葡萄叶片衰老过程中不同光质对其光合和叶绿体超微结构的影响
|
摘要
以栽植于日光温室中适宜延迟栽培的‘贝达’砧‘意大利’葡萄为试材,自2013年8月1日始至落叶(2014年1月31日)止每天分别进行红光和蓝光不同光质延长光照至24:00补光处理,以不补光作为对照,研究不同光质补光处理对葡萄叶片衰老过程中叶片的外观形态特征、叶绿素含量、净光合速率、F_v/F_m、可溶性蛋白质含量和叶绿体超微结构的影响,为叶片抗衰老技术的研发提供理论依据。研究结果表明:补充红光显著提高了叶片的叶绿素含量、净光合速率、F_v/F_m值和可溶性蛋白质含量,叶绿体基粒片层排列整齐有序,明显减缓了叶绿体超微结构的解体,显著延缓叶片衰老;补充蓝光处理前期显著降低了叶片的叶绿素含量、净光合速率、F_v/F_m值和可溶性蛋白质含量,破坏了叶绿体的超微结构,加速了植株的衰老进程;但在叶片衰老后期,叶绿素含量、净光合速率、F_v/F_m值和可溶性蛋白质含量逐渐高于对照,并且叶绿体的基粒片层比对照排列整齐有序,在一定程度上延缓了叶片衰老。补充蓝光处理的叶片可溶性蛋白质含量一直显著高于补充红光和对照。综上,补充红光处理有效延缓了叶片衰老进程,延长了叶片的生理功能期。
The experiment was carried out in 2013 at the fruit garden of the Chinese Academy of Agricultural Sciences in Liaoning,which was equipped with a core grape technology demonstration garden‘.Italy'grape cultivars were grafted on Beta as test materials. And their row spacing was 0.7 m,line spacing was 2 m. The lean dragon trunk matches the V shape leaf curtain,and used the integrated management of water and fertilizer,other management was the same as the routine. Since the beginning of August 1 st to the defoliation day before dark for half an hour to fill 24:00,use red light and blue light to deal with the grape cultivars,with no-fill-light grapes as contrast,studying morphological characteristics,chlorophyll content,net photosynthetic rate,F_v/F_m value,soluble proteins content and chloroplast ultrastructure of grape leaves in aging process,to provide theoretical basis for the research and development of leaf anti-aging technology. The research result showed that compared with the control group,red light treatment increased the chlorophyll content,net photosynthetic rate,F_v/F_m value and soluble proteins content of leaf,the chloroplast grana were arranged in order,the disintegration of chloroplast ultrastructure was obviously slowed down,leaf senescence was delayed significantly. Blue light treatment significantly reduced the leaf chlorophyll content,net photosynthetic rate,soluble proteins content and F_v/F_m value,the ultrastructure of chloroplasts was destroyed,and the aging process of grapes was accelerated. But in the later stage of leaf senescence,the chlorophyll content of leaf,the net photosynthetic rate,soluble proteins content and F_v/F_m value of the blue light treatment was higher than that of the control. Moreover,the grana lamellae of leaf chloroplasts are arranged in order,leaf senescence was delayed in some degree. The content of soluble proteins in leaf of blue light treatment was significantly higher than that of red light treatment and control group. Summing up the above,the red light as supplement light had the best effect,the leaf senescence process was delayed effectively,and the physiological function of leaves was prolonged.
