赤霉素对黄瓜幼苗在根区亚低温下生长与氮积累的影响
|
摘要
为了探讨赤霉素(GA)对黄瓜在根区亚低温下氮(N)吸收和代谢的影响机理,以营养液栽培的‘中农26’黄瓜为试材,研究了外源GA对黄瓜在根区亚低温下叶片叶绿素荧光参数、光合参数和碳(C)、氮代谢平衡调节酶活性的影响。结果表明:与根区适温处理相比,根区亚低温使黄瓜幼苗生长受抑,叶片光系统Ⅱ(PSⅡ)的光化学活性、光合放氧速率、净光合速率(Pn)、磷酸烯醇式丙酮酸羧化酶(PEPCase)和蔗糖磷酸合成酶(SPS)活性等降低,单株C与N积累量的比值(C/N)升高,N代谢消耗的能量减少。外源GA处理缓解了根区亚低温对黄瓜生长的抑制,使PSⅡ光化学活性、光合放氧速率、Pn、PEPCase和SPS活性都有不同程度的提高,C/N值和N代谢消耗的能量恢复至与根区适温处理相当的水平。表明,外源GA可以通过增强光能捕获和CO_2同化能力,调节N、C代谢平衡,促进根区亚低温下黄瓜对N的吸收与代谢。
The purpose of this paper is to clarify the effect of gibberellin(GA)on the energy and carbon(C)skeleton supply of nitrogen(N)metabolism of cucumber(Cucumis sativus L.)under suboptimal root-zone temperatures and to provide some theoretic basis for searching an effective way to regulate the N use efficiency of cucumber.Cucumber seedlings at 2-leaf stage were treated with root-zone temperature(RT)22 ℃,RT 16 ℃ or RT 16 ℃ in the presence of 5 μmol·L~(-1) GA_3(GA)for 8 d.Then the items,including individual plant dry mass,chlorophyll fluorescence parameters,photosynthesis parameters,activity of phosphoenolpyruvate carboxylase(PEPCase)and sucrose phosphate synthase(SPS),N and C accumulation,and energy distribution between N and C metabolism were tested.The results showed that exposure of cucumber seedlings to a RT of 16 ℃ led to a significant reduction in the photochemistry activity of PSⅡ,O_2 evolution rate,activity of PEPCase and SPS,and energy distribution in N metabolism,while a rise in C/N.However,application of GA_3 showed prominent effects on mitigating the stress of suboptimal RT.Furthermore,GA treatment increased energy and C skeleton distribution in nitrogen metabolism.In conclusion,GA increased N absorption of cucumber under suboptimal RT conditions through increasing the energy and C skeleton supply.
The purpose of this paper is to clarify the effect of gibberellin(GA)on the energy and carbon(C)skeleton supply of nitrogen(N)metabolism of cucumber(Cucumis sativus L.)under suboptimal root-zone temperatures and to provide some theoretic basis for searching an effective way to regulate the N use efficiency of cucumber.Cucumber seedlings at 2-leaf stage were treated with root-zone temperature(RT)22 ℃,RT 16 ℃ or RT 16 ℃ in the presence of 5 μmol·L~(-1) GA_3(GA)for 8 d.Then the items,including individual plant dry mass,chlorophyll fluorescence parameters,photosynthesis parameters,activity of phosphoenolpyruvate carboxylase(PEPCase)and sucrose phosphate synthase(SPS),N and C accumulation,and energy distribution between N and C metabolism were tested.The results showed that exposure of cucumber seedlings to a RT of 16 ℃ led to a significant reduction in the photochemistry activity of PSⅡ,O_2 evolution rate,activity of PEPCase and SPS,and energy distribution in N metabolism,while a rise in C/N.However,application of GA_3 showed prominent effects on mitigating the stress of suboptimal RT.Furthermore,GA treatment increased energy and C skeleton distribution in nitrogen metabolism.In conclusion,GA increased N absorption of cucumber under suboptimal RT conditions through increasing the energy and C skeleton supply.
