扁桃蔗糖合成酶对幼果生理脱落的响应研究
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摘要
【目的】为扁桃蔗糖合成酶基因功能与幼果生理落果的关系。【方法】以新疆主栽品种纸皮扁桃生理落果期的正常果和即将脱落果实为试材,采用qRT-PCR法分析SuSy基因的表达模式,测定糖组分浓度和蔗糖合成酶活性。【结果】扁桃果实SuSy基因在不同发育期均有表达,且均在快速落果期(盛花后7～22 d)的高峰期(盛花后17 d)表达量达低值,而后逐渐升高;果实中葡萄糖浓度﹥果糖浓度﹥蔗糖蔗糖浓度,正常果的葡萄糖和果糖浓度均大于即将脱落果且与SuSy分解方向酶活性变化趋势一致,即将脱落果的蔗糖浓度始终大于正常果,即将脱落果中的蔗糖代谢出现异常;在落果高峰期(盛花后17 d),SuSy分解方向酶活性达峰值,之后随着营养竞争缓和而逐步降低,且正常果酶活性始终大于即将脱落果;SuSy合成方向酶活性在正常果中持续走低,直至落果后期库源竞争趋于缓和后缓慢升高,而即将脱落果的合成方向酶活性却在落果高峰期达峰值。【结论】扁桃幼果生理落果期SuSy酶主要在分解方向上起作用,其活性变化异常造成部分果实糖代谢紊乱导致脱落。
【Objective】 To ascertain the relationship between the function of sucrose synthase gene and physiological fruit drop in almond.【Method】In this study, the normal fruit and soon falling fruit of 'Zhipi' almond, a main cultivar in Xinjiang, were used as the test material. The expression pattern of SuSy gene was analyzed by qRT-PCR and the concentration of sugar components and the activity of sucrose synthase were also measured.【Result】The results showed that:(1) SuSy gene expression was observed at all stages of fruit development, the expression level of SuSy was low at the peak period of physiological falling fruits(7-22 days after full bloom), and then gradually increased, indicating that SuSy expression was inhibited by nutritional competition;(2) In almond fruits, the proportion of the sugar components was as follows: glucose > fructose > sucrose. Contents of both glucose and fructose in normal fruits were greater than those in the impending shedding fruits, and the variation trends of these two sugars were consistent with the changes of SuSy enzyme activity in the decomposition direction. However, sucrose content in the impending shedding fruits was always higher than that in the normal fruits, indicating that the sucrose metabolism in the impending shedding fruit was disturbed;(3) During the peak period of fruit drop(17 days after full bloom), SuSy decomposition direction enzyme reached its peak, and then decreased gradually with the relaxation of nutrition competition, and in addition, SuSy activity in normal fruit enzyme was always higher than that in the impending shedding fruits. SuSy activity in synthetic direction continued to decline and begun to increase slowly at the later stage of falling fruits as the competition between sink and source tended to relax. However, in the impending shedding fruits, SuSy activity in synthetic direction reached the highest level at the peak of falling fruits.【Conclusion】It can be concluded that SuSy plays a role in the direction of decomposition during the almond physiological falling stage, and the abnormal change in SuSy activity results in sugar metabolism disorder in some fruits which causes the fruit shedding.
【Objective】 To ascertain the relationship between the function of sucrose synthase gene and physiological fruit drop in almond.【Method】In this study, the normal fruit and soon falling fruit of 'Zhipi' almond, a main cultivar in Xinjiang, were used as the test material. The expression pattern of SuSy gene was analyzed by qRT-PCR and the concentration of sugar components and the activity of sucrose synthase were also measured.【Result】The results showed that:(1) SuSy gene expression was observed at all stages of fruit development, the expression level of SuSy was low at the peak period of physiological falling fruits(7-22 days after full bloom), and then gradually increased, indicating that SuSy expression was inhibited by nutritional competition;(2) In almond fruits, the proportion of the sugar components was as follows: glucose > fructose > sucrose. Contents of both glucose and fructose in normal fruits were greater than those in the impending shedding fruits, and the variation trends of these two sugars were consistent with the changes of SuSy enzyme activity in the decomposition direction. However, sucrose content in the impending shedding fruits was always higher than that in the normal fruits, indicating that the sucrose metabolism in the impending shedding fruit was disturbed;(3) During the peak period of fruit drop(17 days after full bloom), SuSy decomposition direction enzyme reached its peak, and then decreased gradually with the relaxation of nutrition competition, and in addition, SuSy activity in normal fruit enzyme was always higher than that in the impending shedding fruits. SuSy activity in synthetic direction continued to decline and begun to increase slowly at the later stage of falling fruits as the competition between sink and source tended to relax. However, in the impending shedding fruits, SuSy activity in synthetic direction reached the highest level at the peak of falling fruits.【Conclusion】It can be concluded that SuSy plays a role in the direction of decomposition during the almond physiological falling stage, and the abnormal change in SuSy activity results in sugar metabolism disorder in some fruits which causes the fruit shedding.