The experiment was carried out in 2013 at the fruit garden of the Chinese Academy of Agricultural Sciences in Liaoning,which was equipped with a core grape technology demonstration garden‘.Italy'grape cultivars were grafted on Beta as test materials. And their row spacing was 0.7 m,line spacing was 2 m. The lean dragon trunk matches the V shape leaf curtain,and used the integrated management of water and fertilizer,other management was the same as the routine. Since the beginning of August 1 st to the defoliation day before dark for half an hour to fill 24:00,use red light and blue light to deal with the grape cultivars,with no-fill-light grapes as contrast,studying morphological characteristics,chlorophyll content,net photosynthetic rate,F_v/F_m value,soluble proteins content and chloroplast ultrastructure of grape leaves in aging process,to provide theoretical basis for the research and development of leaf anti-aging technology. The research result showed that compared with the control group,red light treatment increased the chlorophyll content,net photosynthetic rate,F_v/F_m value and soluble proteins content of leaf,the chloroplast grana were arranged in order,the disintegration of chloroplast ultrastructure was obviously slowed down,leaf senescence was delayed significantly. Blue light treatment significantly reduced the leaf chlorophyll content,net photosynthetic rate,soluble proteins content and F_v/F_m value,the ultrastructure of chloroplasts was destroyed,and the aging process of grapes was accelerated. But in the later stage of leaf senescence,the chlorophyll content of leaf,the net photosynthetic rate,soluble proteins content and F_v/F_m value of the blue light treatment was higher than that of the control. Moreover,the grana lamellae of leaf chloroplasts are arranged in order,leaf senescence was delayed in some degree. The content of soluble proteins in leaf of blue light treatment was significantly higher than that of red light treatment and control group. Summing up the above,the red light as supplement light had the best effect,the leaf senescence process was delayed effectively,and the physiological function of leaves was prolonged.
引文
Andersson A,Keskitalo J,Sj?din A,Bhalerao R,Sterky F,Wissel K,Tandre K,Aspeborg H,Moyle R,Ohmiya Y,Bhalerao R,Brunner A,Gustafsson P,Karlsson J,Lundeberg J,Nilsson O,Sandberg G,Strauss S,Sundberg B,Uhlen M,Jansson S,Nilsson P.2004.A transcriptional time table of autumn senescence.Genome Biology,5(4):R24.
Borras L,Maddonni G A,Otegui M E.2003.Leaf senescence in maize hybrids:plant population,row spacing and kernel set effects.Field Crops Research,82(1):13-26.
Brouwer B,Ziolkowska A,Bagard M,Oliver K,Gardestrom P.2012.The impact of light intensity on shade induced leaf senescence.Plant,Cell and Environment,35(6):1084-1098.
Buchanan W V,Ainsworth C.1997.Leaf senescence in Brassica napus:cloning of senescence related genes by subtractive hybridisation.Plant Molecular Biology,33(5):821-834.
Butler R D,Simon E W.1971.Ultrastructural aspects of senescence in plants.Adv Gerontol Res,3:73-129.
Ding Meng-qiu,Wen Shi-wen,Lu Wei-ping,Lu Da-lei.2017.Effects of weak-light stress after pollination on grain filling and leaf senescence in sweet maize.Journal of Nuclear Agricultural Sciences,31(5):964-971.(in Chinese)丁梦秋,闻诗文,陆卫平,陆大雷.2017.结实期弱光胁迫对甜玉米籽粒灌浆和叶片衰老的影响.核农学报,31(5):964-971.
Gan S,Amasino R M.1997.Making sense of senescence(molecular genetic regulation and manipulation of leaf senescence).Plant Physiology,113(2):313-319.
Han Biao,Chen Guo-xiang,Gao Zhi-ping,Wei Xiao-dong,Xie Kai-bin,Yang Xian-song.2010.The changes of PSⅡchlorophyll fluorescence dynamic characteristic during leaf senescence of ginkgo.Acta Horticulturae Sinica,37(2):173-178.(in Chinese)韩彪,陈国祥,高志萍,魏晓东,解凯彬,杨贤松.2010.银杏叶片衰老过程中PSⅡ荧光动力学特性变化.园艺学报,37(2):173-178.
Jiang Jian-fu,Zhang Ying,Tian Zhi-shuo,Fan Xiu-cai,Liu Chong-huai.2012.Observation on microstructure and ultrastructure of leaves in Vitis L.Acta Bot Boreal-Occient Sin,32(7):1365-1371.(in Chinese)姜建福,张颖,田智硕,樊秀彩,刘崇怀.2012.葡萄属植物叶片显微和超微结构观察.西北植物学报,32(7):1365-1371.
