堇菜叶片草酸钙晶体与水分维持的关系
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摘要
随着全球气候变化加重,干旱强度和持续时间逐渐增加,严重影响植物生长和作物产量。喀斯特为典型的干旱和高钙生境,植物叶片富集大量的草酸钙晶体,而该晶体与植物耐旱性之间的关系并不清楚。该研究以喀斯特适生植物堇菜(Viola verecumda)为材料,土壤进行自然干旱,分析堇菜叶片的草酸钙晶体变化特征与水分之间的关系。结果表明:在土壤自然干旱条件下,堇菜主要通过细胞内束缚水的释放,维持细胞内水分平衡;而在干旱后期,叶片通过关闭气孔,将部分自由水转变为束缚水,防止水分流失。此外,草酸钙晶体的密度与束缚水含量具有极其显著的强正相关线性回归关系(r=0.825 3,P<0.000 1),表明草酸钙晶体作为主要的束缚水物质。因此,堇菜植物在耐旱过程中可能协调草酸钙晶体和气孔的生理行为忍耐干旱胁迫。
As the global climate changes dramatically,the increasing intensity and duration of drought seriously affects plant growth and crop yield. Karst is a typical arid and high-calcium habitat,and plant leaves in karst areas contain a large amount of calcium oxalate crystals,while the relationship between the crystal and the drought tolerance of plants is not clear. In this study,we took Viola verecumda grown in karst area as material,and analyzed the relationship between the changing characteristics of calcium oxalate cyrstals and water in its leaves,by subjecting the leaves to natural dry soil. The results showed that in the early stage of drought stress,V. verecumda maintained the intracellular water balance mainly through the release of intracellular bound water. In the later stage of drought,the leaves mainly prevented water loss by closing the stoma and converting part of the free water into bound water. In addition,the density of calcium oxalate crystals had an significant positive correlation with the bound water content( r = 0. 825 3,P<0. 000 1),suggesting that calcium oxalate crystals act as the main matter binding water. Therefore,V. verecumda may coordinate the physiological behavior of calcium oxalate crystals and stomatal during drought tolerance.
As the global climate changes dramatically,the increasing intensity and duration of drought seriously affects plant growth and crop yield. Karst is a typical arid and high-calcium habitat,and plant leaves in karst areas contain a large amount of calcium oxalate crystals,while the relationship between the crystal and the drought tolerance of plants is not clear. In this study,we took Viola verecumda grown in karst area as material,and analyzed the relationship between the changing characteristics of calcium oxalate cyrstals and water in its leaves,by subjecting the leaves to natural dry soil. The results showed that in the early stage of drought stress,V. verecumda maintained the intracellular water balance mainly through the release of intracellular bound water. In the later stage of drought,the leaves mainly prevented water loss by closing the stoma and converting part of the free water into bound water. In addition,the density of calcium oxalate crystals had an significant positive correlation with the bound water content( r = 0. 825 3,P<0. 000 1),suggesting that calcium oxalate crystals act as the main matter binding water. Therefore,V. verecumda may coordinate the physiological behavior of calcium oxalate crystals and stomatal during drought tolerance.
引文
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MACNISH AJ,IRVING DE,JOYCE DC,et al.,2003.Identification of intracellular calcium oxalate crystals in Chamelaucium uncinatum(Myrtaceae)[J].Aust J Bot,51(5):565-572.
PENG XL,2011.The droughe tolerance of three gesneriaceae species in danxia landscape[D].Xiamen:Xiamen University:1-53.[彭小玲,2011.丹霞地貌3种苦苣苔科植物的耐旱性研究[D].厦门:厦门大学:1-53.]
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TOOULAKOU G,GIANNOPOULOS A,NIKOLOPOULOS D,et al.,2016.Alarm photosynthesis:Calcium oxalate crystals as an internal CO2source in plants[J].Plant Physiol,171(4):2577-2585.
TURNER NC,WRIGHT GC,SIDDIQUE KHM,2001.Adaptation of grain legumes(pulses)to water-limited environments[J].Advan Agron,71(1):193-231.
VASILYEVA HT,DONG RR,1956.The relationship between the ratio of free water and bound water in plant leaves and drought resistance of plants[J].Plant Physiol J,2(3):50-53.[瓦西里也娃,董任瑞,1956.论植物叶片中自由水和束缚水的比例与植物抗旱性的关系[J].植物生理学报,2(3):50-53.]
WARD D,SPIEGEL M,SALTZ D,1997.Gazelle herbivory and interpopulation differences in calcium oxalate content of leaves of a desert lily[J].J Chem Ecol,23(2):333-346.