引文
[1] 李疆,胡芳名,李文胜, 等. 扁桃的栽培及研究概况[J].果树学报,2002,(5):346-350.LI Jiang, HU Fang-ming, LI Wen-sheng, et al. (2002). Advances of Almond Growing and Research [J]. Journal of Science, (5):346-350. (in Chinese)
[2] 刘玉冰, 张腾国, 李新荣,等. 红砂(Reaumuria soongorica)忍耐极度干旱的保护机制:叶片脱落和茎中蔗糖累积[J]. 中国科学, 2006, 36(4):328-333.LIU Yu-bing, ZHANG Teng-guo, LI Xin-rong, et al. (2006). Protective mechanism of desiccation tolerance in Reaumuria soongorica: Leaf abscission and sucrose accumulation in the stem [J]. Science China, 36(4):328-333.
[3] 吕忠恕,王邦锡,盧崇恩, 等. 苹果和梨幼果脱落期中的代谢变化[J].园艺学报,1963, (3):261-270.Lü Zong-su, WANG Bang-xi, LU Chong-en, et al. (1963). Metabolic changes in apple and pear young fruit during shedding [J]. Acta Horticulturae Sinica, (3):261-270. (in Chinese)
[4] Aurelio Gómez-Cadenas, Mehouachi, J. , Tadeo, F. R. , Primo-Millo, E. , & Talon, M. . (2000). Hormonal regulation of fruitlet abscission induced by carbohydrate shortage in citrus. Planta, 210(4): 636-643.
[5] 付崇毅. 日光温室柑橘诱导成花及落果机理研究[D].呼和浩特:内蒙古农业大学博士学论文,2013.FU Chong-yi. (2013). Study on Mechanism of Flower Induction and Fruitlet Abscission of Citrus under Solar Greenhouse[D]. PhD Dissertation. Inner Mongolia Agricultural University, Hohhot. (in Chinese)
[6] 杨子琴,李茂,章笑赟, 等. 饥饿胁迫对龙眼果实脱落及糖代谢的影响[J].果树学报,2011,28(3):428-432.YANG Zi-qin, LI Mao, ZHANG Xiao-yun, et al. (2011). Effects of starvation stress on fruit abscission and sugar metabolism in longan [J]. Journal of Fruit Science, 28(3):428-432. (in Chinese)
[7] Finazzo, S. F., Davenport, T. L., & Schaffer, B. (1994). Partitioning of photoassimilates in avocado (Persea americana Mill.) during flowering and fruit set. Tree Physiology, 14(2): 153-164.
[8] 黄永敬,马培恰,吴文, 等 . 砂糖橘夏梢生长对果实糖代谢及脱落的影响[J].园艺学报,2013,40(10):1 869-1 876.HUANG Yong-jing, MA Pei-qia, WU Wen, et al. (2013). Effects of Summer Shoot Growth on Sugar Metabolism and Abscission of Fruitlet in'Shatangju' Mandarin [J]. Acta Horticulturae Sinica, 40(10):1,869-1,876. (in Chinese)
[9] 曾庆钱,蔡岳文,严振, 等. 化橘红生理落果与若干生理生化指标的关系[J].亚热带植物科学,2008,(3):19-21.ZENG Qing-qian, CAI Yue-wen, YAN Zhen, et al. (2008). Relationship between Physiological Drop and Some Physiological and Biochemical Indexes of Citrus grandis 'Tomentosa' [J]. Subtropical Plant Science, (3):19-21. (in Chinese)
[10] Echeverria, E. (1990). Developmental transition from enzymatic to acid hydrolysis of sucrose in acid limes (Citrus aurantifolia). Plant Physiology, 92(1): 168-171.
