lignin content and activity of related enzymes in the endocarp during the walnut shell development period[J].Horticultural Plant Journal,2016,2(3):141-146.
[13]王安娜,徐山宝,刘小彦,吴立根.福林法测定猕猴桃多酚含量的研究[J].食品科学,2008,29(7):398-401.WANG Anna,XU Shanbao,LIU Xiaoyan,WU Ligen.Study on determination of content of polyphenol in kiwifruit by folin method[J].Food Science,2008,29(7):398-401.
[14]刘小阳,李玲,高贵珍,蔡永萍,宗梅.光照对砀山酥梨果实发育过程中POD、PAL、PPO酶活性的影响研究[J].激光生物学报,2008,17(3):295-298.LIU Xiaoyang,LI Ling,GAO Guizhen,CAI Yongping,ZONG Mei.Influence of light intensity on POD,PAL and PPO activity of Pyrus bretschneideri cv.Dangshan su fruit in its growth phase[J].The Influence of Laser Biology Journal,2008 17(3):295-298.
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