WANG DL,ZHU SQ,HUANG BL,2005.Preliminary study on types and quantitative assessment of karst rocky desertification in Guizhou Province,China[J].Acta Ecol Sin,25(5):115-121.[王德炉,朱守谦,黄宝龙,2005.贵州喀斯特石漠化类型及程度评价[J].生态学报,25(5):115-121.]
WU LL,WANG DL,LI ZY,2018.Physiological response of Ligustrum robustum to continuous drought and rewatering[J].Plant Physiol J,54(5):837-844.[吴丽丽,王德炉,李自玉,2018.粗壮女贞对持续性干旱及复水的生理响应[J].植物生理学报,54(5):837-844.]
XU GB,ZENG WZ,WAN DG,1995.Plant resources of ciola and prospects of exploitation in Sichuan Province[J].Chin Med Mat,18(5):227-231.[徐国兵,曾万章,万德光,1995.四川省堇菜属药用植物资源与开发前景[J].中药材,18(5):227-231.]
YI Y,SU ZM,ZHANG XM,et al.,2018.Calcium oxalate biosynthesis in plant and its defense functions[J].J Guizhou Norm Univ(Nat Sci Ed),36(4):1-7.[乙引,苏志孟,张习敏,等,2018.草酸钙在植物中的生物合成及其防御功能[J].贵州师范大学学报(自然科学版),36(4):1-7.]
ZHANG LG,ZENG FJ,LIU B,et al.,2012.Study of the photosynthesis characteristics and physical signs of four plants at the desert-oasis ecotone[J].Acta Pratac Sin,21(1):103-111.[张利刚,曾凡江,刘波,等,2012.绿洲-荒漠过渡带四种植物光合及生理特征的研究[J].草业学报,21(1):103-111.]
ZHU GL,MA Y,HAN L,et al.,2014.Current status of research on morphological structure,biological function and formation mechanism of plant crystals[J].Acta Ecol Sin,34(22):6429-6439.[朱广龙,马茵,韩蕾,等,2014.植物晶体的形态结构、生物功能及形成机制研究进展[J].生态学报,34(22):6429-6439.]
ZHU GL,WEI XZ,2016.Leaf morphological plasticity of Ziziphus jujuba var.spinosa in response to natural drought gradient ecotopes[J].Acta Ecol Sin,36(19):6178-6187.[朱广龙,魏学智,2016.酸枣叶片结构可塑性对自然梯度干旱生境的适应特征[J].生态学报,36(19):6178-6187.]
CHEN L,YUAN HJ,WEI SZ,2008.Analysis of the bound water/free water and transpiration intensity of Sedum plants[J].Agric Technol,28(6):80-81.[陈莉,原红娟,尉淑珍,2008.景天植物的束缚水/自由水与蒸腾强度的分析[J].农业与技术,28(6):80-81.]
FANG Y,XIONG L,2015.General mechanisms of drought response and their application in drought resistance improvement in plants[J].Cell Mol Life Sci,72(4):673-689.
FRANCESCHI VR,NAKATA PA,2005.Calcium oxalate in plants:Formation and function[J].Ann Rev Plant Biol,56(1):41-71.
GUO ZC,ZENG FJ,LIU B,et al.,2015.Photosynthetic characteristics of Alhagi sparsifolia and Tamarix ramosissima and the relevant environment factors in different periods[J].Acta Bot Boreal-Occident Sin,35(1):189-198.[郭自春,曾凡江,刘波,等,2015.疏叶骆驼刺和多枝柽柳不同时期光合特性日变化及其与环境因子的关系[J].西北植物学报,35(1):189-198.]
LAWLOR DW,2013.Genetic engineering to improve plant performance under drought:Physiological evaluation of achievements,limitations,and possibilities[J].J Exp Bot,64(1):83-108.
LUO LJ,2010.Breeding for water-saving and drought-resistance rice(WDR)in China[J].J Exp Bot,61(13):3509-3517.
LI YL,2008.Medicinal plant resources of viola and its utiliazation in Shandong Province[J].N Hortic,32(1):237-238.[李彦连,2008.山东省堇菜属药用植物资源及利用[J].北方园艺,32(1):237-238.]
LI Y,JIANG TM,WANG J,2009.Seasonal drought characteristic of guizhou karst mountainous area and its counterneasures:Taking Tongzi County as the example[J].Guizhou Agric Sci,37(5):43-46.[李渝,蒋太明,王静,2009.贵州喀斯特山区季节性干旱特征及对策:以桐梓县为例[J].贵州农业科学,37(5):43-46.]
LIU JQ,1982.The xeromorphic structure of different typical plants in deserts of China[J].Chin J Plant Ecol,6(4):314-319.[刘家琼,1982.我国荒漠不同生态类型植物的旱生结构[J].植物生态学报,6(4):314-319.]
