叶生态特征及其相关性对下垫面热效应的生态权衡
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摘要
为研究植物叶性状及其相关关系对城市下垫面热效应的响应及其生态权衡策略,以生长在6种典型城市下垫面(沥青、水泥、荷兰砖、大理石、嵌草砖和自然地表)环境中的3种常见绿化树种(洋白蜡、国槐和臭椿)为研究对象,测定其叶功能性状指标。结果表明:比叶面积、叶绿素含量、气孔面积、气孔开度及叶氮磷比均随着环境地表温度的增大总体上呈现先增加、后减小的趋势,城市下垫面从大到小依次为嵌草砖地表、自然地表、大理石地表、水泥地表、荷兰砖地表、沥青地表。而叶干物质含量、叶组织密度、气孔密度及叶脉密度、叶氮含量、叶磷含量则呈相反的变化规律,城市下垫面从大到小依次为沥青地表、荷兰砖地表、水泥地表、大理石地表、自然地表、嵌草砖地表。叶性状间表现了一定的相关关系,其中,比叶面积与叶干物质含量、叶组织密度与气孔密度间呈极显著的负相关关系(P <0. 01),与叶氮含量、叶磷含量呈显著的正相关关系(P <0. 01)。叶干物质含量与叶组织密度、叶脉密度和叶氮含量存在极显著的正相关关系(P <0. 01)。气孔密度与气孔面积、气孔开度间存在显著的负相关关系(P <0. 05),而与叶脉密度间存在显著的正相关关系(P <0. 05)。气孔面积与叶氮含量间存在显著的正相关关系(P <0. 05),与叶组织密度呈极显著的正相关关系(P <0. 01)。随着地表温度的升高,植物叶性状间的相关性呈现先增、后减的趋势,小幅度升温总体上有利于加强叶功能性状间的相关性,而温度过高则明显降低其相关关系。城市生态系统中,植物叶片具有低的比叶面积、叶绿素含量、气孔面积、气孔开度和叶氮磷比,高的叶干物质含量、叶组织密度、气孔密度、叶氮含量和叶磷含量,进一步验证了城市生态系统中叶经济谱的存在,并在该谱系中属于"快速投资-收益"型。
The object was to study the response of plant leaf traits and their related relationships to the thermal environment effect of urban underlying surface and its ecological trade-off strategy. Results showed that specific leaf area,chlorophyll content,stomatal area,stomatal aperture and leaf nitrogenphosphorus ratio were all increased firstly and then decreased with the increase of environmental surface temperature,which was expressed as grass-planting bricks surface,grass surface,marble surface,cement surface,brick surface,asphalt surface from large to small. While the leaf dry matter content,leaf tissue density,stomatal density,leaf nitrogen content,leaf phosphorus content and leaf vein density showed opposite changes,which were expressed as asphalt surface,brick surface,cement surface,marble surface,grass surface and grass-planting brick surface from large to small. There was a certain correlation between plant leaf functional traits. Specific leaf area showed a significant negative correlation with leaf dry matter content,leaf tissue density and stomatal density(P < 0. 01),and a significantpositive correlation with leaf nitrogen content and leaf phosphorus content(P < 0. 01). Leaf dry matter content showed a significant positive correlation with leaf tissue density,leaf vein density and leaf nitrogen content(P < 0. 01). Stomatal density was significantly and negatively correlated with stomatal aperture and stomatal area(P < 0. 05),and significantly and positively correlated with leaf vein density(P < 0. 05). Leaf vein density was significantly and positively correlated with leaf nitrogen content(P <0. 05) and leaf tissue density(P < 0. 01). With the increase of surface temperature,the correlation between plant leaf traits was firstly increased and then decreased. The small temperature increase was generally beneficial to strengthen the correlation between leaf functional traits,which significantly reduced its correlation relationship when the temperature was too high. In urban ecosystems,plant leaves showed low specific leaf area,chlorophyll content,stomatal area,stomatal aperture and leaf nitrogen-phosphorus ratio,and high leaf dry matter content,leaf tissue density,stomatal density,leaf nitrogen content and leaf phosphorus content. The research further verified the existence of the economics spectrum of the lower temperature of high temperature stress habitat on the underlying surface of the city,and it was a"quick investment-return"type in the global leaf economics spectrum,which provided an important theoretical basis for the selection and configuration of urban greening plants and urban planning. The study further validated the existence of economics spectrum in the urban system,which was a "quick investment-benefit"type.