引文
Ai Xi-zhen,Wang Xiu-feng,Cui Zhi-feng,Wang Zhen-lin.2006.Effect of calcium on photosynthesis of cucumber under low light intensity and sub-optimal temperature.Scientia Agricultura Sinica,39(9):1865-1871.(in Chinese)艾希珍,王秀峰,崔志峰,王振林.2006.钙对弱光亚适温下黄瓜光合作用的影响.中国农业科学,39(9):1865-1871.
Ashraf M,Karim F,Rasul E.2002.Interactive effects of gibberellic acid(GA3)and salt stress on growth,ion accumulation and photosynthetic capacity of two spring wheat(Triticum aestivum L.)cultivars differing in salt tolerance.Plant Growth Regulation,36(1):49-59.
Bai L Q,Deng H H,Zhang X C,Yu X C,Li Y S.2016.Gibberellin is involved in inhibition of cucumber growth and nitrogen uptake at suboptimal root-zone temperatures.PLOS ONE,11(5):e0156188..
Bloom A J,Caldwell R M,Finazzo J,Warner R L,Weissbart J.1989.Oxygen and carbon dioxide fluxes from barley shoots depend on nitrate assimilation.Plant Physiology,91(1):352-356.
Champigny M L.1995.Integration of photosynthetic carbon and nitrogen metabolism in higher plants.Photosynthesis Research,46:117-127.
Chen Ran-ran.2012.Influences of different levels of nitrogen on carbon and nitrogen metabolisms in transgenic rice plants expressing maize PEPCgene[M.D.Dissertation].Beijing:Chinese Academy of Agricultural Sciences.(in Chinese)陈冉冉.2012.不同氮素水平对转玉米PEPC基因水稻碳、氮代谢的影响[硕士论文].北京:中国农业科学院.
Guo Shi-rong.2007.Science of soil-less culture.Beijing:China Agriculture Press.郭世荣.2007.无土栽培学.北京:中国农业出版社.
Huppe H C,Turpin D H.2003.Integration of carbon and nitrogen metabolism in plant and algal cells.Annual Review of Plant Biology,45(1):577-607.
Khan N A,Ansari H R,Samiullah.1998.Effect of gibberellic acid spray during ontogeny of mustard on growth,nutrient uptake and yield characteristics.Journal of Agronomy&Crop Science,181(1):61-63.
Khan N A,Mir R,Khan M,Javid S,Samiullah.2002.Effects of gibberellic acid spray on nitrogen yield efficiency of mustard grown with different nitrogen levels.Plant Growth Regulation,38(3):243-247.
Kurai T,Wakayama M,Abiko T,Yanagisawa S,Aoki N,Ohsugi R.2011.Introduction of the ZmD of1 gene into rice enhances carbon and nitrogen assimilation under low-nitrogen conditions.Plant Biotechnology Journal,9(8):826-837.
Latzko E,Kelly G J.1983.The many-faceted function of phosphoenolpyruvate carboxylase in C3 plants.Physiologie Vegetable,21(5):805-815.
Liang Wen-juan,Wang Mei-ling,Ai Xi-zhen,Sun Xiao-qi,Wang Hong-tao.2008.Photosynthesis adaptation of cucumber seedlings to suboptimal temperature and low light intensity stress.Transactions of the Chinese Society of Agricultural Engineering,24(8):240-244.(in Chinese)梁文娟,王美玲,艾希珍,孙晓琦,王洪涛.2008.黄瓜幼苗光合作用对亚适温弱光胁迫的适应性.农业工程学报,24(8):240-244.