[11] Zhou L, Paull R E. (2001).Sucrose Metabolism During Papaya (Carica papaya) Fruit Growth and Ripening. Am Soc Hortic Sci, 126(3): 351-357.
[12] Chengappa, S., Guilleroux, M., Phillips, W., & Shields, R. (1999). Transgenic tomato plants with decreased sucrose synthase are unaltered in starch and sugar accumulation in the fruit. Plant Molecular Biology, 40(2): 213-221.
[13] 卢合全,沈法富,刘凌霄, 等. 植物蔗糖合成酶功能与分子生物学研究进展[J].中国农学通报,2005 (7):34-37, 57.LU He-quan, SHEN Fa-fu, LIU Ling-xiao, et al. (2005). Recent Advances in Study on Plant Sucrose Synthase[J]. Chinese Agricultural Science Bulletin, (7):34-37, 57. (in Chinese)
[14] Marc-André D'Aoust, Yelle, S., & Binh Nguyen-Quoc. (1999). Antisense inhibition of tomato fruit sucrose synthase decreases fruit setting and the sucrose unloading capacity of young fruit. Plant Cell, 11(12): 2,407-2,418.
[14] 张丽丽,刘威生,刘有春, 等.高效液相色谱法测定5个杏品种的糖和酸[J].果树学报,2010,27(1):119-123.ZHANG Li-li, LIU Wei-sheng, LIU You-chun, et al. (2010). Determination of Sugar and Acid in Five Apricot Varieties by High Performance Liquid Chromatography [J]. The Fruit Tree Journal 27(1): 119-123. (In Chinese)
[15] Keller, F., & Ludlow, M. M. (1993). Carbohydrate metabolism in drought-stressed leaves of pigeonpea (cajanus cajan). Journal of Experimental Botany, 44(265):1,351-1,359.
[16] 杨波,车玉红,郭春苗. 扁桃生理落果期不同组织激素浓度的动态变化及其对落果的影响[J].西北植物学报,2015,35(1):118-124.YANG Bo, CHE Yu-hong, GUO Chun-miao. (2015). ynamic changes of hormones in the different tissue of Almond during the physiological fruit drop and its effect on fruit drop [J]. Acta Botanica Boreali-Occidentalia Sinica, 35(1):118-124. (in Chinese)
[17] Miron, D., & Schaffer, A. A. (1991). Sucrose phosphate synthase, sucrose synthase, and invertase activities in developing fruit of lycopersicon esculentum mill. and the sucrose accumulating Lycopersicon hirsutum humb. and bonpl. Plant Physiology, 95(2): 623-627.
[18] 秦翠鲜,桂意云,陈忠良,等.植物蔗糖合成酶基因研究进展[J].分子植物育种,2018,16(12):3 907-3 914.QIN Cui-xiang, GUI Yi-yun, CHEN Zhong-liang, et al. (2018). The Progress of Studies on Sucrose Synthase Genes in Plants [J]. Molecular Plant Breeding, 16(12):3,907-3,914. (in Chinese)
[19] 赵智中,张上隆,徐昌杰,等.蔗糖代谢相关酶在温州蜜柑果实糖积累中的作用[J].园艺学报,2001,(2):112-118.ZHAO Zhi-zhong, ZHANG Shang-long, XU Chang-jie, et al. (2001). The Role of Sucrose Metabolism Related Enzymes in Sugar Accumulation in Citrus Fruits of Wenzhou [J]. Acta Horticulturae Sinica, (2):112-118. (in Chinese)
[20] 龚荣高, 张光伦, 吕秀兰,等. 脐橙果实糖积累与蔗糖代谢相关酶关系的研究[J]. 四川农业大学学报, 2004, 22(1):34-36.GONG Rong-gao, ZHANG Guang-lun, Lü Xiu-lang, et al. (2004). Study on the Relationship between the Sugar Accumulation and Its Metabolizing Enzymes in Navel Orange Fruit [J]. Journal of Sichuan Agricultural University, 22(1):34-36. (in Chinese)
[21] 张春华, 俞明亮, 马瑞娟,等. 桃不同发育时期主要糖类含量和蔗糖合成酶基因表达水平的动态变化[J].江苏农业学报, 2014,(6):1 456-1 463.ZHANG Chun-hua, YU Ming-liang, MA Rui-juan, et al. (2014). Dynamic changes of carbohydrate content and expression level of sucrose synthase genes in peach (Prunus persica) at different developmental stages [J]. Jiangsu Journal of Agricultural Sciences, (6):1,456-1,463. (in Chinese)
[22] Elling, & Lothar. (1995). Effect of metal ions on sucrose synthase from rice grains-a study on enzyme inhibition and enzyme topography. Glycobiology, 5(2): 201-206.
