热带季雨林不同小生境大戟科植物幼树的叶片结构、耐旱性和光合能力之间的相关性

doi:10.3724/SP.J.1258.2014.00028

摘要/Abstract

摘要：

植物的叶片结构和功能性状受到自身、环境和系统发育的影响。该研究选取西双版纳20 hm²热带雨林动态监测大样地内18种分布格局不同的大戟科植物, 测量了幼树叶片的解剖结构、水分关系特征、最大光合能力和暗呼吸, 主要探讨了叶片结构对植物耐旱性和光合能力的影响, 耐旱性和光合能力之间的权衡关系, 以及环境水分条件对植物功能性状相关性的影响。结果表明: 1)生境内植物表现出一定的结构和功能的趋同性, 分布在山脊和山坡的种比沟谷种具有更强的耐失水能力; 2)去除了系统发育的影响后, 一些关键性状(特别是叶片密度和膨压丧失点时的水势、饱和渗透势等)之间存在跨生境尺度上的相关关系, 植物叶片结构同时影响了植物的耐失水能力和光合能力, 植物叶片自身的结构限制导致了植物的耐旱性(高的叶片密度、比叶质量)和光合能力(低的叶片密度、比叶质量)存在反向进化关系; 3)如果研究的植物类群亲缘关系较近, 传统的Pearson相关分析不能很好地揭示其性状间的相关关系, 因而必须采用系统发育独立对照差作相关分析。大戟科植物的结构和功能在水分梯度和光梯度上的生态位分化也从功能性状的角度为热带季雨林能维持高生物多样性, 保持植物物种长期共存提供了一个可能的解释。

关键词: 暗呼吸, 生境, 叶片解剖结构, 光合作用, 压力-容积曲线

Abstract:

Aims Leaf structural and functional traits have been extensively studied to explain community assembly mechanisms, species distributions, niche differentiations, and even ecosystem services functions. However, these traits are influenced by both environment and phylogeny, showing correlations or trade-offs among them. In this study, we assessed the impacts of leaf structure on drought tolerance and photosynthetic potential, and the trade-off between drought tolerance and photosynthetic capacity, to provide an explanation for species coexistence and the maintenance of high biodiversity in tropical rainforests.
Methods We chose 18 species in the Euphobiaceae family differing in distribution patterns along topographic gradients in a 20 hm² forest dynamics monitoring plot (FDP) in Xishuangbanna. We measured leaf anatomy, leaf water relations characteristics, maximum photosynthetic rate, and dark respiration, and used two different methods—the traditional Pearson correlation and phylogenetic independent contrasts—to analyze the relationships among those traits.
Important findings We found that: 1) species showed convergence in structures and functions within specific habitat; species on ridge or slope had a stronger water loss-tolerance abilities than species in the valley. 2) Correlations among some key traits (specifically, leaf density, water potential at turgor loss point, and water potential at full turgor, etc.) were found among habitats; plants adjusted leaf structure to influence simultaneously plant water loss-tolerance abilities and photosynthetic capability, which may result in a trade-off between drought tolerance (high leaf density, leaf mass per area) and photosynthetic capability (low leaf density, leaf mass per area). 3) The phylogenetic independent contrasts must be used when analyzing correlations among the traits of genetically related species due to the weakness of traditional Pearson analysis. The ecological niche differentiation to water and light gradients as revealed by the present study provides a potential explanation for the high diversity of the seasonal tropical rainforest.