The object was to study the response of plant leaf traits and their related relationships to the thermal environment effect of urban underlying surface and its ecological trade-off strategy. Results showed that specific leaf area,chlorophyll content,stomatal area,stomatal aperture and leaf nitrogenphosphorus ratio were all increased firstly and then decreased with the increase of environmental surface temperature,which was expressed as grass-planting bricks surface,grass surface,marble surface,cement surface,brick surface,asphalt surface from large to small. While the leaf dry matter content,leaf tissue density,stomatal density,leaf nitrogen content,leaf phosphorus content and leaf vein density showed opposite changes,which were expressed as asphalt surface,brick surface,cement surface,marble surface,grass surface and grass-planting brick surface from large to small. There was a certain correlation between plant leaf functional traits. Specific leaf area showed a significant negative correlation with leaf dry matter content,leaf tissue density and stomatal density(P < 0. 01),and a significantpositive correlation with leaf nitrogen content and leaf phosphorus content(P < 0. 01). Leaf dry matter content showed a significant positive correlation with leaf tissue density,leaf vein density and leaf nitrogen content(P < 0. 01). Stomatal density was significantly and negatively correlated with stomatal aperture and stomatal area(P < 0. 05),and significantly and positively correlated with leaf vein density(P < 0. 05). Leaf vein density was significantly and positively correlated with leaf nitrogen content(P <0. 05) and leaf tissue density(P < 0. 01). With the increase of surface temperature,the correlation between plant leaf traits was firstly increased and then decreased. The small temperature increase was generally beneficial to strengthen the correlation between leaf functional traits,which significantly reduced its correlation relationship when the temperature was too high. In urban ecosystems,plant leaves showed low specific leaf area,chlorophyll content,stomatal area,stomatal aperture and leaf nitrogen-phosphorus ratio,and high leaf dry matter content,leaf tissue density,stomatal density,leaf nitrogen content and leaf phosphorus content. The research further verified the existence of the economics spectrum of the lower temperature of high temperature stress habitat on the underlying surface of the city,and it was a"quick investment-return"type in the global leaf economics spectrum,which provided an important theoretical basis for the selection and configuration of urban greening plants and urban planning. The study further validated the existence of economics spectrum in the urban system,which was a "quick investment-benefit"type.
引文
1 CORNELISSEN J H C,LAVOREL S,GARNIER E,et al.A handbook of protocols for standardized and easy measurement of plant functional traits worldwide[J].Australian Journal of Botany,2003,51(4):335-380.
2 DIAZ S,CABIDO M.Vive la difference:plant functional diversity matters to ecosystem processes[J].Irish Historical Studies,2001,16(11):646-655.
3 VIOLLE C,NAVAS M L,VILE D,et al.Let the concept of trait be functional[J].Oikos,2007,116(5):882-892.
4 WRIGHT I J,REICH P B,WESTOBY M,et al.The worldwide leaf economics spectrum[J].Nature,2004,428(6985):821.
5 WRIGHT I J,WESTOBY M,REICH P B.Convergence towards higher leaf mass per area in dry and nutrient-poor habitats has different consequences for leaf life span[J].Journal of Ecology,2002,90(3):534-543.
6 DAVID D,ACKERLY D D.Community assembly,niche conservatism,and adaptive evolution in changing environments[J].International Journal of Plant Sciences,2010,164(Supp.3):165-184.
7 SHIPLEY B,LECHOWICZ M J,WRIGHT I,et al.Fundamental trade-offs generating the worldwide leaf economics spectrum[J].Ecology,2006,87(3):535-541.
8 PRIYADARSINI R,HIEN W N,DAVID C K W.Microclimatic modeling of the urban thermal environment of Singapore to mitigate urban heat island[J].Solar Energy,2008,82(8):727-745.