Kowallik W.1982.Blue light effects on respiration.Annual Review of Plant Physiology,33:51-72.
Leong T Y,Goodchild D J,Anderson J M.1985.Effect of light quality on the composition,function,and structure of photosynthetic thylakoid membranes of Asplenium australasicum(Sm.)Hook.Plant Physiology,78(3):561-567.
Li He-sheng.2000.Principles and techniques of plant physiology and biochemistry experiments.Beijing:Higher Education Press:182-185.(in Chinese)李合生.2000.植物生理生化实验原理和技术.北京:高等教育出版社:182-185.
Lim P O,Woo H R,Nam H G.2003.Molecular genetics of leaf senescence in Arabidopsis.Trends in Plant Science,8(6):272-278.
Liu Rui-xian,Wang You-hua,Chen Bing-lin,Guo Wen-qi,Zhou Zhi-guo.2008.Effects of nitrogen levels on photosynthesis and chlorophyll fluorescence characteristics under drought stress in cotton flowering and boll forming stage.Acta Agronomic Sinica,34(4):675-683.(in Chinese)刘瑞显,王友华,陈兵林,郭文琦,周治国.2008.花铃期干旱胁迫下氮素水平对棉花光合作用与叶绿素荧光特性的影响.作物学报,34(4):675-683.
Lu Ding-zhi,Pan Yu-cai,Ma Yue-fang,Lin Zong-da,Bao Wei-qun.1988.Physiological and biochemical studies in leaf senescenceat heading and grain formation stage in hybrid rice.Scientia Agricultura Sinica,21(3):21-26.(in Chinese)陆定志,潘裕才,马跃芳,林宗达,鲍为群.1988.杂交水稻抽穗结实期间叶片衰老的生理生化研究.中国农业科学,21(3):21-26.
Lu Su-ping,Zhao Ming-qin,Liu Peng-fei,Li Yu-e,Fu Bo.2016.Effect of light quality on endogenous hormone content and lipid peroxidation of membrane system during aging process of flue-cured tobacco leaf.Journal of Agricultural Science and Technology,18(6):44-51.(in Chinese)卢素萍,赵铭钦,刘鹏飞,李玉娥,付博.2016.光质对烤烟叶片衰老过程中内源激素含量和膜脂过氧化的影响.中国农业科技导报,18(6):44-51.
Mao Jian-fei,Li Kai-wei,Yang Zai-qiang,Du Jin-shan.2018.Effects of low Light stress and strong light restoration on photosynthetic and senescence characteristics of leaves of greenhouse red globe grape.Jiangsu Agricultural Science,46(6):105-111.(in Chinese)毛剑飞,李凯伟,杨再强,杜金杉.2018.弱光胁迫及强光恢复对设施红地球葡萄叶片光合及衰老特性的影响.江苏农业科学,46(6):105-111.
Maxwell K,Johnson G N.2000.Chlorophyll fluorescence a practical guide.Journal of Experimental Botany,51(345):659-668.
Oh S A,Lee S Y,Chung I K,Lee C H,Nam H G.1996.A senescence-associated gene of Arabidopsis thaliana is distinctively regulated during natural and artificially induced leaf senescence.Plant Molecular Biology,30(4):739-754.
Ramalho J C,Marques N C,Semedo J N,Matos M C,Quartin V L.2002.Photosynthetic performance and pigment compositeon of leaves from two tropical species is determined by light quality.Plant Biology,4(1):112-120.
Saebo A,Krekling T,Appelgren M.1995.Light quality affects photosynthesis and leaf anatomy of birch plantlets in vitro.Plant Cell,Tissue and Organ Culture,41(2):177-185.