Key words: dark respiration, habitat, leaf structure, photosynthesis, pressure-volume curve

孙善文, 章永江, 曹坤芳. 热带季雨林不同小生境大戟科植物幼树的叶片结构、耐旱性和光合能力之间的相关性. 植物生态学报, 2014, 38(4): 311-324. DOI: 10.3724/SP.J.1258.2014.00028

SUN Shan-Wen, ZHANG Yong-Jiang, CAO Kun-Fang. Correlations among leaf structure, drought tolerance and photosynthetic capacity in saplings of Euphorbiaceae from different micro-habitats in a seasonal tropical rainforest. Chinese Journal of Plant Ecology, 2014, 38(4): 311-324. DOI: 10.3724/SP.J.1258.2014.00028

导出引用管理器 EndNote|Ris|BibTeX

链接本文: https://www.plant-ecology.com/CN/10.3724/SP.J.1258.2014.00028

https://www.plant-ecology.com/CN/Y2014/V38/I4/311

图/表 10

参考文献 66

[1]	Aranda I, Castro L, Pardos M, Gil L, Pardos JA (2005). Effects of the interaction between drought and shade on water relations, gas exchange and morphological traits in cork oak (Quercus suber L.) seedlings. Forest Ecology and Management, 210, 117-129.
[2]	Bacelar EA, Correia CM, Moutinho-Pereira JM, Goncalves BC, Lopes JI, Torres-Pereira JM (2004). Sclerophylly and leaf anatomical traits of five field-grown olive cultivars growing under drought conditions. Tree Physiology, 24, 233-239. URL PMID
[3]	Bartlett MK, Scoffoni C, Ardy R, Zhang Y, Sun SW, Cao KF, Sack L (2012a). Rapid determination of comparative drought tolerance traits: using an osmometer to predict turgor loss point. Methods in Ecology and Evolution, 3, 880-888.
[4]	Bartlett MK, Scoffoni C, Sack L (2012b). The determinants of leaf turgor loss point and prediction of drought tolerance of species and biomes: a global meta-analysis. Ecology Letters, 15, 393-405. URL PMID
[5]	Becker P, Wong M (1994). Drought induced mortality in tropical heath forest. Journal of Tropical Sciences, 5, 416-417.
[6]	Brodersen CR, Vogelmann TC (2007). Do epidermal lens cells facilitate the absorptance of diffuse light? American Journal of Botang, 94, 1061-1066.
[7]	Brown C, Burslem DFRP, Illian JB, Bao L, Brockelman W, Cao M, Chang LW, Dattaraja HS, Davies S, Gunatilleke CVS, Gunatilleke IAUN, Huang J, Kassim AR, LaFrankie JV, Lian J, Lin L, Ma K, Mi X, Nathalang A, Noor S, Ong P, Sukumar R, Su SH, Sun IF, Suresh HS, Tan S, Thompson J, Uriarte M, Valencia R, Yap SL, Ye W, Law R (2013). Multispecies coexistence of trees in tropical forests: spatial signals of topographic niche differentiation increase with environmental heterogeneity. Proceedings of the Royal Society B: Biological Sciences, 280, 20130502. DOI URL PMID
[8]	Bucci S, Goldstein G, Meinzer F, Scholz F, Franco A, Bust- amante M (2004). Functional convergence in hydraulic architecture and water relations of tropical savanna trees: from leaf to whole plant. Tree Physiology, 24, 891-899. URL PMID
[9]	Cao KF (2000). Leaf anatomy and chlorophyll content of 12 woody species in contrasting light conditions in a bornean heath forest. Canadian Journal of Botany, 78, 1245-1253. DOI URL
[10]	Cao M, Zou XM, Warren M, Zhu H (2006). Tropical forests of Xishuangbanna, China. Biotropica, 38, 306-309. DOI URL
[11]	Chartzoulakis K, Patakas A, Kofidis G, Bosabalidis A, Nastou A (2002). Water stress affects leaf anatomy, gas exchange, water relations and growth of two avocado cultivars. Scientia Horticulturae, 95, 39-50. DOI URL
[12]	Choat B, Medek DE, Stuart SA, Pasquet-Kok J, Egerton JJG, Salari H, Sack L, Ball MC (2011). Xylem traits mediate a trade-off between resistance to freeze-thaw-induced embolism and photosynthetic capacity in overwintering evergreens. New Phytologist, 191, 996-1005.
[13]	Cutler J, Rains D, Loomis R (1977). The importance of cell size in the water relations of plants. Physiologia Plantarum, 40, 255-260.
[14]	DeLucia EH, Nelson K, Vogelmann TC, Smith WK (1996). Contribution of intercellular reflectance to photosynthesis in shade leaves. Plant, Cell & Environment, 19, 159-170.
[15]	Evans J, Vogelmann T, Williams W, Gorton H (2004). Chloroplast to leaf. In: Smith W, Vogelmann T, Critchley C eds. Photosynthetic Adaptation. Springer, New York. 178. 15-41.
[16]	Felsenstein J (1985). Phylogenies and the comparative method. The American Naturalist, 125, 1-15.
[17]	Feng YL, Lei YB, Wang RF, Callaway RM, Valiente-Banuet A, Inderjit , Li YP, Zheng YL (2009). Evolutionary tradeoffs for nitrogen allocation to photosynthesis versus cell walls in an invasive plant. Proceedings of the National Academy of Sciences of the United States of America, 106, 1853-1856. URL PMID
[18]	Fu PL, Jiang YJ, Wang AY, Brodribb TJ, Zhang JL, Zhu SD, Cao KF (2012). Stem hydraulic traits and leaf water-stress tolerance are co-ordinated with the leaf phenology of angiosperm trees in an asian tropical dry karst forest. Annals of Botany, 110, 189-199. DOI URL PMID
