青藏高原和内蒙古高原典型草地植物叶片肾型和哑铃型气孔器气孔特征及其与环境的关系
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摘要
该研究利用植物制片技术,以中国青藏高原和内蒙古高原典型草地常见种或优势种植物的叶片为研究对象,通过比较分析叶片哑铃型气孔器和肾型气孔器的特征及其与环境因子的关系,揭示植物叶片两类气孔器对环境因子的响应策略。结果表明:(1)叶片哑铃型气孔器气孔指标的变异系数小于肾型气孔器。(2)叶片哑铃型气孔器气孔指标与环境气候指标的关系弱于肾型气孔器。(3)叶片哑铃型气孔器气孔特征与环境关系在叶片上下表面之间存在显著差异,而肾型气孔器气孔特征与环境因子的关系在叶片上下表面之间无显著差异。(4)叶片哑铃型气孔器的气孔特征与降水关系密切,而肾型气孔器气孔特征与温度关系密切。(5)同一种气孔器的气孔特征在两个地区(青藏高原和内蒙古高原)间存在显著差异。研究认为,肾型气孔器和哑铃型气孔器的气孔特征及其与环境之间的关系存在差异,在分析气孔特征时有必要将肾形与哑铃形保卫细胞形成的气孔器加以区分,该研究结果有助于进一步理解中国草地植物叶气孔特征对气候变化的响应与适应策略。
The stomatal characters of dominant grassland species in the Tibetan Plateau and Inner Mongolia Plateau were investigated by using plant micro-techniques.We aimed to disentangle the stomatal characters and their relationships with environmental factors and reveal the adaptation strategies in two types of stomatal apparatus,which are composed of kidney-shaped and dumb-bell-shaped guard cells.The results showed:(1)the coefficient of variation of stomatal characters of dumb-bell-shaped stomatal apparatus was smaller than that of kidney-shaped stomatal apparatus.(2)The relationships between stomatal characters of dumb-bell-shaped stomatal apparatus and environmental factors were weaker than that of kidney-shaped stomatal apparatus.(3)The environmental factors showed different effects on the adaxial and abaxial stomatal characters in dumb-bell-shaped stomatal apparatus,while in kidney-shaped stomatal apparatusthere were no different effects on the adaxial and abaxial stomatal characters.(4)The stomatal characters were closely correlated to precipitation in dumb-bell-shaped stomatal apparatus,while in kidney-shaped stomatal apparatus they were closely correlated to temperature.(5)The stomatal characters of the same stomata had significant differences in the two regions(Tibetan and Inner Mongolia Plateau).We came to the conclusion that there were significant differences in stomatal characters and the relationships with environmental factors between kidney-shaped and dumb-bell-shaped stomatal apparatus.The results indicate that it is necessary to distinguish the types of stomata when analyzing stomatal characters,and will be helpful for further understanding the adaptation strategies of leaf stomatal characters of grassland plants in China under the background of global climate change.
The stomatal characters of dominant grassland species in the Tibetan Plateau and Inner Mongolia Plateau were investigated by using plant micro-techniques.We aimed to disentangle the stomatal characters and their relationships with environmental factors and reveal the adaptation strategies in two types of stomatal apparatus,which are composed of kidney-shaped and dumb-bell-shaped guard cells.The results showed:(1)the coefficient of variation of stomatal characters of dumb-bell-shaped stomatal apparatus was smaller than that of kidney-shaped stomatal apparatus.(2)The relationships between stomatal characters of dumb-bell-shaped stomatal apparatus and environmental factors were weaker than that of kidney-shaped stomatal apparatus.(3)The environmental factors showed different effects on the adaxial and abaxial stomatal characters in dumb-bell-shaped stomatal apparatus,while in kidney-shaped stomatal apparatusthere were no different effects on the adaxial and abaxial stomatal characters.(4)The stomatal characters were closely correlated to precipitation in dumb-bell-shaped stomatal apparatus,while in kidney-shaped stomatal apparatus they were closely correlated to temperature.(5)The stomatal characters of the same stomata had significant differences in the two regions(Tibetan and Inner Mongolia Plateau).We came to the conclusion that there were significant differences in stomatal characters and the relationships with environmental factors between kidney-shaped and dumb-bell-shaped stomatal apparatus.The results indicate that it is necessary to distinguish the types of stomata when analyzing stomatal characters,and will be helpful for further understanding the adaptation strategies of leaf stomatal characters of grassland plants in China under the background of global climate change.