Liu Jun,Cao Zhi-fu,Huang Yan-nan,Gao Li-hong.2007.Nutrients absorption properties of cucumber in long-season cultivation in solar greenhouse.Scientia Agricultura Sinica,40(9):2109-2113.(in Chinese)刘军,曹之富,黄延楠,高丽红.2007.日光温室黄瓜冬春茬栽培氮磷钾吸收特性研究.中国农业科学,40(9):2109-2113
Liu Yu-mei.2006.Regulation of photosynthesis of cucumber under suboptimal temperature and suboptimal light[Ph.D.Dissertation].Tai’an:Shandong Agricultural University.(in Chinese)刘玉梅.2006.亚适温较弱光照条件下调控黄瓜光合作用的原理与技术研究[博士论文].泰安:山东农业大学.
Ma Dong-yun,Guo Tian-cai,Song Xiao,Wang Chen-yang,Han Qiao-xiao,Yue Yan-jun,Cha Fei-na.2010.Effects of nitrogen fertilizer application on Ru BP carboxylase activity and chlorophyll fluorescence parameters in flag leaves of winter wheat.Acta Botanica Boreali-Occidentalis Sinica,30(11):2197-2202.(in Chinese)马冬云,郭天财,宋晓,王晨阳,韩巧霞,岳艳军,查菲娜.2010.施氮对冬小麦旗叶Ru BP羧化酶活性及叶绿素荧光参数的影响.西北植物学报,30(11):2197-2202.
Migocka M,Warzybok A,K?obus G.2013.The genomic organization and transcriptional pattern of genes encoding nitrate transporters 1(NRT1)in cucumber.Plant&Soil,364(1-2):245-260.
Ren Zhi-yu,Wang Xiu-feng.2002.Effects of root zone temperature on contents of mineral elements and absorbent functions of cucumber seedlings root.Acta Agriculturae Boreali-Sinica,17(4):13-17.(in Chinese)任志雨,王秀峰.2002.根区温度对黄瓜幼苗矿质元素含量及根系吸收功能的影响.华北农学报,17(4):13-17.
Shuichi Y,Ai A,Hiroaki K,Uchimiya H,Miwa T.2004.Metabolic engineering with Dof1 transcription factor in plants:improved nitrogen assimilation and growth under low-nitrogen conditions.Proceedings of the National Academy of Sciences,101(20):7833-7838.
Tian Ji-chun.2000.Carbon and nitrogen metabolism of wheat varieties with different protein content[Ph.D.Dissertation].Beijing:China Agricultural University.(in Chinese)田纪春.2000.不同蛋白质含量小麦品种氮、碳代谢特性的研究[博士论文].北京:中国农业大学.
Tian Ji-chun,Wang Xue-chen,Liu Guang-tian.2001.The coupling and regulation between photosynthesis and nitrogen,carbon metabolism in plant.Chinese Bulletin of Life Sciences,13(4):145-147.(in Chinese)田纪春,王学臣,刘广田.2001.植物光合作用与光合氮、碳代谢的偶联及调节.生命科学,13(4):145-147.
Tsai D S,Arteca R N.1985.Effects of root applications of gibberellic acid on photosynthesis and growth in C3 and C4 plants.Photosynthesis Research,6(2):147-157.
Wang Yong-jian,Zhang Hai-ying,Zhang Feng,Xu Yong,Cao Wan-hong,Kang Guo-bin.2001.Effects of low temperature and low light intensity stress on photosynthesis in seedlings of different cucumber varieties.Acta Horticulturae Sinica,28(3):230-234.(in Chinese)王永健,张海英,张峰,许勇,曹婉虹,康国斌.2001.低温弱光对不同黄瓜品种幼苗光合作用的影响.园艺学报,28(3):230-234.
Wu Xiu,Lu Xiao-min.2015.Effects of brassinolide on the antioxidant system and photosynthesis of cucumber seedlings under suboptimal temperature,light and salt environment.Chinese Journal of Applied Ecology,26(9):2751-2757.(in Chinese)吴秀,陆晓民.2015.亚适宜温光盐环境下油菜素内酯对黄瓜幼苗抗氧化系统及光合作用的影响.应用生态学报,26(9):2751-2757.