Simeonova E,Garstka M,Koziol-Lipinska J,Mostowska A.2004.Monitoring the mitochondrial transmembrane pote-ntial with the JC-1fluorochrome in programmed cell death during mesophyll leaf senescence.Protoplasma,223(2-4):143-153.
Strasser R J,Tsimilli M M,Srivastava A.2004.Analysis of the chlorophyll a fluorescence transient//Papageorgiou G.Advances in Photosynthesis and Respiration.Netherlands:KAP Press:321-362.
Wang H,Gu M,Cui J,Shi K,Zhou Y,Yu J.2009.Effects of light quality on CO2 assimilation,chlorophyll-fluorescence quenching,expression of Calvin cycle genes and carbohydrate accumulation in Cucumis sativus.Journal of Photochemistry and Photobiology B:Biology,96(1):30-37.
Wang Hai-bo,Wang Shuai,Wang Xiao-di,Shi Xiang-bin,Wang Bao-liang,Zheng Xiao-cui,Wang Zhi-qiang,Liu Feng-zhi.2017.Effects of light quality on leaf senescence and endogenous hormones content in grapevine under protected cultivation.Chinese Journal of Applied Ecology,28(11):3535-3543.(in Chinese)王海波,王帅,王孝娣,史祥宾,王宝亮,郑晓翠,王志强,刘凤之.2017.光质对设施葡萄叶片衰老与内源激素含量的影响.应用生态学报,28(11):3535-3543.
Wang Jian-ying,Yao Xiao-hua,Zhang Zhi-bin.2011.Research progress on mechanism and regulation of plant leaf senescence.Journal of Anhui Agri Sci,39(31):19036-19038,19058.(in Chinese)王建勇,姚晓华,张志斌.2011.植物叶片衰老机理与调控研究进展.安徽农业科学,39(31):19036-19038,19058.
Wang Shuai,Liu Feng-zhi,Wang Hai-bo.2015.Study on leaf senescence physiology and anti-aging technology of grape cultured in protected and delayed cultivation[M.D.Dissertation].Beijing:Chinese Academy of Agricultural Sciences.(in Chinese)王帅,刘凤之,王海波.2015.设施葡萄延迟栽培叶片衰老生理及抗衰老技术研究[硕士论文].北京:中国农业科学院.
Wang Shuai,Wang Xiaodi,Shi Xiangbin,Wang Baoliang,Zheng Xiaocui,Wang Haibo,Liu Fengzhi.2016.Red and blue lights significantly affect photosynthetic properties and ultrastructure of mesophyll cells in senescing grape leaves.Horticultural Plant Journal,2(2):82-90.
Wen Han,Nie Fan.2000.Effects of drought on senescence of flag leaf and yield factors in rice.Journal of Anhui Agricultural University,27(2):135-137.(in Chinese)文汉,聂凡.2000.干旱对水稻抽穗后旗叶衰老和产量构成因子的影响.安徽农业大学学报,27(2):135-137.
Xiong Bing-lin,Wang Shi-wen,Wang Xin-yue,Chen Dao-qian,Yin Li-na,Deng Xi-ping.2016.Effects of nitrogenous fertilizer on leaf senescence of maize and the associate with carbon/nitrogen balance under drought stress.Journal of Maize Sciences,24(3):138-146.(in Chinese)熊炳霖,王仕稳,王鑫月,陈道钳,殷俐娜,邓西平.2016.干旱胁迫下氮肥对玉米叶片衰老影响及与碳氮平衡的关系.玉米科学,24(3):138-146.
Xu Chao-hua,Li Jun-ying,Cui Ming-kun,Ma Er-dong,Huang Guo-bin,Gong Ming.2013.Effects of supplemental lighting on growth and photo-synthesis of tobacco leaves.Acta Bot Boreal-Occient Sin,33(4):763-770.(in Chinese)徐超华,李军营,崔明昆,马二登,黄国宾,龚明.2013.延长光照时间对烟草叶片生长发育及光合特性的影响.西北植物学报,33(4):763-770.