9 DHAKAL S,HANAKI K.Improvement of urban thermal environment by managing heat discharge sources and surface modification in Tokyo[J].Energy&Buildings,2002,34(1):13-23.
10 GORCHAKOV G I,KOPROV B M,SHUKUROV K A.Wind effect on aerosol transport from the underlying surface[J].Izvestiya Atmospheric&Oceanic Physics,2004,40(6):679-694.
11 STONE B,NORMAN J M.Land use planning and surface heat island formation:a parcel-based radiation flux approach[J].Atmospheric Environment,2006,40(19):3561-3573.
12 TAHA H.Meso-urban meteorological and photochemical modeling of heat island mitigation[J].Atmospheric Environment,2008,42(38):8795-8809.
13 KONIJNENDIJK C C,RANDRUP T B.Urban forestry&urban greening[J].Urban Forestry&Urban Greening,2002,1(1):1-4.
14 CASE J L,CROSSON W L,KUMAR S V,et al.Impacts of high resolution land surface initialization on regional sensible weather forecasts from the WRF model[J].Journal of Hydrometeorology,2008,9(6):1249-1266.
15 DASH P,GTTSCHE F M,OLESEN F S,et al.Separating surface emissivity and temperature using two-channel spectral indices and emissivity composites and comparison with a vegetation fraction method[J].Remote Sensing of Environment,2005,96(1):1-17.
16 HURTALOVT,MATEJKA F.Surface characteristics and energy fluxes above different plant canopies[J].Agricultural&Forest Meteorology,1999,99(22):491-499.
17 SALAMANCA F,MARTILLI A.A new building energy model coupled with an urban canopy parameterization for urban climate simulations-part II.validation with one dimension off-line simulations[J].Theoretical and Applied Climatology,2010,99(3/4):345-356.
18 JOHNSON J D,O'CONNOR K A,HANSEN M K,et al.Effects of prior stress on LPS-induced cytokine and sickness responses[J].Am.J.Physiol.Regul.Integr.Comp.Physiol.,2003,284(2):22-32.
19 KATTGE J,DAZ S,LAVOREL S,et al.TRY-a global database of plant traits[J].Global Change Biology,2011,17(9):2905-2935.
20 LOEUILLE N.Consequences of adaptive foraging in diverse communities[J].Functional Ecology,2010,24(1):18-27.
21 ROSSATTO D R,TAKAHASHI F S C,SILVA L D C R,et al.Leaf functional traits in sun and shade leaves of gallery forest trees in Distrito Federal,Brazil[J].Acta Botanica Brasilica,2010,24(3):640-647.
22朱济友,于强,刘亚培,等.植物功能性状及其叶经济谱对城市热环境的响应[J].北京林业大学学报,2018,40(9):1-10.ZHU Jiyou,YU Qiang,LIU Yapei,et al.Response of plant functional traits and leaf economics spectrum to urban thermal environment[J].Journal of Beijing Forestry University,2018,40(9):1-10.(in Chinese)
23朱济友,徐程扬,吴鞠.基于e Cognition植物叶片气孔密度及气孔面积快速测算方法[J].北京林业大学学报,2018,40(5):37-45.ZHU Jiyou,XU Chengyang,WU Ju.Fast estimation of stomatal density and stomatal area of plant leaves based on e Cognition[J].Journal of Beijing Forestry University,2018,40(5):37-45.(in Chinese)
24张大龙,张中典,李建明.环境因子对温室甜瓜蒸腾的驱动和调控效应研究[J/OL].农业机械学报,2015,46(11):137-144.http:∥www.j-csam.org/jcsam/ch/reader/abstract_view.aspx?file_no=20151119&flag=1&journal_id=jcsam.DOI:10.6041/j.issn.1000-1298.2015.11.019.ZHANG Dalong,ZHANG Zhongdian,LI Jianming.Co-ordination of environmental factors in driving and regulating transpiration rate of greenhouse grown muskmelon[J/OL].Transactions of the Chinese Society for Agricultural Machinery,2015,46(11):137-144.(in Chinese)
25 DIAZPEREZ J C,PHATAK S C,GIDDINGS D,et al.Root zone temperature,plant growth,and fruit yield of tomatillo as affected by plastic film mulch[J].Hortscience,2005,40(5):1312-1319.