Yang Quan-yong,Wang Xiu-feng,Han Yu-rui,Yan Jing-jing,Wei Min,Yang Feng-juan,Shi Qing-hua,Li Yan.2015.Effects of NO-3stress on cucumber seedling growth and magnesium absorbing under suboptimal temperature.Chinese Journal of Applied Ecology,26(5):1343-1350.(in Chinese)杨全勇,王秀峰,韩宇睿,杨静静,魏珉,杨凤娟,史庆华,李岩.2015.亚适温和硝酸盐胁迫对黄瓜幼苗生长及镁吸收的影响.应用生态学报,26(5):1343-1350.
Yin Lu-lu,Yu Xian-chang,Wang Ying-hua,Xu Zhen-hang,Li Kun,Han Dao-jie.2007.Effect of 5-aminolevulinic acid on chilling-tolerance in cucumber seedlings.Acta Agriculturae Boreali-Occidentalis Sinica,16(4):166-169.(in Chinese)尹璐璐,于贤昌,王英华,许贞杭,李坤,韩道杰.2007.5-氨基乙酰丙酸对黄瓜幼苗抗冷性的影响.西北农业学报,16(4):166-169.
Zhang Meng,Wang Xiu-feng,Zhang Fan-yang,Wei Min,Shi Qing-hua,Yang Feng-juan,Li Yan.2014.Effects of chlorophyll in iron on osmotic adjustment and activities of antioxidantive enzymes in cucumber seedlings under suboptimal temperature.Chinese Journal of Applied Ecology,25(12):3527-3532.(in Chinese)张蒙,王秀峰,张帆洋,魏珉,史庆华,杨凤娟,李岩.2014.叶绿酸铁对亚适温条件下黄瓜幼苗渗透调节物质及抗氧化酶活性的影响.应用生态学报,25(12):3527-3532.
Zhang Ying,Jiang Wei-jie,Yu Hong-jun,Yang Xue-yong.2012.Exogenous abscisic acid alleviates low temperature-induced oxidative damage inseedlings of Cucumis sativus L.Transactions of the Chinese Society of Agricultural Engineering,28(26):221-228.(in Chinese)张颖,蒋卫杰,余宏军,杨学勇.2012.外源ABA缓解黄瓜幼苗中低温诱导的氧化损伤.农业工程学报,28(26):221-228.
Ashraf M,Karim F,Rasul E.2002.Interactive effects of gibberellic acid(GA3)and salt stress on growth,ion accumulation and photosynthetic capacity of two spring wheat(Triticum aestivum L.)cultivars differing in salt tolerance.Plant Growth Regulation,36(1):49-59.
Bai L Q,Deng H H,Zhang X C,Yu X C,Li Y S.2016.Gibberellin is involved in inhibition of cucumber growth and nitrogen uptake at suboptimal root-zone temperatures.PLOS ONE,11(5):e0156188..
Bloom A J,Caldwell R M,Finazzo J,Warner R L,Weissbart J.1989.Oxygen and carbon dioxide fluxes from barley shoots depend on nitrate assimilation.Plant Physiology,91(1):352-356.
Champigny M L.1995.Integration of photosynthetic carbon and nitrogen metabolism in higher plants.Photosynthesis Research,46:117-127.
Chen Ran-ran.2012.Influences of different levels of nitrogen on carbon and nitrogen metabolisms in transgenic rice plants expressing maize PEPCgene[M.D.Dissertation].Beijing:Chinese Academy of Agricultural Sciences.(in Chinese)陈冉冉.2012.不同氮素水平对转玉米PEPC基因水稻碳、氮代谢的影响[硕士论文].北京:中国农业科学院.
Guo Shi-rong.2007.Science of soil-less culture.Beijing:China Agriculture Press.郭世荣.2007.无土栽培学.北京:中国农业出版社.
Huppe H C,Turpin D H.2003.Integration of carbon and nitrogen metabolism in plant and algal cells.Annual Review of Plant Biology,45(1):577-607.