Xu Li,Liu Shi-qi,Qi Lian-dong,Liang Qing-ling,Yu Wen-yan.2007.Effect of light quality on leaf lettuce photosynthesis and chlorophyll fluorescence.Chinese Agricultural Science Bulletin,23(1):96-100.(in Chinese)许莉,刘世琦,齐连东,梁庆玲,于文艳.2007.不同光质对叶用莴苣光合作用及叶绿素荧光的影响.中国农学通报,23(1):96-100.
Yan Xiao-hua,Yu Ji-hua.2016.Effects of supplemental LED light on photosynthetic pigment contentsand antioxidant enzyme activities of cucumber seedling leaves.Journal of Desert Research,36(2):392-398.(in Chinese)闫晓花,郁继华.2016.LED补光对温室黄瓜幼苗抗衰老及抗氧化酶系统的影响.中国沙漠,36(2):392-398.
Zhai Rong-rong,Feng Yue,Cao Li-yong,Cheng Shi-hua,Wu Wei-ming.2011.Review of rice leaf senescence.Chinese Rice,17(1):7-12.(in Chinese)翟荣荣,冯跃,曹立勇,程式华,吴伟明.2011.水稻叶片衰老研究进展.中国稻米,17(1):7-12.
Zhang Jie,Li Tian-lai.2005.Effects of daytime sub-high temperature on photosynthesis and chloroplast ultrastructure of tomato leaves in greenhouse.Acta Horticulturae Sinica,32(4):614-619.(in Chinese)张洁,李天来.2005.日光温室亚高温对番茄光合作用及叶绿体超微结构的影响.园艺学报,32(4):614-619.
Zheng Shao-qing,Zeng Guang-wen.1990.Senescence of plant leaf and the molecular approach for its delaying.Plant Physiology Journal,35(1):152-157.(in Chinese)郑少青,曾广文.1990.植物叶片衰老及其延缓的分子途径.植物生理学通讯,35(1):152-157.
Zhu Xue-mei,Ke Yong-pei,Shao Ji-rong,Lin Li-jin,Yang Yuan-xiang.2005.Effects of high temperature stress on activated oxygen metabolism of the leaves of the heavy panicle type of medium india hybrid rice.Seed,24(3):25-27,32.(in Chinese)朱雪梅,柯永培,邵继荣,林立金,杨远祥.2005.高温胁迫对重穗型水稻品种叶片活性氧代谢的影.种子,24(3):25-27,32.
Zimmermann P,Zentgraf U.2005.The correlation between oxidative stress and leaf senescence during plant development.Cellular and Molecular Biology Letters,10(3):515-534.
Zou Qi.2000.Experimental guidance of plant physiology.Beijing:Chinese Agricultural Press:72-75.(in Chinese)邹琦.2000.植物生理学实验指导.北京:中国农业出版社:72-75.
Borras L,Maddonni G A,Otegui M E.2003.Leaf senescence in maize hybrids:plant population,row spacing and kernel set effects.Field Crops Research,82(1):13-26.
Brouwer B,Ziolkowska A,Bagard M,Oliver K,Gardestrom P.2012.The impact of light intensity on shade induced leaf senescence.Plant,Cell and Environment,35(6):1084-1098.
Buchanan W V,Ainsworth C.1997.Leaf senescence in Brassica napus:cloning of senescence related genes by subtractive hybridisation.Plant Molecular Biology,33(5):821-834.
Butler R D,Simon E W.1971.Ultrastructural aspects of senescence in plants.Adv Gerontol Res,3:73-129.
Ding Meng-qiu,Wen Shi-wen,Lu Wei-ping,Lu Da-lei.2017.Effects of weak-light stress after pollination on grain filling and leaf senescence in sweet maize.Journal of Nuclear Agricultural Sciences,31(5):964-971.(in Chinese)丁梦秋,闻诗文,陆卫平,陆大雷.2017.结实期弱光胁迫对甜玉米籽粒灌浆和叶片衰老的影响.核农学报,31(5):964-971.
