阔叶红松林6种阔叶树单个叶片叶面积和叶干质量的垂直变异及模型构建
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摘要
快捷、准确地测定植物叶片尺度上的叶面积(LA)和叶干质量(LDM)对于探讨植物性状对气候变化的响应机制至关重要,但适于测定区域尺度上不同阔叶植物单个叶片LA和LDM的经验模型尚未提出.本研究以中国东北4个分布区阔叶红松林内的白桦、紫椴、山杨、枫桦、水曲柳和裂叶榆6种阔叶树种为研究对象,分别测定其不同冠层高度(上层、中层和下层)叶片的叶长、叶宽、叶厚、LA及LDM.以叶长与叶宽之比(叶长宽比)的中位数为标准将6种阔叶树分为两组,检验每组树种不同冠层高度对构建预测LA和LDM的经验模型是否存在显著影响;构建适于预测区域尺度上不同冠层单个叶片LA和LDM的经验模型,验证其预测精度;并进一步评估构建的经验模型预测其他区域相同阔叶树种LA和LDM的适用性.结果表明:整体上6种阔叶树单个叶片的LA随冠层高度的降低呈显著增大趋势,而部分树种的LDM呈下降趋势;冠层高度对构建预测LA和LDM的经验模型具有显著影响;构建的经验模型预测两组阔叶树种不同冠层单个叶片LA和LDM的平均精度分别为95%和83%,且基于这些模型预测其他区域相应树种LA和LDM的平均精度分别为94%和80%,表明本研究构建的经验模型在我国东北区域具有普遍适用性.
Rapidly and accurately predicting leaf area(LA) and leaf dry mass(LDM) are essential for exploring the response of plant traits to climate change. Empirical models suitable for predicting LA and LDM of a single leaf for various broadleaved tree species at the regional scale have not been proposed. We selected six broadleaved tree species in four mixed broadleaved-Korean pine(Pinus koraiensis) forests in northeastern China, including Betula platyphylla, Tilia amurensis, Populus davidiana, Betula costata, Fraxinus mandshurica and Ulmus laciniata, and measured leaf length, leaf width, leaf thickness, LA and LDM at different canopy layers(top, middle, and low). Using the median of leaf length and width ratio as the classification criterion, the six species were sorted into two groups. We tested whether different canopy layers for each group of broadleaved tree species had significant impacts on the empirical model for predicting LA and LDM. We constructed empirical models suitable for predicting LA and LDM of a single leaf at different canopy layers at the regional scale, and verified their forecast accuracy, and further evaluated their applicability for predicting LA and LDM of same broadleaved tree species in other regions. These results showed that the LA of a single leaf increased significantly with the decreases of canopy height for the six tree species, while the LDM of some broadleaved tree species showed a downward trend. The canopy height had significant impacts on constructing the empirical model for LA and LDM. The average forecast accuracy of empirical model was 95% and 83% for LA and LDM of a single leaf across canopy layers for two groups of broadleaved tree species, respectively. The average forecast accuracy was 94% and 80% for predicting LA and LDM of corresponding broadleaved tree species in other regions, respectively, indicating that the empirical models constructed in this study had a universal applicability in Northeast China.
Rapidly and accurately predicting leaf area(LA) and leaf dry mass(LDM) are essential for exploring the response of plant traits to climate change. Empirical models suitable for predicting LA and LDM of a single leaf for various broadleaved tree species at the regional scale have not been proposed. We selected six broadleaved tree species in four mixed broadleaved-Korean pine(Pinus koraiensis) forests in northeastern China, including Betula platyphylla, Tilia amurensis, Populus davidiana, Betula costata, Fraxinus mandshurica and Ulmus laciniata, and measured leaf length, leaf width, leaf thickness, LA and LDM at different canopy layers(top, middle, and low). Using the median of leaf length and width ratio as the classification criterion, the six species were sorted into two groups. We tested whether different canopy layers for each group of broadleaved tree species had significant impacts on the empirical model for predicting LA and LDM. We constructed empirical models suitable for predicting LA and LDM of a single leaf at different canopy layers at the regional scale, and verified their forecast accuracy, and further evaluated their applicability for predicting LA and LDM of same broadleaved tree species in other regions. These results showed that the LA of a single leaf increased significantly with the decreases of canopy height for the six tree species, while the LDM of some broadleaved tree species showed a downward trend. The canopy height had significant impacts on constructing the empirical model for LA and LDM. The average forecast accuracy of empirical model was 95% and 83% for LA and LDM of a single leaf across canopy layers for two groups of broadleaved tree species, respectively. The average forecast accuracy was 94% and 80% for predicting LA and LDM of corresponding broadleaved tree species in other regions, respectively, indicating that the empirical models constructed in this study had a universal applicability in Northeast China.