Khan N A,Ansari H R,Samiullah.1998.Effect of gibberellic acid spray during ontogeny of mustard on growth,nutrient uptake and yield characteristics.Journal of Agronomy&Crop Science,181(1):61-63.
Khan N A,Mir R,Khan M,Javid S,Samiullah.2002.Effects of gibberellic acid spray on nitrogen yield efficiency of mustard grown with different nitrogen levels.Plant Growth Regulation,38(3):243-247.
Kurai T,Wakayama M,Abiko T,Yanagisawa S,Aoki N,Ohsugi R.2011.Introduction of the ZmD of1 gene into rice enhances carbon and nitrogen assimilation under low-nitrogen conditions.Plant Biotechnology Journal,9(8):826-837.
Latzko E,Kelly G J.1983.The many-faceted function of phosphoenolpyruvate carboxylase in C3 plants.Physiologie Vegetable,21(5):805-815.
Liang Wen-juan,Wang Mei-ling,Ai Xi-zhen,Sun Xiao-qi,Wang Hong-tao.2008.Photosynthesis adaptation of cucumber seedlings to suboptimal temperature and low light intensity stress.Transactions of the Chinese Society of Agricultural Engineering,24(8):240-244.(in Chinese)梁文娟,王美玲,艾希珍,孙晓琦,王洪涛.2008.黄瓜幼苗光合作用对亚适温弱光胁迫的适应性.农业工程学报,24(8):240-244.
Liu Jun,Cao Zhi-fu,Huang Yan-nan,Gao Li-hong.2007.Nutrients absorption properties of cucumber in long-season cultivation in solar greenhouse.Scientia Agricultura Sinica,40(9):2109-2113.(in Chinese)刘军,曹之富,黄延楠,高丽红.2007.日光温室黄瓜冬春茬栽培氮磷钾吸收特性研究.中国农业科学,40(9):2109-2113
Liu Yu-mei.2006.Regulation of photosynthesis of cucumber under suboptimal temperature and suboptimal light[Ph.D.Dissertation].Tai’an:Shandong Agricultural University.(in Chinese)刘玉梅.2006.亚适温较弱光照条件下调控黄瓜光合作用的原理与技术研究[博士论文].泰安:山东农业大学.
Ma Dong-yun,Guo Tian-cai,Song Xiao,Wang Chen-yang,Han Qiao-xiao,Yue Yan-jun,Cha Fei-na.2010.Effects of nitrogen fertilizer application on Ru BP carboxylase activity and chlorophyll fluorescence parameters in flag leaves of winter wheat.Acta Botanica Boreali-Occidentalis Sinica,30(11):2197-2202.(in Chinese)马冬云,郭天财,宋晓,王晨阳,韩巧霞,岳艳军,查菲娜.2010.施氮对冬小麦旗叶Ru BP羧化酶活性及叶绿素荧光参数的影响.西北植物学报,30(11):2197-2202.
Migocka M,Warzybok A,K?obus G.2013.The genomic organization and transcriptional pattern of genes encoding nitrate transporters 1(NRT1)in cucumber.Plant&Soil,364(1-2):245-260.
Ren Zhi-yu,Wang Xiu-feng.2002.Effects of root zone temperature on contents of mineral elements and absorbent functions of cucumber seedlings root.Acta Agriculturae Boreali-Sinica,17(4):13-17.(in Chinese)任志雨,王秀峰.2002.根区温度对黄瓜幼苗矿质元素含量及根系吸收功能的影响.华北农学报,17(4):13-17.
Shuichi Y,Ai A,Hiroaki K,Uchimiya H,Miwa T.2004.Metabolic engineering with Dof1 transcription factor in plants:improved nitrogen assimilation and growth under low-nitrogen conditions.Proceedings of the National Academy of Sciences,101(20):7833-7838.
Tian Ji-chun.2000.Carbon and nitrogen metabolism of wheat varieties with different protein content[Ph.D.Dissertation].Beijing:China Agricultural University.(in Chinese)田纪春.2000.不同蛋白质含量小麦品种氮、碳代谢特性的研究[博士论文].北京:中国农业大学.