Gan S,Amasino R M.1997.Making sense of senescence(molecular genetic regulation and manipulation of leaf senescence).Plant Physiology,113(2):313-319.
Han Biao,Chen Guo-xiang,Gao Zhi-ping,Wei Xiao-dong,Xie Kai-bin,Yang Xian-song.2010.The changes of PSⅡchlorophyll fluorescence dynamic characteristic during leaf senescence of ginkgo.Acta Horticulturae Sinica,37(2):173-178.(in Chinese)韩彪,陈国祥,高志萍,魏晓东,解凯彬,杨贤松.2010.银杏叶片衰老过程中PSⅡ荧光动力学特性变化.园艺学报,37(2):173-178.
Jiang Jian-fu,Zhang Ying,Tian Zhi-shuo,Fan Xiu-cai,Liu Chong-huai.2012.Observation on microstructure and ultrastructure of leaves in Vitis L.Acta Bot Boreal-Occient Sin,32(7):1365-1371.(in Chinese)姜建福,张颖,田智硕,樊秀彩,刘崇怀.2012.葡萄属植物叶片显微和超微结构观察.西北植物学报,32(7):1365-1371.
Kowallik W.1982.Blue light effects on respiration.Annual Review of Plant Physiology,33:51-72.
Leong T Y,Goodchild D J,Anderson J M.1985.Effect of light quality on the composition,function,and structure of photosynthetic thylakoid membranes of Asplenium australasicum(Sm.)Hook.Plant Physiology,78(3):561-567.
Li He-sheng.2000.Principles and techniques of plant physiology and biochemistry experiments.Beijing:Higher Education Press:182-185.(in Chinese)李合生.2000.植物生理生化实验原理和技术.北京:高等教育出版社:182-185.
Lim P O,Woo H R,Nam H G.2003.Molecular genetics of leaf senescence in Arabidopsis.Trends in Plant Science,8(6):272-278.
Liu Rui-xian,Wang You-hua,Chen Bing-lin,Guo Wen-qi,Zhou Zhi-guo.2008.Effects of nitrogen levels on photosynthesis and chlorophyll fluorescence characteristics under drought stress in cotton flowering and boll forming stage.Acta Agronomic Sinica,34(4):675-683.(in Chinese)刘瑞显,王友华,陈兵林,郭文琦,周治国.2008.花铃期干旱胁迫下氮素水平对棉花光合作用与叶绿素荧光特性的影响.作物学报,34(4):675-683.
Lu Ding-zhi,Pan Yu-cai,Ma Yue-fang,Lin Zong-da,Bao Wei-qun.1988.Physiological and biochemical studies in leaf senescenceat heading and grain formation stage in hybrid rice.Scientia Agricultura Sinica,21(3):21-26.(in Chinese)陆定志,潘裕才,马跃芳,林宗达,鲍为群.1988.杂交水稻抽穗结实期间叶片衰老的生理生化研究.中国农业科学,21(3):21-26.
Lu Su-ping,Zhao Ming-qin,Liu Peng-fei,Li Yu-e,Fu Bo.2016.Effect of light quality on endogenous hormone content and lipid peroxidation of membrane system during aging process of flue-cured tobacco leaf.Journal of Agricultural Science and Technology,18(6):44-51.(in Chinese)卢素萍,赵铭钦,刘鹏飞,李玉娥,付博.2016.光质对烤烟叶片衰老过程中内源激素含量和膜脂过氧化的影响.中国农业科技导报,18(6):44-51.
Mao Jian-fei,Li Kai-wei,Yang Zai-qiang,Du Jin-shan.2018.Effects of low Light stress and strong light restoration on photosynthetic and senescence characteristics of leaves of greenhouse red globe grape.Jiangsu Agricultural Science,46(6):105-111.(in Chinese)毛剑飞,李凯伟,杨再强,杜金杉.2018.弱光胁迫及强光恢复对设施红地球葡萄叶片光合及衰老特性的影响.江苏农业科学,46(6):105-111.