引文
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[3] Alton PB.The sensitivity of models of gross primary productivity to meteorological and leaf area forcing:A comparison between a Penman-Monteith ecophysiological approach and the MODIS light-use efficiency algorithm.Agricultural and Forest Meteorology,2016,217:11-24
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[5] Leverenz JW,Hinckley TM.Shoot structure,leaf area index and productivity of evergreen conifer stands.Tree Physiology,1990,6:135-149
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[9] Nelson AS,Wagner RG,Day ME,et al.Light absorption and light-use efficiency of juvenile white spruce trees in natural stands and plantations.Forest Ecology and Management,2016,376:158-165
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[11] Wright IJ,Reich PB,Westoby M,et al.The worldwide leaf economics spectrum.Nature,2004,428:821-827
[12] Duursma RA,Falster DS.Leaf mass per area,not total leaf area,drives differences in above ground biomass distribution among woody plant functional types.New Phytologist,2016,212:368-376
[13] Okajima Y,Taneda H,Noguchi K,et al.Optimum leaf size predicted by a novel leaf energy balance model incorporating dependencies of photosynthesis on light and temperature.Ecological Research,2012,27:333-346
[14] Marron N,Villar M,Dreyer E,et al.Diversity of leaf traits related to productivity in 31 Populus deltoides×Popu-lus nigra clones.Tree Physiology,2005,25:425-435
[15] He Y-D (何应德).Image Analysis based Pixel Area Calculation of Tree Leaf.Master Thesis.Beijing:Beijing Forestry University,2011 (in Chinese)
[16] Liu ZL,Chen JM,Jin GZ,et al.Estimating seasonal variations of leaf area index using litterfall collection and optical methods in four mixed evergreen-deciduous forests.Agricultural and Forest Meteorology,2015,209:36-48
[17] Daughtry CST.Direct measurements of canopy structure.Remote Sensing Reviews,1990,5:45-60
[18] Peksen E.Non-destructive leaf area estimation model for faba bean (Vicia faba L.).Scientia Horticulturae,2007,113:322-328
[19] Cristofori V,Rouphael Y,Mendoza DE,et al.A simple model for estimating leaf area of hazelnut from linear measurements.Scientia Horticulturae,2007,113:221-225
[20] Keramatlou I,Sharifani M,Sabouri H,et al.A simple linear model for leaf area estimation in Persian walnut (Juglans regia L.).Scientia Horticulturae,2015,184:36-39
[21] Rouphael Y,Mouneimne AH,Ismail A,et al.Modeling individual leaf area of rose (Rosa hybrida L.) based on leaf length and width measurement.Photosynthetica,2010,48:9-15
[22] Rouphael Y,Colla G,Fanasca S,et al.Leaf area estimation of sunflower leaves from simple linear measurements.Photosynthetica,2007,45:306-308
[23] Liu ZL,Zhu Y,Li FR,et al.Non-destructively predicting leaf area,leaf mass and specific leaf area based on a linear mixed-effect model for broadleaf species.Ecological Indicators,2017,78:340-350
[24] Tondjo K,Brancheriau L,Sabatier SA,et al.Non-destructive measurement of leaf area and dry biomass in Tectona grandis.Trees,2015,29:1625-1631
[25] Wang Y-J (王彦君),Jin G-Z (金光泽),Liu Z-L (刘志理).Construction of empirical models for leaf area and leaf mass of two broadleaf species in Xiaoxing’an Mountains,China.Chinese Journal of Applied Ecology (应用生态学报),2018,29(6):1745-1752 (in Chinese)
[26] Takenaka A.Effects of leaf blade narrowness and petiole length on the light capture efficiency of a shoot.Ecological Research,1994,9:109-114
[27] Li D-Z (李德志),Zang R-G (臧润国).The research advances on the structure and function of forest canopy,as well as their temporal and spatial changes.World Forestry Research (世界林业研究),2004,17(3):12-16 (in Chinese)