引文
[1]WILLMER C M,FRICKER M.Stomata[M].London:Springer.Chapman&Hall,1996.
[2]ROELFSEMA M R,KOLLIST H.Tiny pores with a global impact[J].New Phytologist,2013,197(1):11-15.
[3]HETHERINGTON A M,WOODWARD F I.The role of stomata in sensing and driving environmental change[J].Nature,2003,424(6 951):901-908.
[4]LAKE J A,QUICK W P,BEERLING D J,et al.Plant development:signals from mature to new leaves[J].Nature,2001,411(6 834):154.
[5]BUCKLEY T N.The control of stomata by water balance[J].New Phytologist,2005,168(2):275-292.
[6]XU Z Z,ZHOU G S.Responses of leaf stomatal density to water status and its relationship with photosynthesis in a grass[J].Journal of Experimental Botany,2008,59(12):3 317-3 325.
[7]LUOMALA E M,LAITINEN K,SUTINEN S,et al.Stomatal density,anatomy and nutrient concentrations of Scots pine needles are affected by elevated CO2 and temperature[J].Plant,Cell&Environment,2005,28(6):733-749.
[8]FRASER L H,GREENALL A,CARLYLE C,et al.Adaptive phenotypic plasticity of Pseudoroegneria spicata:response of stomatal density,leaf area and biomass to changes in water supply and increased temperature[J].Annals of Botany,2009,103(5):769-775.
[9]WOODWARD F I.Stomatal numbers are sensitive to increases in CO2from pre-industrial levels[J].Nature,1987,327(6 123):617-618.
[10]FRANKS P J,BEERLING D J.Maximum leaf conductance driven by CO2effects on stomatal size and density over geologic time[J].Proceedings of the National Academy of Sciences of the United States of America,2009,106(25):10 343-10 347.
[11]MOTT K A.Opinion:stomatal responses to light and CO2depend on the mesophyll[J].Plant,Cell&Environment,2009,32(11):1 479-1 486.
[12]KRNER C,BANNISTER P,MARK A F.Altitudinal variation in stomatal conduncance,nitrogen content and leaf anatomy in different plant life forms in New Zealand[J].Oecologia,1986,69(4):577-588.
[13]QIANG W Y,WANG X L,CHEN T,et al.Variation of stomatal density and carbon isotope values of Picea crassifolia at different altitudes in the Qilian Mountains[J].Trees,2003,17(3):258-262.
[14]KESSLER M,SIORAK Y,WUNDERLICH M,et al.Patterns of morphological leaf traits among pteridophytes along humidity and temperature gradients in the Bolivian Andes.[J].Functional Plant Biology,2007,34(11):963-971.
[15]CASSON S A,HETHERINGTON A M.Environmental regulation of stomatal development[J].Current Opinion in Plant Biology,2010,13(1),90-95.
[16]CASSON S,GRAY J E.Influence of environmental factors on stomatal development[J].New Phytologist,2008,178(1),9-23.
[17]HERNANDEZ M L,PASSAS H J,SMITH L G.Clonal analysis of epidermal patterning during maize leaf development[J].Developmental Biology,1999,216(2):646-658.
[18]陈佐忠.中国典型草原生态系统[M].北京:科学出版社,2000:9-11.
[19]MA J J,JI C J,HAN M,et al.Comparative analyses of leaf anatomy of dicotyledonous species in Tibetan and Inner Mongolian grasslands[J].Science China Life Sciences,2012,55(1):68-79.
[20]中国科学院内蒙古宁夏综合考察队.内蒙古植被[M].北京:科学出版社,1985:4-9.