Maxwell K,Johnson G N.2000.Chlorophyll fluorescence a practical guide.Journal of Experimental Botany,51(345):659-668.
Oh S A,Lee S Y,Chung I K,Lee C H,Nam H G.1996.A senescence-associated gene of Arabidopsis thaliana is distinctively regulated during natural and artificially induced leaf senescence.Plant Molecular Biology,30(4):739-754.
Ramalho J C,Marques N C,Semedo J N,Matos M C,Quartin V L.2002.Photosynthetic performance and pigment compositeon of leaves from two tropical species is determined by light quality.Plant Biology,4(1):112-120.
Saebo A,Krekling T,Appelgren M.1995.Light quality affects photosynthesis and leaf anatomy of birch plantlets in vitro.Plant Cell,Tissue and Organ Culture,41(2):177-185.
Simeonova E,Garstka M,Koziol-Lipinska J,Mostowska A.2004.Monitoring the mitochondrial transmembrane pote-ntial with the JC-1fluorochrome in programmed cell death during mesophyll leaf senescence.Protoplasma,223(2-4):143-153.
Strasser R J,Tsimilli M M,Srivastava A.2004.Analysis of the chlorophyll a fluorescence transient//Papageorgiou G.Advances in Photosynthesis and Respiration.Netherlands:KAP Press:321-362.
Wang H,Gu M,Cui J,Shi K,Zhou Y,Yu J.2009.Effects of light quality on CO2 assimilation,chlorophyll-fluorescence quenching,expression of Calvin cycle genes and carbohydrate accumulation in Cucumis sativus.Journal of Photochemistry and Photobiology B:Biology,96(1):30-37.
Wang Hai-bo,Wang Shuai,Wang Xiao-di,Shi Xiang-bin,Wang Bao-liang,Zheng Xiao-cui,Wang Zhi-qiang,Liu Feng-zhi.2017.Effects of light quality on leaf senescence and endogenous hormones content in grapevine under protected cultivation.Chinese Journal of Applied Ecology,28(11):3535-3543.(in Chinese)王海波,王帅,王孝娣,史祥宾,王宝亮,郑晓翠,王志强,刘凤之.2017.光质对设施葡萄叶片衰老与内源激素含量的影响.应用生态学报,28(11):3535-3543.
Wang Jian-ying,Yao Xiao-hua,Zhang Zhi-bin.2011.Research progress on mechanism and regulation of plant leaf senescence.Journal of Anhui Agri Sci,39(31):19036-19038,19058.(in Chinese)王建勇,姚晓华,张志斌.2011.植物叶片衰老机理与调控研究进展.安徽农业科学,39(31):19036-19038,19058.
Wang Shuai,Liu Feng-zhi,Wang Hai-bo.2015.Study on leaf senescence physiology and anti-aging technology of grape cultured in protected and delayed cultivation[M.D.Dissertation].Beijing:Chinese Academy of Agricultural Sciences.(in Chinese)王帅,刘凤之,王海波.2015.设施葡萄延迟栽培叶片衰老生理及抗衰老技术研究[硕士论文].北京:中国农业科学院.
Wang Shuai,Wang Xiaodi,Shi Xiangbin,Wang Baoliang,Zheng Xiaocui,Wang Haibo,Liu Fengzhi.2016.Red and blue lights significantly affect photosynthetic properties and ultrastructure of mesophyll cells in senescing grape leaves.Horticultural Plant Journal,2(2):82-90.
Wen Han,Nie Fan.2000.Effects of drought on senescence of flag leaf and yield factors in rice.Journal of Anhui Agricultural University,27(2):135-137.(in Chinese)文汉,聂凡.2000.干旱对水稻抽穗后旗叶衰老和产量构成因子的影响.安徽农业大学学报,27(2):135-137.