[19]	Garland T, Harvey PH, Ives AR (1992). Procedures for the analysis of comparative data using phylogenetically independent contrasts. Systematic Biology, 41, 18-32.
[20]	Green DS, Kruger EL (2001). Light-mediated constraints on leaf function correlate with leaf structure among deciduous and evergreen tree species. Tree Physiology, 21, 1341-1346. DOI URL PMID
[21]	Hanba YT, Miyazawa SI, Terashima I (1999). The influence of leaf thickness on the CO₂ transfer conductance and leaf stable carbon isotope ratio for some evergreen tree species in Japanese warm-temperate forests. Functional Ecology, 13, 632-639.
[22]	Hassiotou F, Renton M, Ludwig M, Evans JR, Veneklaas EJ (2010). Photosynthesis at an extreme end of the leaf trait spectrum: How does it relate to high leaf dry mass per area and associated structural parameters? Journal of Experimental Botany, 61, 3015-3028. DOI URL PMID
[23]	Hessini K, Martínez JP, Gandour M, Albouchi A, Soltani A, Abdelly C (2009). Effect of water stress on growth, osmo- tic adjustment, cell wall elasticity and water-use efficiency in Spartina alterniflora. Environmental and Experimental Botany, 67, 312-319.
[24]	Hu YH, Sheng DY, Xiang YZ, Yang ZJ, Xu DP, Zhang NN, Shi LL (2013). The environment, not space, dominantly structures the landscape patterns of the richness and composition of the tropical understory vegetation. PLoS ONE, 8, e81308. URL PMID
[25]	Karabourniotis G (1998). Light-guiding function of foliar sclereids in the evergreen sclerophyll phillyrea latifolia: a quantitative approach. Journal of Experimental Botany, 49, 739-746.
[26]	Karabourniotis G, Bornman JF (1999). Penetration of UV-A, UV-B and blue light through the leaf trichome layers of two xeromorphic plants, olive and oak, measured by optical fibre microprobes. Physiologia Plantarum, 105, 655-661.
[27]	Karabourniotis G, Papastergiou N, Kabanopoulou E, Fasseas C (1994). Foliar sclereids of Olea europaea may function as optical fibres. Canadian Journal of Botany, 72, 330-336.
[28]	Kubiske ME, Abrams MD (1990). Pressure-volume relation- ships in non-rehydrated tissue at various water deficits. Plant, Cell & Environment, 13, 995-1000.
[29]	Kursar TA, Engelbrecht BMJ, Burke A, Tyree MT, Omari BE, Giraldo JP (2009). Tolerance to low leaf water status of tropical tree seedlings is related to drought performance and distribution. Functional Ecology, 23, 93-102.
[30]	Lan GY, Getzin S, Wiegand T, Hu YH, Xie GS, Zhu H, Cao M (2012). Spatial distribution and interspecific associations of tree species in a tropical seasonal rain forest of China. PLoS ONE, 7, e46074. DOI URL PMID
[31]	Lan YG, Hu YH, Cao M, Zhu H, Wang H, Zhou SS, Deng SS, Deng XB, Cui JY, Huang JG, Liu LY, Xu HL, Song JP, He YC (2008). Establishment of Xishuangbanna tropical forest dynamics plot: species compositions and spatial distribution patterns. Journal of Plant Ecology (Chinese Version), 32, 287-298. (in Chinese with English abstract)
	[ 兰国玉, 胡跃华, 曹敏, 朱华, 王洪, 周仕顺, 邓晓保, 崔景云, 黄建国, 刘林云, 许海龙, 宋军平, 何有才 (2008). 西双版纳热带森林动态监测样地——树种组成与空间分布格局. 植物生态学报, 32, 287-298.]
[32]	Lin LX, Comita LS, Zheng Z, Cao M (2012). Seasonal differentiation in density-dependent seedling survival in a tropical rain forest. Journal of Ecology, 100, 905-914.
[33]	Markesteijn L, Poorter L, Paz H, Sack L, Bongers F (2011). Ecological differentiation in xylem cavitation resistance is associated with stem and leaf structural traits. Plant, Cell & Environment, 34, 137-148.
[34]	Mediavilla S, Escudero A, Heilmeier H (2001). Internal leaf anatomy and photosynthetic resource-use efficiency: inter- specific and intraspecific comparisons. Tree Physiology, 21, 251-259. DOI URL PMID
[35]	Meinzer FC (2003). Functional convergence in plant responses to the environment. Oecologia, 134, 1-11. DOI URL PMID
[36]	Moore JP, Vicré-Gibouin M, Farrant JM, Driouich A (2008). Adaptations of higher plant cell walls to water loss: drought vs desiccation. Physiologia Plantarum, 134, 237-245. URL PMID
[37]	Morgan JM (1984). Osmoregulation and water stress in higher plants. Annual Review of Plant Physiology, 35, 299-319.
[38]	Niinemets Ü (1999). Research review. Components of leaf dry mass per area-thickness and density-alter leaf photo- synthetic capacity in reverse directions in woody plants. New Phytologist, 144, 35-47.
[39]	Niinemets Ü (2001). Global-scale climatic controls of leaf dry mass per area, density, and thickness in trees and shrubs. Ecology, 82, 453-469.
[40]	Niinemets Ü, Diaz-Espejo A, Flexas J, Galmes J, Warren CR (2009). Role of mesophyll diffusion conductance in constraining potential photosynthetic productivity in the field. Journal of Experimental Botany, 60, 2249-2270. DOI URL PMID