[21]杨元合,朴世龙.青藏高原草地植被覆盖变化及其与气候因子的关系[J].植物生态学报,2006,30(1):1-8.YANG Y H,PIAO S L.Variations in grassland vegetation cover in relation to climatic factors on the Tibetan Plateau[J].Journal of Plant Ecology,2006,30(1):1-8.
[22]封涛,胡东.单子叶植物叶片气孔观察方法改良的研究[J].植物研究,2008,28(1):82-84.FENG T,HU D.An improving method for observation of the stomata on monocotyledon[J].Bulletin of Botanical Research,2008,28(1):82-84.
[23]RASHKE K.Movement of stomata[J].Encyclopedia of Plant Physiology,1979,7:383-441.
[24]TIAN M,YU G R,HE N P,et al.Leaf morphological and anatomical traits from tropical to temperate coniferous forests:mechanisms and influencing factors[J].Scientific Reports,2016,6(6):19 703.
[25]HAWORTH M,ELLIOTT-KINGSTON C,MCELWAIN J C.Stomatal control as a driver of plant evolution[J].Journal of Experimental Botany,2011,62(8):2 419-2 423.
[26]DRAKE P L,FROEND R H,FRANKS P J.Smaller,faster stomata:scaling of stomatal size,rate of response,and stomatal conductance[J].Journal of Experimental Botany,2013,64(2):495-505.
[27]费松林,方精云,樊拥军,等.贵州梵净山亮叶水青冈叶片和木材的解剖学特征及其与生态因子的关系[J].植物学报(英文版),1999,41(9):1 002-1 009.FEI S L,FANG J Y,FAN Y J,et al.Anatomical characteristics of leaves and woods of Fagus lucida and their relationship to ecological factors in Mountain Fanjingshan,Guizhou,China[J].Acta Botanica Sinica,1999,41(9):1 002-1 009.
[28]KRNER C.Alpine plant life:functional plant ecology of high mountain ecosystems[J].Mountain Research&Development,1999,21(2):202.
[29]施海燕,王一峰,王剑虹.青藏高原不同海拔地7种风毛菊属植物叶片解剖结构的研究[J].植物研究,2009,29(3):289-294.SHI H Y,WANG Y F,WANG J H.Studies on the anatomical structures of leaves of seven Saussurea D C.Species on the Qinghai-Tibetan plateau with different altitude[J].Bulletin of Botanical Research,2009,29(3):289-294.
[2]ROELFSEMA M R,KOLLIST H.Tiny pores with a global impact[J].New Phytologist,2013,197(1):11-15.
[3]HETHERINGTON A M,WOODWARD F I.The role of stomata in sensing and driving environmental change[J].Nature,2003,424(6 951):901-908.
[4]LAKE J A,QUICK W P,BEERLING D J,et al.Plant development:signals from mature to new leaves[J].Nature,2001,411(6 834):154.
[5]BUCKLEY T N.The control of stomata by water balance[J].New Phytologist,2005,168(2):275-292.
[6]XU Z Z,ZHOU G S.Responses of leaf stomatal density to water status and its relationship with photosynthesis in a grass[J].Journal of Experimental Botany,2008,59(12):3 317-3 325.
[7]LUOMALA E M,LAITINEN K,SUTINEN S,et al.Stomatal density,anatomy and nutrient concentrations of Scots pine needles are affected by elevated CO2 and temperature[J].Plant,Cell&Environment,2005,28(6):733-749.
[8]FRASER L H,GREENALL A,CARLYLE C,et al.Adaptive phenotypic plasticity of Pseudoroegneria spicata:response of stomatal density,leaf area and biomass to changes in water supply and increased temperature[J].Annals of Botany,2009,103(5):769-775.
[9]WOODWARD F I.Stomatal numbers are sensitive to increases in CO2from pre-industrial levels[J].Nature,1987,327(6 123):617-618.
[10]FRANKS P J,BEERLING D J.Maximum leaf conductance driven by CO2effects on stomatal size and density over geologic time[J].Proceedings of the National Academy of Sciences of the United States of America,2009,106(25):10 343-10 347.
[11]MOTT K A.Opinion:stomatal responses to light and CO2depend on the mesophyll[J].Plant,Cell&Environment,2009,32(11):1 479-1 486.