26 DODD I C,HE J,TURNBULL C G,et al.The influence of supra-optimal root-zone temperatures on growth and stomatal conductance in Capsicum annuum L[J].Journal of Experimental Botany,2000,51(343):239-248.
27 KRAFT N J B,ACKERLY D D.Functional trait and phylogenetic tests of community assembly across spatial scales in an Amazonian forest[J].Ecological Monographs,2010,80(3):401-422.
28 YANG J,CI X,LU M,et al.Functional traits of tree species with phylogenetic signal co-vary with environmental niches in two large forest dynamics plots[J].Journal of Plant Ecology,2014,7(2SI):115-125.
29 PEASE A A,WINEMILLER K O.Functional diversity and trait-environment relationships of stream fish assemblages in a large tropical catchment[J].Freshwater Biology,2012,57(5):1060-1075.
30 WALKER A P,BECKERMAN A P,GU L,et al.The relationship of leaf photosynthetic traits Vcmaxand Jmaxto leaf nitrogen,leaf phosphorus,and specific leaf area:a meta analysis and modeling study[J].Ecology&Evolution,2014,4(16):3218-3235.
31 KIMM H,RYU Y.Seasonal variations in photosynthetic parameters and leaf area index in an urban park[J].Urban Forestry&Urban Greening,2015,14(4):1059-1067.
32 WESTOBY M,FALSTER D S,MOLES A T,et al.Plant ecological strategies:some leading dimensions of variation between species[J].Annual Review of Ecology&Systematics,2002,33(1):125-159.
33 LAVOREL S,GARNIER E.Predicting changes in community composition and ecosystem functioning from plant traits:revisiting the Holy Grail[J].Functional Ecology,2010,16(5):545-556.
34 ORDOEZ J C,BODEGOM P M V,WITTE J P M,et al.A global study of relationships between leaf traits,climate and soil measures of nutrient fertility[J].Global Ecology&Biogeography,2010,18(2):137-149.
35 GENG Y,WANG Z,LIANG C,et al.Effect of geographical range size on plant functional traits and the relationships between plant,soil and climate in Chinese grasslands[J].Global Ecology&Biogeography,2012,21(4):416-427.
36 LIENIN P,KLEYER M.Plant leaf economics and reproductive investment are responsive to gradients of land use intensity[J].Agriculture Ecosystems&Environment,2011,145(1):67-76.
37 REICH P B.The world-wide‘fast-slow’plant economics spectrum:a traits manifesto[J].Journal of Ecology,2014,102(2):275-301.
38 FRESCHET G T,CORNELISSEN J H C,VAN LOGTESTIJN R S P,et al.Evidence of the‘plant economics spectrum’in a subarctic flora[J].Journal of Ecology,2010,98(2):362-373.
39 WILSON K B,BALDOCCHI D D,HANSON P J.Spatial and seasonal variability of photosynthetic parameters and their relationship to leaf nitrogen in a deciduous forest[J].Tree Physiology,2000,20(9):565-578.
40 ADLER P B,SALGUEROGMEZ R,COMPAGNONI A,et al.Functional traits explain variation in plant life history strategies[J].Proceedings of the National Academy of Science,2014,111(2):740-745.
2 DIAZ S,CABIDO M.Vive la difference:plant functional diversity matters to ecosystem processes[J].Irish Historical Studies,2001,16(11):646-655.
3 VIOLLE C,NAVAS M L,VILE D,et al.Let the concept of trait be functional[J].Oikos,2007,116(5):882-892.
4 WRIGHT I J,REICH P B,WESTOBY M,et al.The worldwide leaf economics spectrum[J].Nature,2004,428(6985):821.
5 WRIGHT I J,WESTOBY M,REICH P B.Convergence towards higher leaf mass per area in dry and nutrient-poor habitats has different consequences for leaf life span[J].Journal of Ecology,2002,90(3):534-543.