[41]	Nikolopoulos D, Liakopoulos G, Drossopoulos I, Karabour- niotis G (2002). The relationship between anatomy and photosynthetic performance of heterobaric leaves. Plant Physiology, 129, 235-243. DOI URL PMID
[42]	Onoda Y, Hikosaka K, Hirose T (2004). Allocation of nitrogen to cell walls decreases photosynthetic nitrogen-use efficiency. Functional Ecology, 18, 419-425.
[43]	Parkhurst DF (1994). Diffusion of CO₂ and other gases inside leaves. New Phytologist, 126, 449-479.
[44]	Patakas A, Nikolaou N, Zioziou E, Radoglou K, Noitsakis B (2002). The role of organic solute and ion accumulation in osmotic adjustment in drought-stressed grapevines. Plant Science, 163, 361-367.
[45]	Poorter H, Evans JR (1998). Photosynthetic nitrogen-use efficiency of species that differ inherently in specific leaf area. Oecologia, 116, 26-37. URL PMID
[46]	Poorter H, Niinemets Ü, Poorter L, Wright IJ, Villar R (2009). Causes and consequences of variation in leaf mass per area (LMA): a meta-analysis. New Phytologist, 182, 565-588.
[47]	Poulson ME, Vogelmann TC (1990). Epidermal focussing and effects upon photosynthetic light-harvesting in leaves of oxalis. Plant, Cell & Environment, 13, 803-811.
[48]	Reich P, Wright I, Cavender-Bares J, Craine J, Oleksyn J, Westoby M, Walters M (2003). The evolution of plant functional variation: traits, spectra, and strategies. International Journal of Plant Sciences, 164, S143-S164.
[49]	Richardson AD, Berlyn GP (2002). Changes in foliar spectral reflectance and chlorophyll fluorescence of four temperate species following branch cutting. Tree Physiology, 22, 499-506. DOI URL PMID
[50]	Sack L, Frole K (2006). Leaf structural diversity is related to hydraulic capacity in tropical rain forest trees. Ecology, 87, 483-491. DOI URL PMID
[51]	Sack L, Pasquet-Kok J (2011). Leaf pressure-volume curve parameters. http://prometheuswiki.publish.csiro.au/tiki- index.php?page=Pressure-volume+curves. Cited: 23 Dec. 2013.
[52]	Savé R, Biel C, de Herralde F (2000). Leaf pubescence, water relations and chlorophyll fluorescence in two subspecies of Lotus creticus L. Biologia Plantarum, 43, 239-244. DOI URL
[53]	Scafaro AP, von Caemmerer S, Evans JR, Atwell BJ (2011). Temperature response of mesophyll conductance in cultiv- ated and wild oryza species with contrasting mesophyll cell wall thickness. Plant, Cell & Environment, 34, 1999-2008.
[54]	Smith T, Huston M (1989). A theory of the spatial and temporal dynamics of plant communities. Plant Ecology, 83, 49-69.
[55]	Smith WK, Vogelmann TC, DeLucia EH, Bell DT, Shepherd KA (1997). Leaf form and photosynthesis. BioScience, 47, 785-793.
[56]	Swenson NG, Enquist BJ (2007). Ecological and evolutionary determinants of a key plant functional trait: wood density and its community-wide variation across latitude and elevation. American Journal of Botany, 94, 451-459. DOI URL PMID
[57]	Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S (2011). MEGA5: molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution, 28, 2731-2739. URL PMID
[58]	Terashima I, Hanba YT, Tholen D, Niinemets U (2011). Leaf functional anatomy in relation to photosynthesis. Plant Physiology (Rockville), 155, 108-116. DOI URL PMID
[59]	Tomas M, Flexas J, Copolovici L, Galmes J, Hallik L, Medrano H, Ribas-Carbo M, Tosens T, Vislap V, Niinemets Ü (2013). Importance of leaf anatomy in determining mesophyll diffusion conductance to CO₂ across species: quantitative limitations and scaling up by models. Journal of Experimental Botany, 64, 2269-2281. DOI URL PMID
[60]	Vogelmann TC (1993). Plant tissue optics. Annual Review of Plant Physiology and Plant Molecular Biology, 44, 231-251.
[61]	Vogelmann TC, Bornman JF, Yates DJ (1996a). Focusing of light by leaf epidermal cells. Physiologia Plantarum, 98, 43-56.
[62]	Vogelmann TC, Martin G (1993). The functional-significance of palisade tissue—Penetration of directional versus diffuse light. Plant, Cell & Environment, 16, 65-72.
[63]	Vogelmann TC, Nishio JN, Smith WK (1996b). Leaves and light capture: light propagation and gradients of carbon fixation within leaves. Trends in Plant Science, 1, 65-70.
[64]	Walters M, Reich P (2000). Trade-offs in low-light CO₂ exchange: a component of variation in shade tolerance among cold temperate tree seedlings. Functional Ecology, 14, 155-165.
[65]	Winter H, Robinson DG, Heldt HW (1993). Subcellular volumes and metabolite concentrations in barley leaves. Planta, 191, 180-190.
[66]	Wright IJ, Reich PB, Westoby M, Ackerly DD, Baruch Z, Bongers F, Cavender-Bares J, Chapin T, Cornelissen JH, Diemer M (2004). The worldwide leaf economics spectrum. Nature, 428, 821-827. DOI URL PMID