Xiong Bing-lin,Wang Shi-wen,Wang Xin-yue,Chen Dao-qian,Yin Li-na,Deng Xi-ping.2016.Effects of nitrogenous fertilizer on leaf senescence of maize and the associate with carbon/nitrogen balance under drought stress.Journal of Maize Sciences,24(3):138-146.(in Chinese)熊炳霖,王仕稳,王鑫月,陈道钳,殷俐娜,邓西平.2016.干旱胁迫下氮肥对玉米叶片衰老影响及与碳氮平衡的关系.玉米科学,24(3):138-146.
Xu Chao-hua,Li Jun-ying,Cui Ming-kun,Ma Er-dong,Huang Guo-bin,Gong Ming.2013.Effects of supplemental lighting on growth and photo-synthesis of tobacco leaves.Acta Bot Boreal-Occient Sin,33(4):763-770.(in Chinese)徐超华,李军营,崔明昆,马二登,黄国宾,龚明.2013.延长光照时间对烟草叶片生长发育及光合特性的影响.西北植物学报,33(4):763-770.
Xu Li,Liu Shi-qi,Qi Lian-dong,Liang Qing-ling,Yu Wen-yan.2007.Effect of light quality on leaf lettuce photosynthesis and chlorophyll fluorescence.Chinese Agricultural Science Bulletin,23(1):96-100.(in Chinese)许莉,刘世琦,齐连东,梁庆玲,于文艳.2007.不同光质对叶用莴苣光合作用及叶绿素荧光的影响.中国农学通报,23(1):96-100.
Yan Xiao-hua,Yu Ji-hua.2016.Effects of supplemental LED light on photosynthetic pigment contentsand antioxidant enzyme activities of cucumber seedling leaves.Journal of Desert Research,36(2):392-398.(in Chinese)闫晓花,郁继华.2016.LED补光对温室黄瓜幼苗抗衰老及抗氧化酶系统的影响.中国沙漠,36(2):392-398.
Zhai Rong-rong,Feng Yue,Cao Li-yong,Cheng Shi-hua,Wu Wei-ming.2011.Review of rice leaf senescence.Chinese Rice,17(1):7-12.(in Chinese)翟荣荣,冯跃,曹立勇,程式华,吴伟明.2011.水稻叶片衰老研究进展.中国稻米,17(1):7-12.
Zhang Jie,Li Tian-lai.2005.Effects of daytime sub-high temperature on photosynthesis and chloroplast ultrastructure of tomato leaves in greenhouse.Acta Horticulturae Sinica,32(4):614-619.(in Chinese)张洁,李天来.2005.日光温室亚高温对番茄光合作用及叶绿体超微结构的影响.园艺学报,32(4):614-619.
Zheng Shao-qing,Zeng Guang-wen.1990.Senescence of plant leaf and the molecular approach for its delaying.Plant Physiology Journal,35(1):152-157.(in Chinese)郑少青,曾广文.1990.植物叶片衰老及其延缓的分子途径.植物生理学通讯,35(1):152-157.
Zhu Xue-mei,Ke Yong-pei,Shao Ji-rong,Lin Li-jin,Yang Yuan-xiang.2005.Effects of high temperature stress on activated oxygen metabolism of the leaves of the heavy panicle type of medium india hybrid rice.Seed,24(3):25-27,32.(in Chinese)朱雪梅,柯永培,邵继荣,林立金,杨远祥.2005.高温胁迫对重穗型水稻品种叶片活性氧代谢的影.种子,24(3):25-27,32.
Zimmermann P,Zentgraf U.2005.The correlation between oxidative stress and leaf senescence during plant development.Cellular and Molecular Biology Letters,10(3):515-534.
Zou Qi.2000.Experimental guidance of plant physiology.Beijing:Chinese Agricultural Press:72-75.(in Chinese)邹琦.2000.植物生理学实验指导.北京:中国农业出版社:72-75.