6 DAVID D,ACKERLY D D.Community assembly,niche conservatism,and adaptive evolution in changing environments[J].International Journal of Plant Sciences,2010,164(Supp.3):165-184.
7 SHIPLEY B,LECHOWICZ M J,WRIGHT I,et al.Fundamental trade-offs generating the worldwide leaf economics spectrum[J].Ecology,2006,87(3):535-541.
8 PRIYADARSINI R,HIEN W N,DAVID C K W.Microclimatic modeling of the urban thermal environment of Singapore to mitigate urban heat island[J].Solar Energy,2008,82(8):727-745.
9 DHAKAL S,HANAKI K.Improvement of urban thermal environment by managing heat discharge sources and surface modification in Tokyo[J].Energy&Buildings,2002,34(1):13-23.
10 GORCHAKOV G I,KOPROV B M,SHUKUROV K A.Wind effect on aerosol transport from the underlying surface[J].Izvestiya Atmospheric&Oceanic Physics,2004,40(6):679-694.
11 STONE B,NORMAN J M.Land use planning and surface heat island formation:a parcel-based radiation flux approach[J].Atmospheric Environment,2006,40(19):3561-3573.
12 TAHA H.Meso-urban meteorological and photochemical modeling of heat island mitigation[J].Atmospheric Environment,2008,42(38):8795-8809.
13 KONIJNENDIJK C C,RANDRUP T B.Urban forestry&urban greening[J].Urban Forestry&Urban Greening,2002,1(1):1-4.
14 CASE J L,CROSSON W L,KUMAR S V,et al.Impacts of high resolution land surface initialization on regional sensible weather forecasts from the WRF model[J].Journal of Hydrometeorology,2008,9(6):1249-1266.
15 DASH P,GTTSCHE F M,OLESEN F S,et al.Separating surface emissivity and temperature using two-channel spectral indices and emissivity composites and comparison with a vegetation fraction method[J].Remote Sensing of Environment,2005,96(1):1-17.
16 HURTALOVT,MATEJKA F.Surface characteristics and energy fluxes above different plant canopies[J].Agricultural&Forest Meteorology,1999,99(22):491-499.
17 SALAMANCA F,MARTILLI A.A new building energy model coupled with an urban canopy parameterization for urban climate simulations-part II.validation with one dimension off-line simulations[J].Theoretical and Applied Climatology,2010,99(3/4):345-356.
18 JOHNSON J D,O'CONNOR K A,HANSEN M K,et al.Effects of prior stress on LPS-induced cytokine and sickness responses[J].Am.J.Physiol.Regul.Integr.Comp.Physiol.,2003,284(2):22-32.
19 KATTGE J,DAZ S,LAVOREL S,et al.TRY-a global database of plant traits[J].Global Change Biology,2011,17(9):2905-2935.
20 LOEUILLE N.Consequences of adaptive foraging in diverse communities[J].Functional Ecology,2010,24(1):18-27.
21 ROSSATTO D R,TAKAHASHI F S C,SILVA L D C R,et al.Leaf functional traits in sun and shade leaves of gallery forest trees in Distrito Federal,Brazil[J].Acta Botanica Brasilica,2010,24(3):640-647.