种 Species	分布 Distribution	丰度 Abundance (Ind.·hm^-2)
长梗三宝木 Trigonostemon thyrsoideus	沟谷 Valley	40.45
粉绿野桐 Mallotus garrettii	沟谷 Valley	34.15
勐腊核果木 Drypetes hoaensis	沟谷 Valley	28.35
棒柄花 Cleidion brevipetiolatum	沟谷 Valley	48.55
秋枫 Bischofia javanica	沟谷 Valley	1.65
缅桐 Sumbaviopsis albicans	山坡 Slope	23.00
轮叶戟 Lasiococca comberi	山坡 Slope	9.20
风轮桐 Epiprinus siletianus	山坡 Slope	6.20
网脉核果木 Drypetes perreticulata	山坡 Slope	1.50
土蜜树 Bridelia tomentosa	山坡 Slope	1.05
木奶果 Baccaurea ramilflora	遍及整个样地 Throughout the entire sample plot	160.60
山地五月茶 Antidesma montanum	遍及整个样地 Throughout the entire sample plot	22.75
日本五月茶 Antidesma japonicum	遍及整个样地 Throughout the entire sample plot	13.75
尾叶血桐 Macaranga kurzii	山脊 Ridge	0.40
越南巴豆 Croton kongensis	山脊 Ridge	7.35
银背巴豆 Croton argyratus	山脊 Ridge	3.00
椴叶山麻杆 Alchornea tiliifolia	山脊 Ridge	18.75
云南银柴 Aporusa yunnanensis	山脊 Ridge	26.40