[2] 刘玉冰, 张腾国, 李新荣,等. 红砂(Reaumuria soongorica)忍耐极度干旱的保护机制:叶片脱落和茎中蔗糖累积[J]. 中国科学, 2006, 36(4):328-333.LIU Yu-bing, ZHANG Teng-guo, LI Xin-rong, et al. (2006). Protective mechanism of desiccation tolerance in Reaumuria soongorica: Leaf abscission and sucrose accumulation in the stem [J]. Science China, 36(4):328-333.
[3] 吕忠恕,王邦锡,盧崇恩, 等. 苹果和梨幼果脱落期中的代谢变化[J].园艺学报,1963, (3):261-270.Lü Zong-su, WANG Bang-xi, LU Chong-en, et al. (1963). Metabolic changes in apple and pear young fruit during shedding [J]. Acta Horticulturae Sinica, (3):261-270. (in Chinese)
[4] Aurelio Gómez-Cadenas, Mehouachi, J. , Tadeo, F. R. , Primo-Millo, E. , & Talon, M. . (2000). Hormonal regulation of fruitlet abscission induced by carbohydrate shortage in citrus. Planta, 210(4): 636-643.
[5] 付崇毅. 日光温室柑橘诱导成花及落果机理研究[D].呼和浩特:内蒙古农业大学博士学论文,2013.FU Chong-yi. (2013). Study on Mechanism of Flower Induction and Fruitlet Abscission of Citrus under Solar Greenhouse[D]. PhD Dissertation. Inner Mongolia Agricultural University, Hohhot. (in Chinese)
[6] 杨子琴,李茂,章笑赟, 等. 饥饿胁迫对龙眼果实脱落及糖代谢的影响[J].果树学报,2011,28(3):428-432.YANG Zi-qin, LI Mao, ZHANG Xiao-yun, et al. (2011). Effects of starvation stress on fruit abscission and sugar metabolism in longan [J]. Journal of Fruit Science, 28(3):428-432. (in Chinese)
[7] Finazzo, S. F., Davenport, T. L., & Schaffer, B. (1994). Partitioning of photoassimilates in avocado (Persea americana Mill.) during flowering and fruit set. Tree Physiology, 14(2): 153-164.
[8] 黄永敬,马培恰,吴文, 等 . 砂糖橘夏梢生长对果实糖代谢及脱落的影响[J].园艺学报,2013,40(10):1 869-1 876.HUANG Yong-jing, MA Pei-qia, WU Wen, et al. (2013). Effects of Summer Shoot Growth on Sugar Metabolism and Abscission of Fruitlet in'Shatangju' Mandarin [J]. Acta Horticulturae Sinica, 40(10):1,869-1,876. (in Chinese)
[9] 曾庆钱,蔡岳文,严振, 等. 化橘红生理落果与若干生理生化指标的关系[J].亚热带植物科学,2008,(3):19-21.ZENG Qing-qian, CAI Yue-wen, YAN Zhen, et al. (2008). Relationship between Physiological Drop and Some Physiological and Biochemical Indexes of Citrus grandis 'Tomentosa' [J]. Subtropical Plant Science, (3):19-21. (in Chinese)
[10] Echeverria, E. (1990). Developmental transition from enzymatic to acid hydrolysis of sucrose in acid limes (Citrus aurantifolia). Plant Physiology, 92(1): 168-171.
[11] Zhou L, Paull R E. (2001).Sucrose Metabolism During Papaya (Carica papaya) Fruit Growth and Ripening. Am Soc Hortic Sci, 126(3): 351-357.
[12] Chengappa, S., Guilleroux, M., Phillips, W., & Shields, R. (1999). Transgenic tomato plants with decreased sucrose synthase are unaltered in starch and sugar accumulation in the fruit. Plant Molecular Biology, 40(2): 213-221.