[28] Vierling LA,Wessman CA.Photosynthetically active radiation heterogeneity within a monodominant Congolese rain forest canopy.Agricultural and Forest Meteorology,2000,103:265-278
[29] Liu XD,Zhu CQ,Lei JP,et al.The distribution of photosynthetic radiation in the canopy of poplar plantation.Scientia Silvae Sinicae,2000,36:2-7
[30] Jia J-P (贾俊平),He X-Q (何晓群),Jin Y-J (金勇进).Statistics.6th Ed.Beijing:China Renmin University Press,2015 (in Chinese)
[31] Akaike H.Information theory and an extension of the maximum likelihood principle.International Symposium on Information Theory,1973,1:267-281
[32] Marquardit DW.Generalized inverses,ridge regression,biased linear estimation,and nonlinear estimation.Technometrics,1970,12:591-612
[33] Liu Z-L (刘志理),Jin G-Z (金光泽).Estimation of seasonal changes in leaf area index based on optical methods in spruce-fir valley forest.Chinese Journal of Applied Ecology (应用生态学报),2014,25(12):3420-3428 (in Chinese)
[34] Chiang LH,Pell RJ,Seasholtz MB.Exploring process data with the use of robust outlier detection algorithms.Journal of Process Control,2003,13:437-449
[35] R Development Core Team.R:A Language and Environment for Statistical Computing.R Foundation for Statistical Computing [EB/OL].(2015-03-09) [2018-11-28].http://www.R-project.org
[36] Pinheiro J,Bates D,DebRoy S,et al.NLDMe:Linear and Nonlinear Mixed Effects Models.R Package Version 3.1-131 [EB/OL].(2017-02-06) [2018-11-28].https://CRAN.R-project.org/package=nlme
[37] Salazar JCS,Muňoz LDMM,Bautista EHD,et al.Non-destructive estimation of the leaf weight and leaf area in cacao (Theobroma cacao).Scienti Horticulturae,2018,229:19-24
[38] Li J-Q (李俊清).Forest Ecology.2nd Ed.Beijing:Higher Education Press,2010 (in Chinese)
[39] Parkhurst DF,Loucks OL.Optimal leaf size in relation to environment.Journal of Ecology,1972,60:505-537
[40] Poorter H,Pepin S,Rijkers T,et al.Construction costs,chemical composition and payback time of high-and low-irradiance leaves.Journal of Experimental Botany,2006,57:355-371
[41] Rijkers T,Pons TL,Bongers F.The effect of tree height and light availability on photosynthetic leaf traits of four neotropical species differing in shade tolerance.Functional Ecology,2000,14:77-86
[42] Boardman NK.Comparative photosynthesis of sun and shade plants.Annual Review of Plant Physiology,1977,28:355-377
[43] Tai NH,Hung TT,Ahn TI,et al.Estimation of leaf area,fresh weight,and dry weight of paprika (Capsicum annuum) using leaf length and width in Rockwool-Based soilless culture.Horticulture,Environment,and Biotechnology,2009,50:422-426
[44] Meng FQ,Zhang GF,Li XC,et al.Growth synchrony between leaves and stems during twig development differs among plant functional types of subtropical rainforest woody species.Tree Physiology,2015,35:621-631
[45] Bell A.Plant Form:An Illustrated Guide to Flowering Plant Morphology.London:Oxford University Press,1991
[46] Brito-Rocha E,Schilling AC,Dos AL,et al.Regression models for estimating leaf area of seedlings and adult individuals of neotropical rainforest tree species.Brazilian Journal of Biology,2016,76:983-989
[2] Peppe DJ,Royer DL,Cariglino B,et al.Sensitivity of leaf size and shape to climate:Global patterns and paleoclimatic applications.New Phytologist,2011,190:724-739
[3] Alton PB.The sensitivity of models of gross primary productivity to meteorological and leaf area forcing:A comparison between a Penman-Monteith ecophysiological approach and the MODIS light-use efficiency algorithm.Agricultural and Forest Meteorology,2016,217:11-24