种 Species	分布 Distribution	丰度 Abundance (Ind.·hm^-2)
长梗三宝木 Trigonostemon thyrsoideus	沟谷 Valley	40.45
粉绿野桐 Mallotus garrettii	沟谷 Valley	34.15
勐腊核果木 Drypetes hoaensis	沟谷 Valley	28.35
棒柄花 Cleidion brevipetiolatum	沟谷 Valley	48.55
秋枫 Bischofia javanica	沟谷 Valley	1.65
缅桐 Sumbaviopsis albicans	山坡 Slope	23.00
轮叶戟 Lasiococca comberi	山坡 Slope	9.20
风轮桐 Epiprinus siletianus	山坡 Slope	6.20
网脉核果木 Drypetes perreticulata	山坡 Slope	1.50
土蜜树 Bridelia tomentosa	山坡 Slope	1.05
木奶果 Baccaurea ramilflora	遍及整个样地 Throughout the entire sample plot	160.60
山地五月茶 Antidesma montanum	遍及整个样地 Throughout the entire sample plot	22.75
日本五月茶 Antidesma japonicum	遍及整个样地 Throughout the entire sample plot	13.75
尾叶血桐 Macaranga kurzii	山脊 Ridge	0.40
越南巴豆 Croton kongensis	山脊 Ridge	7.35
银背巴豆 Croton argyratus	山脊 Ridge	3.00
椴叶山麻杆 Alchornea tiliifolia	山脊 Ridge	18.75
云南银柴 Aporusa yunnanensis	山脊 Ridge	26.40