[12]KRNER C,BANNISTER P,MARK A F.Altitudinal variation in stomatal conduncance,nitrogen content and leaf anatomy in different plant life forms in New Zealand[J].Oecologia,1986,69(4):577-588.
[13]QIANG W Y,WANG X L,CHEN T,et al.Variation of stomatal density and carbon isotope values of Picea crassifolia at different altitudes in the Qilian Mountains[J].Trees,2003,17(3):258-262.
[14]KESSLER M,SIORAK Y,WUNDERLICH M,et al.Patterns of morphological leaf traits among pteridophytes along humidity and temperature gradients in the Bolivian Andes.[J].Functional Plant Biology,2007,34(11):963-971.
[15]CASSON S A,HETHERINGTON A M.Environmental regulation of stomatal development[J].Current Opinion in Plant Biology,2010,13(1),90-95.
[16]CASSON S,GRAY J E.Influence of environmental factors on stomatal development[J].New Phytologist,2008,178(1),9-23.
[17]HERNANDEZ M L,PASSAS H J,SMITH L G.Clonal analysis of epidermal patterning during maize leaf development[J].Developmental Biology,1999,216(2):646-658.
[18]陈佐忠.中国典型草原生态系统[M].北京:科学出版社,2000:9-11.
[19]MA J J,JI C J,HAN M,et al.Comparative analyses of leaf anatomy of dicotyledonous species in Tibetan and Inner Mongolian grasslands[J].Science China Life Sciences,2012,55(1):68-79.
[20]中国科学院内蒙古宁夏综合考察队.内蒙古植被[M].北京:科学出版社,1985:4-9.
[21]杨元合,朴世龙.青藏高原草地植被覆盖变化及其与气候因子的关系[J].植物生态学报,2006,30(1):1-8.YANG Y H,PIAO S L.Variations in grassland vegetation cover in relation to climatic factors on the Tibetan Plateau[J].Journal of Plant Ecology,2006,30(1):1-8.
[22]封涛,胡东.单子叶植物叶片气孔观察方法改良的研究[J].植物研究,2008,28(1):82-84.FENG T,HU D.An improving method for observation of the stomata on monocotyledon[J].Bulletin of Botanical Research,2008,28(1):82-84.
[23]RASHKE K.Movement of stomata[J].Encyclopedia of Plant Physiology,1979,7:383-441.
[24]TIAN M,YU G R,HE N P,et al.Leaf morphological and anatomical traits from tropical to temperate coniferous forests:mechanisms and influencing factors[J].Scientific Reports,2016,6(6):19 703.
[25]HAWORTH M,ELLIOTT-KINGSTON C,MCELWAIN J C.Stomatal control as a driver of plant evolution[J].Journal of Experimental Botany,2011,62(8):2 419-2 423.
[26]DRAKE P L,FROEND R H,FRANKS P J.Smaller,faster stomata:scaling of stomatal size,rate of response,and stomatal conductance[J].Journal of Experimental Botany,2013,64(2):495-505.
[27]费松林,方精云,樊拥军,等.贵州梵净山亮叶水青冈叶片和木材的解剖学特征及其与生态因子的关系[J].植物学报(英文版),1999,41(9):1 002-1 009.FEI S L,FANG J Y,FAN Y J,et al.Anatomical characteristics of leaves and woods of Fagus lucida and their relationship to ecological factors in Mountain Fanjingshan,Guizhou,China[J].Acta Botanica Sinica,1999,41(9):1 002-1 009.
[28]KRNER C.Alpine plant life:functional plant ecology of high mountain ecosystems[J].Mountain Research&Development,1999,21(2):202.
[29]施海燕,王一峰,王剑虹.青藏高原不同海拔地7种风毛菊属植物叶片解剖结构的研究[J].植物研究,2009,29(3):289-294.SHI H Y,WANG Y F,WANG J H.Studies on the anatomical structures of leaves of seven Saussurea D C.Species on the Qinghai-Tibetan plateau with different altitude[J].Bulletin of Botanical Research,2009,29(3):289-294.