[13] 卢合全,沈法富,刘凌霄, 等. 植物蔗糖合成酶功能与分子生物学研究进展[J].中国农学通报,2005 (7):34-37, 57.LU He-quan, SHEN Fa-fu, LIU Ling-xiao, et al. (2005). Recent Advances in Study on Plant Sucrose Synthase[J]. Chinese Agricultural Science Bulletin, (7):34-37, 57. (in Chinese)
[14] Marc-André D'Aoust, Yelle, S., & Binh Nguyen-Quoc. (1999). Antisense inhibition of tomato fruit sucrose synthase decreases fruit setting and the sucrose unloading capacity of young fruit. Plant Cell, 11(12): 2,407-2,418.
[14] 张丽丽,刘威生,刘有春, 等.高效液相色谱法测定5个杏品种的糖和酸[J].果树学报,2010,27(1):119-123.ZHANG Li-li, LIU Wei-sheng, LIU You-chun, et al. (2010). Determination of Sugar and Acid in Five Apricot Varieties by High Performance Liquid Chromatography [J]. The Fruit Tree Journal 27(1): 119-123. (In Chinese)
[15] Keller, F., & Ludlow, M. M. (1993). Carbohydrate metabolism in drought-stressed leaves of pigeonpea (cajanus cajan). Journal of Experimental Botany, 44(265):1,351-1,359.
[16] 杨波,车玉红,郭春苗. 扁桃生理落果期不同组织激素浓度的动态变化及其对落果的影响[J].西北植物学报,2015,35(1):118-124.YANG Bo, CHE Yu-hong, GUO Chun-miao. (2015). ynamic changes of hormones in the different tissue of Almond during the physiological fruit drop and its effect on fruit drop [J]. Acta Botanica Boreali-Occidentalia Sinica, 35(1):118-124. (in Chinese)
[17] Miron, D., & Schaffer, A. A. (1991). Sucrose phosphate synthase, sucrose synthase, and invertase activities in developing fruit of lycopersicon esculentum mill. and the sucrose accumulating Lycopersicon hirsutum humb. and bonpl. Plant Physiology, 95(2): 623-627.
[18] 秦翠鲜,桂意云,陈忠良,等.植物蔗糖合成酶基因研究进展[J].分子植物育种,2018,16(12):3 907-3 914.QIN Cui-xiang, GUI Yi-yun, CHEN Zhong-liang, et al. (2018). The Progress of Studies on Sucrose Synthase Genes in Plants [J]. Molecular Plant Breeding, 16(12):3,907-3,914. (in Chinese)
[19] 赵智中,张上隆,徐昌杰,等.蔗糖代谢相关酶在温州蜜柑果实糖积累中的作用[J].园艺学报,2001,(2):112-118.ZHAO Zhi-zhong, ZHANG Shang-long, XU Chang-jie, et al. (2001). The Role of Sucrose Metabolism Related Enzymes in Sugar Accumulation in Citrus Fruits of Wenzhou [J]. Acta Horticulturae Sinica, (2):112-118. (in Chinese)
[20] 龚荣高, 张光伦, 吕秀兰,等. 脐橙果实糖积累与蔗糖代谢相关酶关系的研究[J]. 四川农业大学学报, 2004, 22(1):34-36.GONG Rong-gao, ZHANG Guang-lun, Lü Xiu-lang, et al. (2004). Study on the Relationship between the Sugar Accumulation and Its Metabolizing Enzymes in Navel Orange Fruit [J]. Journal of Sichuan Agricultural University, 22(1):34-36. (in Chinese)
[21] 张春华, 俞明亮, 马瑞娟,等. 桃不同发育时期主要糖类含量和蔗糖合成酶基因表达水平的动态变化[J].江苏农业学报, 2014,(6):1 456-1 463.ZHANG Chun-hua, YU Ming-liang, MA Rui-juan, et al. (2014). Dynamic changes of carbohydrate content and expression level of sucrose synthase genes in peach (Prunus persica) at different developmental stages [J]. Jiangsu Journal of Agricultural Sciences, (6):1,456-1,463. (in Chinese)
[22] Elling, & Lothar. (1995). Effect of metal ions on sucrose synthase from rice grains-a study on enzyme inhibition and enzyme topography. Glycobiology, 5(2): 201-206.