性状 Trait	山脊 Ridge	山坡 Slope	沟谷 Valley	广布种 Cosmopolitan species
SWC (%)	2.660 ± 0.840^ab	1.860 ± 0.330^a	3.280 ± 1.110^ab	4.260 ± 0.770^b
π_o (MPa)	-1.670 ± 0.180^a	-1.740 ± 0.250^a	-1.190 ± 0.220^b	-1.190 ± 0.040^b
π_tlp (MPa)	-1.950 ± 0.180^a	-1.990 ± 0.220^a	-1.400 ± 0.270^b	-1.400 ± 0.020^b
ε (MPa)	15.21 ± 6.540^a	22.32 ± 7.480^a	18.70 ± 6.300^a	11.88 ± 2.190^a
RWC_tlp (%)	84.62 ± 5.280^a	90.70 ± 4.600^ab	92.61 ± 3.560^b	86.99 ± 0.810^ab
LT (mm)	0.670 ± 0.380^a	0.540 ± 0.140^a	0.900 ± 0.300^a	1.100 ± 0.460^a
UET (mm)	0.069 ± 0.036^a	0.061 ± 0.010^a	0.083 ± 0.017^a	0.164 ± 0.053^b
PT (mm)	0.240 ± 0.130^a	0.150 ± 0.060^a	0.190 ± 0.090^a	0.280 ± 0.060^a
ST (mm)	0.300 ± 0.230^a	0.260 ± 0.100^a	0.540 ± 0.220^a	0.550 ± 0.340^a
LET (mm)	0.063 ± 0.022^a	0.063 ± 0.015^a	0.080 ± 0.018^a	0.101 ± 0.023^a
P/S (%)	1.214 ± 0.611^a	0.670 ± 0.345^a	0.432 ± 0.270^a	0.618 ± 0.208^a
LD (g·cm^-3)	940.4 ± 391.2^ab	1 175.0 ± 385.5^a	514.2 ± 94.2^b	529.5 ± 171.1^ab
LMA (g·cm^-2)	51.36 ± 12.810^a	59.56 ± 14.080^a	44.91 ± 14.750^a	53.49 ± 6.640^a
A_a (μmol·m^-2·s^-1)	10.380 ± 2.310^a	8.400 ± 3.410^a	7.690 ± 1.520^a	7.710 ± 0.750^a
A_m (nmol·g^-1·s^-1)	0.210 ± 0.150^a	0.160 ± 0.110^a	0.190 ± 0.110^a	0.150 ± 0.030^a
R (μmol·m^-2·s^-1)	0.600 ± 0.074^a	0.470 ± 0.074^b	0.500 ± 0.064^ab	0.480 ± 0.046^ab
R_m (nmol·g^-1·s^-1)	0.012 ± 0.003^a	0.008 ± 0.003^a	0.012 ± 0.006^a	0.009 ± 0.001^a

性状 Trait	山脊 Ridge	山坡 Slope	沟谷 Valley	广布种 Cosmopolitan species
SWC (%)	2.660 ± 0.840^ab	1.860 ± 0.330^a	3.280 ± 1.110^ab	4.260 ± 0.770^b
π_o (MPa)	-1.670 ± 0.180^a	-1.740 ± 0.250^a	-1.190 ± 0.220^b	-1.190 ± 0.040^b
π_tlp (MPa)	-1.950 ± 0.180^a	-1.990 ± 0.220^a	-1.400 ± 0.270^b	-1.400 ± 0.020^b
ε (MPa)	15.21 ± 6.540^a	22.32 ± 7.480^a	18.70 ± 6.300^a	11.88 ± 2.190^a
RWC_tlp (%)	84.62 ± 5.280^a	90.70 ± 4.600^ab	92.61 ± 3.560^b	86.99 ± 0.810^ab
LT (mm)	0.670 ± 0.380^a	0.540 ± 0.140^a	0.900 ± 0.300^a	1.100 ± 0.460^a
UET (mm)	0.069 ± 0.036^a	0.061 ± 0.010^a	0.083 ± 0.017^a	0.164 ± 0.053^b
PT (mm)	0.240 ± 0.130^a	0.150 ± 0.060^a	0.190 ± 0.090^a	0.280 ± 0.060^a
ST (mm)	0.300 ± 0.230^a	0.260 ± 0.100^a	0.540 ± 0.220^a	0.550 ± 0.340^a
LET (mm)	0.063 ± 0.022^a	0.063 ± 0.015^a	0.080 ± 0.018^a	0.101 ± 0.023^a
P/S (%)	1.214 ± 0.611^a	0.670 ± 0.345^a	0.432 ± 0.270^a	0.618 ± 0.208^a
LD (g·cm^-3)	940.4 ± 391.2^ab	1 175.0 ± 385.5^a	514.2 ± 94.2^b	529.5 ± 171.1^ab
LMA (g·cm^-2)	51.36 ± 12.810^a	59.56 ± 14.080^a	44.91 ± 14.750^a	53.49 ± 6.640^a
A_a (μmol·m^-2·s^-1)	10.380 ± 2.310^a	8.400 ± 3.410^a	7.690 ± 1.520^a	7.710 ± 0.750^a
A_m (nmol·g^-1·s^-1)	0.210 ± 0.150^a	0.160 ± 0.110^a	0.190 ± 0.110^a	0.150 ± 0.030^a
R (μmol·m^-2·s^-1)	0.600 ± 0.074^a	0.470 ± 0.074^b	0.500 ± 0.064^ab	0.480 ± 0.046^ab
R_m (nmol·g^-1·s^-1)	0.012 ± 0.003^a	0.008 ± 0.003^a	0.012 ± 0.006^a	0.009 ± 0.001^a