22朱济友,于强,刘亚培,等.植物功能性状及其叶经济谱对城市热环境的响应[J].北京林业大学学报,2018,40(9):1-10.ZHU Jiyou,YU Qiang,LIU Yapei,et al.Response of plant functional traits and leaf economics spectrum to urban thermal environment[J].Journal of Beijing Forestry University,2018,40(9):1-10.(in Chinese)
23朱济友,徐程扬,吴鞠.基于e Cognition植物叶片气孔密度及气孔面积快速测算方法[J].北京林业大学学报,2018,40(5):37-45.ZHU Jiyou,XU Chengyang,WU Ju.Fast estimation of stomatal density and stomatal area of plant leaves based on e Cognition[J].Journal of Beijing Forestry University,2018,40(5):37-45.(in Chinese)
24张大龙,张中典,李建明.环境因子对温室甜瓜蒸腾的驱动和调控效应研究[J/OL].农业机械学报,2015,46(11):137-144.http:∥www.j-csam.org/jcsam/ch/reader/abstract_view.aspx?file_no=20151119&flag=1&journal_id=jcsam.DOI:10.6041/j.issn.1000-1298.2015.11.019.ZHANG Dalong,ZHANG Zhongdian,LI Jianming.Co-ordination of environmental factors in driving and regulating transpiration rate of greenhouse grown muskmelon[J/OL].Transactions of the Chinese Society for Agricultural Machinery,2015,46(11):137-144.(in Chinese)
25 DIAZPEREZ J C,PHATAK S C,GIDDINGS D,et al.Root zone temperature,plant growth,and fruit yield of tomatillo as affected by plastic film mulch[J].Hortscience,2005,40(5):1312-1319.
26 DODD I C,HE J,TURNBULL C G,et al.The influence of supra-optimal root-zone temperatures on growth and stomatal conductance in Capsicum annuum L[J].Journal of Experimental Botany,2000,51(343):239-248.
27 KRAFT N J B,ACKERLY D D.Functional trait and phylogenetic tests of community assembly across spatial scales in an Amazonian forest[J].Ecological Monographs,2010,80(3):401-422.
28 YANG J,CI X,LU M,et al.Functional traits of tree species with phylogenetic signal co-vary with environmental niches in two large forest dynamics plots[J].Journal of Plant Ecology,2014,7(2SI):115-125.
29 PEASE A A,WINEMILLER K O.Functional diversity and trait-environment relationships of stream fish assemblages in a large tropical catchment[J].Freshwater Biology,2012,57(5):1060-1075.
30 WALKER A P,BECKERMAN A P,GU L,et al.The relationship of leaf photosynthetic traits Vcmaxand Jmaxto leaf nitrogen,leaf phosphorus,and specific leaf area:a meta analysis and modeling study[J].Ecology&Evolution,2014,4(16):3218-3235.
31 KIMM H,RYU Y.Seasonal variations in photosynthetic parameters and leaf area index in an urban park[J].Urban Forestry&Urban Greening,2015,14(4):1059-1067.
32 WESTOBY M,FALSTER D S,MOLES A T,et al.Plant ecological strategies:some leading dimensions of variation between species[J].Annual Review of Ecology&Systematics,2002,33(1):125-159.
33 LAVOREL S,GARNIER E.Predicting changes in community composition and ecosystem functioning from plant traits:revisiting the Holy Grail[J].Functional Ecology,2010,16(5):545-556.
34 ORDOEZ J C,BODEGOM P M V,WITTE J P M,et al.A global study of relationships between leaf traits,climate and soil measures of nutrient fertility[J].Global Ecology&Biogeography,2010,18(2):137-149.
35 GENG Y,WANG Z,LIANG C,et al.Effect of geographical range size on plant functional traits and the relationships between plant,soil and climate in Chinese grasslands[J].Global Ecology&Biogeography,2012,21(4):416-427.
36 LIENIN P,KLEYER M.Plant leaf economics and reproductive investment are responsive to gradients of land use intensity[J].Agriculture Ecosystems&Environment,2011,145(1):67-76.
37 REICH P B.The world-wide‘fast-slow’plant economics spectrum:a traits manifesto[J].Journal of Ecology,2014,102(2):275-301.
38 FRESCHET G T,CORNELISSEN J H C,VAN LOGTESTIJN R S P,et al.Evidence of the‘plant economics spectrum’in a subarctic flora[J].Journal of Ecology,2010,98(2):362-373.
39 WILSON K B,BALDOCCHI D D,HANSON P J.Spatial and seasonal variability of photosynthetic parameters and their relationship to leaf nitrogen in a deciduous forest[J].Tree Physiology,2000,20(9):565-578.
40 ADLER P B,SALGUEROGMEZ R,COMPAGNONI A,et al.Functional traits explain variation in plant life history strategies[J].Proceedings of the National Academy of Science,2014,111(2):740-745.