LIU XD,LI XH,LI WH,et al.,2004.Analysis on difference for drought responses of maize inbred lines at seedling stage[J].J Maize Sci,12(3):63-65.[刘贤德,李晓辉,李文华,等,2004.玉米自交系苗期耐旱性差异分析[J].玉米科学,12(3):63-65.]
MACNISH AJ,IRVING DE,JOYCE DC,et al.,2003.Identification of intracellular calcium oxalate crystals in Chamelaucium uncinatum(Myrtaceae)[J].Aust J Bot,51(5):565-572.
PENG XL,2011.The droughe tolerance of three gesneriaceae species in danxia landscape[D].Xiamen:Xiamen University:1-53.[彭小玲,2011.丹霞地貌3种苦苣苔科植物的耐旱性研究[D].厦门:厦门大学:1-53.]
RASCIO A,PLATANI C,FONZO N,et al.,1992.Bound water in durum wheat under drought stress[J].Plant Physiol,98(3):908-912.
SHAO YJ,SHAN L,2006.Advances in the research on the mechanism of plants[J].Chin J Eco-Agric,14(4):16-20.[邵艳军,山仑,2006.植物耐旱机制研究进展[J].中国生态农业学报,14(4):16-20.]
TOOULAKOU G,GIANNOPOULOS A,NIKOLOPOULOS D,et al.,2016.Alarm photosynthesis:Calcium oxalate crystals as an internal CO2source in plants[J].Plant Physiol,171(4):2577-2585.
TURNER NC,WRIGHT GC,SIDDIQUE KHM,2001.Adaptation of grain legumes(pulses)to water-limited environments[J].Advan Agron,71(1):193-231.
VASILYEVA HT,DONG RR,1956.The relationship between the ratio of free water and bound water in plant leaves and drought resistance of plants[J].Plant Physiol J,2(3):50-53.[瓦西里也娃,董任瑞,1956.论植物叶片中自由水和束缚水的比例与植物抗旱性的关系[J].植物生理学报,2(3):50-53.]
WARD D,SPIEGEL M,SALTZ D,1997.Gazelle herbivory and interpopulation differences in calcium oxalate content of leaves of a desert lily[J].J Chem Ecol,23(2):333-346.
WANG DL,ZHU SQ,HUANG BL,2005.Preliminary study on types and quantitative assessment of karst rocky desertification in Guizhou Province,China[J].Acta Ecol Sin,25(5):115-121.[王德炉,朱守谦,黄宝龙,2005.贵州喀斯特石漠化类型及程度评价[J].生态学报,25(5):115-121.]
WU LL,WANG DL,LI ZY,2018.Physiological response of Ligustrum robustum to continuous drought and rewatering[J].Plant Physiol J,54(5):837-844.[吴丽丽,王德炉,李自玉,2018.粗壮女贞对持续性干旱及复水的生理响应[J].植物生理学报,54(5):837-844.]
XU GB,ZENG WZ,WAN DG,1995.Plant resources of ciola and prospects of exploitation in Sichuan Province[J].Chin Med Mat,18(5):227-231.[徐国兵,曾万章,万德光,1995.四川省堇菜属药用植物资源与开发前景[J].中药材,18(5):227-231.]
YI Y,SU ZM,ZHANG XM,et al.,2018.Calcium oxalate biosynthesis in plant and its defense functions[J].J Guizhou Norm Univ(Nat Sci Ed),36(4):1-7.[乙引,苏志孟,张习敏,等,2018.草酸钙在植物中的生物合成及其防御功能[J].贵州师范大学学报(自然科学版),36(4):1-7.]
ZHANG LG,ZENG FJ,LIU B,et al.,2012.Study of the photosynthesis characteristics and physical signs of four plants at the desert-oasis ecotone[J].Acta Pratac Sin,21(1):103-111.[张利刚,曾凡江,刘波,等,2012.绿洲-荒漠过渡带四种植物光合及生理特征的研究[J].草业学报,21(1):103-111.]
ZHU GL,MA Y,HAN L,et al.,2014.Current status of research on morphological structure,biological function and formation mechanism of plant crystals[J].Acta Ecol Sin,34(22):6429-6439.[朱广龙,马茵,韩蕾,等,2014.植物晶体的形态结构、生物功能及形成机制研究进展[J].生态学报,34(22):6429-6439.]
ZHU GL,WEI XZ,2016.Leaf morphological plasticity of Ziziphus jujuba var.spinosa in response to natural drought gradient ecotopes[J].Acta Ecol Sin,36(19):6178-6187.[朱广龙,魏学智,2016.酸枣叶片结构可塑性对自然梯度干旱生境的适应特征[J].生态学报,36(19):6178-6187.]