[4] Zhang L (张林),Luo T-X (罗天祥).Advances in ecological studies on leaf lifespan and associated leaf traits.Chinese Journal of Plant Ecology (植物生态学报),2004,28(6):844-852 (in Chinese)
[5] Leverenz JW,Hinckley TM.Shoot structure,leaf area index and productivity of evergreen conifer stands.Tree Physiology,1990,6:135-149
[6] Zhao M,Xiang W,Peng C,et al.Simulating age-related changes in carbon storage and allocation in a Chinese fir plantation growing in southern China using the 3-PG model.Forest Ecology and Management,2009,257:1520-1531
[7] Lucht W,Prentice IC,Myneni RB,et al.Climatic control of the high-latitude vegetation greening trend and Pinatubo effect.Science,2002,296:1687-1689
[8] Ewert F.Modelling plant responses to elevated CO2:How important is leaf area index?Annals of Botany,2004,93:619-627
[9] Nelson AS,Wagner RG,Day ME,et al.Light absorption and light-use efficiency of juvenile white spruce trees in natural stands and plantations.Forest Ecology and Management,2016,376:158-165
[10] Cornelissen JHC,Lavorel S,Garnier E,et al.Handbook of protocols for standardised and easy measurement of plant functional traits worldwide.Australian Journal of Botany,2003,51:335-380
[11] Wright IJ,Reich PB,Westoby M,et al.The worldwide leaf economics spectrum.Nature,2004,428:821-827
[12] Duursma RA,Falster DS.Leaf mass per area,not total leaf area,drives differences in above ground biomass distribution among woody plant functional types.New Phytologist,2016,212:368-376
[13] Okajima Y,Taneda H,Noguchi K,et al.Optimum leaf size predicted by a novel leaf energy balance model incorporating dependencies of photosynthesis on light and temperature.Ecological Research,2012,27:333-346
[14] Marron N,Villar M,Dreyer E,et al.Diversity of leaf traits related to productivity in 31 Populus deltoides×Popu-lus nigra clones.Tree Physiology,2005,25:425-435
[15] He Y-D (何应德).Image Analysis based Pixel Area Calculation of Tree Leaf.Master Thesis.Beijing:Beijing Forestry University,2011 (in Chinese)
[16] Liu ZL,Chen JM,Jin GZ,et al.Estimating seasonal variations of leaf area index using litterfall collection and optical methods in four mixed evergreen-deciduous forests.Agricultural and Forest Meteorology,2015,209:36-48
[17] Daughtry CST.Direct measurements of canopy structure.Remote Sensing Reviews,1990,5:45-60
[18] Peksen E.Non-destructive leaf area estimation model for faba bean (Vicia faba L.).Scientia Horticulturae,2007,113:322-328
[19] Cristofori V,Rouphael Y,Mendoza DE,et al.A simple model for estimating leaf area of hazelnut from linear measurements.Scientia Horticulturae,2007,113:221-225
[20] Keramatlou I,Sharifani M,Sabouri H,et al.A simple linear model for leaf area estimation in Persian walnut (Juglans regia L.).Scientia Horticulturae,2015,184:36-39
[21] Rouphael Y,Mouneimne AH,Ismail A,et al.Modeling individual leaf area of rose (Rosa hybrida L.) based on leaf length and width measurement.Photosynthetica,2010,48:9-15
[22] Rouphael Y,Colla G,Fanasca S,et al.Leaf area estimation of sunflower leaves from simple linear measurements.Photosynthetica,2007,45:306-308
[23] Liu ZL,Zhu Y,Li FR,et al.Non-destructively predicting leaf area,leaf mass and specific leaf area based on a linear mixed-effect model for broadleaf species.Ecological Indicators,2017,78:340-350
[24] Tondjo K,Brancheriau L,Sabatier SA,et al.Non-destructive measurement of leaf area and dry biomass in Tectona grandis.Trees,2015,29:1625-1631