Tian Ji-chun,Wang Xue-chen,Liu Guang-tian.2001.The coupling and regulation between photosynthesis and nitrogen,carbon metabolism in plant.Chinese Bulletin of Life Sciences,13(4):145-147.(in Chinese)田纪春,王学臣,刘广田.2001.植物光合作用与光合氮、碳代谢的偶联及调节.生命科学,13(4):145-147.
Tsai D S,Arteca R N.1985.Effects of root applications of gibberellic acid on photosynthesis and growth in C3 and C4 plants.Photosynthesis Research,6(2):147-157.
Wang Yong-jian,Zhang Hai-ying,Zhang Feng,Xu Yong,Cao Wan-hong,Kang Guo-bin.2001.Effects of low temperature and low light intensity stress on photosynthesis in seedlings of different cucumber varieties.Acta Horticulturae Sinica,28(3):230-234.(in Chinese)王永健,张海英,张峰,许勇,曹婉虹,康国斌.2001.低温弱光对不同黄瓜品种幼苗光合作用的影响.园艺学报,28(3):230-234.
Wu Xiu,Lu Xiao-min.2015.Effects of brassinolide on the antioxidant system and photosynthesis of cucumber seedlings under suboptimal temperature,light and salt environment.Chinese Journal of Applied Ecology,26(9):2751-2757.(in Chinese)吴秀,陆晓民.2015.亚适宜温光盐环境下油菜素内酯对黄瓜幼苗抗氧化系统及光合作用的影响.应用生态学报,26(9):2751-2757.
Yang Quan-yong,Wang Xiu-feng,Han Yu-rui,Yan Jing-jing,Wei Min,Yang Feng-juan,Shi Qing-hua,Li Yan.2015.Effects of NO-3stress on cucumber seedling growth and magnesium absorbing under suboptimal temperature.Chinese Journal of Applied Ecology,26(5):1343-1350.(in Chinese)杨全勇,王秀峰,韩宇睿,杨静静,魏珉,杨凤娟,史庆华,李岩.2015.亚适温和硝酸盐胁迫对黄瓜幼苗生长及镁吸收的影响.应用生态学报,26(5):1343-1350.
Yin Lu-lu,Yu Xian-chang,Wang Ying-hua,Xu Zhen-hang,Li Kun,Han Dao-jie.2007.Effect of 5-aminolevulinic acid on chilling-tolerance in cucumber seedlings.Acta Agriculturae Boreali-Occidentalis Sinica,16(4):166-169.(in Chinese)尹璐璐,于贤昌,王英华,许贞杭,李坤,韩道杰.2007.5-氨基乙酰丙酸对黄瓜幼苗抗冷性的影响.西北农业学报,16(4):166-169.
Zhang Meng,Wang Xiu-feng,Zhang Fan-yang,Wei Min,Shi Qing-hua,Yang Feng-juan,Li Yan.2014.Effects of chlorophyll in iron on osmotic adjustment and activities of antioxidantive enzymes in cucumber seedlings under suboptimal temperature.Chinese Journal of Applied Ecology,25(12):3527-3532.(in Chinese)张蒙,王秀峰,张帆洋,魏珉,史庆华,杨凤娟,李岩.2014.叶绿酸铁对亚适温条件下黄瓜幼苗渗透调节物质及抗氧化酶活性的影响.应用生态学报,25(12):3527-3532.
Zhang Ying,Jiang Wei-jie,Yu Hong-jun,Yang Xue-yong.2012.Exogenous abscisic acid alleviates low temperature-induced oxidative damage inseedlings of Cucumis sativus L.Transactions of the Chinese Society of Agricultural Engineering,28(26):221-228.(in Chinese)张颖,蒋卫杰,余宏军,杨学勇.2012.外源ABA缓解黄瓜幼苗中低温诱导的氧化损伤.农业工程学报,28(26):221-228.