[25] Wang Y-J (王彦君),Jin G-Z (金光泽),Liu Z-L (刘志理).Construction of empirical models for leaf area and leaf mass of two broadleaf species in Xiaoxing’an Mountains,China.Chinese Journal of Applied Ecology (应用生态学报),2018,29(6):1745-1752 (in Chinese)
[26] Takenaka A.Effects of leaf blade narrowness and petiole length on the light capture efficiency of a shoot.Ecological Research,1994,9:109-114
[27] Li D-Z (李德志),Zang R-G (臧润国).The research advances on the structure and function of forest canopy,as well as their temporal and spatial changes.World Forestry Research (世界林业研究),2004,17(3):12-16 (in Chinese)
[28] Vierling LA,Wessman CA.Photosynthetically active radiation heterogeneity within a monodominant Congolese rain forest canopy.Agricultural and Forest Meteorology,2000,103:265-278
[29] Liu XD,Zhu CQ,Lei JP,et al.The distribution of photosynthetic radiation in the canopy of poplar plantation.Scientia Silvae Sinicae,2000,36:2-7
[30] Jia J-P (贾俊平),He X-Q (何晓群),Jin Y-J (金勇进).Statistics.6th Ed.Beijing:China Renmin University Press,2015 (in Chinese)
[31] Akaike H.Information theory and an extension of the maximum likelihood principle.International Symposium on Information Theory,1973,1:267-281
[32] Marquardit DW.Generalized inverses,ridge regression,biased linear estimation,and nonlinear estimation.Technometrics,1970,12:591-612
[33] Liu Z-L (刘志理),Jin G-Z (金光泽).Estimation of seasonal changes in leaf area index based on optical methods in spruce-fir valley forest.Chinese Journal of Applied Ecology (应用生态学报),2014,25(12):3420-3428 (in Chinese)
[34] Chiang LH,Pell RJ,Seasholtz MB.Exploring process data with the use of robust outlier detection algorithms.Journal of Process Control,2003,13:437-449
[35] R Development Core Team.R:A Language and Environment for Statistical Computing.R Foundation for Statistical Computing [EB/OL].(2015-03-09) [2018-11-28].http://www.R-project.org
[36] Pinheiro J,Bates D,DebRoy S,et al.NLDMe:Linear and Nonlinear Mixed Effects Models.R Package Version 3.1-131 [EB/OL].(2017-02-06) [2018-11-28].https://CRAN.R-project.org/package=nlme
[37] Salazar JCS,Muňoz LDMM,Bautista EHD,et al.Non-destructive estimation of the leaf weight and leaf area in cacao (Theobroma cacao).Scienti Horticulturae,2018,229:19-24
[38] Li J-Q (李俊清).Forest Ecology.2nd Ed.Beijing:Higher Education Press,2010 (in Chinese)
[39] Parkhurst DF,Loucks OL.Optimal leaf size in relation to environment.Journal of Ecology,1972,60:505-537
[40] Poorter H,Pepin S,Rijkers T,et al.Construction costs,chemical composition and payback time of high-and low-irradiance leaves.Journal of Experimental Botany,2006,57:355-371
[41] Rijkers T,Pons TL,Bongers F.The effect of tree height and light availability on photosynthetic leaf traits of four neotropical species differing in shade tolerance.Functional Ecology,2000,14:77-86
[42] Boardman NK.Comparative photosynthesis of sun and shade plants.Annual Review of Plant Physiology,1977,28:355-377
[43] Tai NH,Hung TT,Ahn TI,et al.Estimation of leaf area,fresh weight,and dry weight of paprika (Capsicum annuum) using leaf length and width in Rockwool-Based soilless culture.Horticulture,Environment,and Biotechnology,2009,50:422-426
[44] Meng FQ,Zhang GF,Li XC,et al.Growth synchrony between leaves and stems during twig development differs among plant functional types of subtropical rainforest woody species.Tree Physiology,2015,35:621-631
[45] Bell A.Plant Form:An Illustrated Guide to Flowering Plant Morphology.London:Oxford University Press,1991
[46] Brito-Rocha E,Schilling AC,Dos AL,et al.Regression models for estimating leaf area of seedlings and adult individuals of neotropical rainforest tree species.Brazilian Journal of Biology,2016,76:983-989