	SWC	π_o	π_tlp	ε	RWC_tlp	LT	UET	PT	ST	LET	P/S	LD	LMA	A_a	A_m	R	R_m
SWC	1	0.81^***	0.82^***	-0.25	0.07	0.62^**	0.61^**	-0.05	0.70^**	0.60^*	-0.81^***	-0.71^***	-0.44	0.25	0.51^*	0.19	0.57^*
π_o	0.70^**	1	0.98^***	-0.25	0.20	0.59^*	0.52^*	-0.23	0.74^***	0.37	-0.76^***	-0.71^***	-0.86^***	0.27	0.63^**	-0.11	0.66^**
π_tlp	0.77^***	0.98^***	1	-0.20	0.28	0.55^*	0.53^*	-0.24	0.69^**	0.41	-0.82^***	-0.70^**	-0.79^***	0.18	0.57^**	-0.17	0.61^**
ε	-0.47	-0.68^**	-0.57^*	1	0.81^***	-0.65^**	-0.45	0.01	-0.74^***	-0.44	0.43	0.24	0.66^**	-0.65^**	-0.74^***	-0.37	-0.69^**
RWC_tlp	-0.05	-0.26	-0.10	0.77^***	1	-0.31	-0.18	-0.05	-0.34	-0.06	-0.12	-0.13	0.47	-0.56^*	-0.54^*	-0.44	-0.52^*
LT	0.64^**	0.60^**	0.60^**	-0.13	0.25	1	0.84^***	0.74^***	0.95^***	0.91^***	-0.56^*	-0.78^***	0.33	0.01	-0.25	-0.03	-0.33
UET	0.66^**	0.55^*	0.56^*	-0.31	0	0.71^**	1	0.71^***	0.69^**	0.85^***	-0.45	-0.62^**	0.23	-0.06	-0.23	-0.10	-0.28
PT	0.65^**	0.42	0.43	-0.23	-0.06	0.51^*	0.27	1	0.49^*	0.69^**	0.20	-0.61^**	0.23	0.40	0.05	0.36	-0.08
ST	0.50^*	0.56^*	0.55^*	-0.03	0.37	0.95^***	0.66^**	0.23	1	0.82^***	-0.70^**	-0.73^***	0.32	-0.13	-0.33	-0.17	-0.37
LET	0.54^*	0.52^*	0.54^*	-0.20	0.20	0.83^***	0.63^**	0.60^*	0.71^**	1	-0.50^*	-0.76^***	0.34	-0.03	-0.25	-0.16	-0.38
P/S	-0.21	-0.37	-0.37	-0.37	-0.68^**	-0.54^*	-0.30	-0.00	-0.70^**	-0.57^*	1	0.69^**	0.44	-0.21	-0.45	0.06	-0.41
LD	-0.70^**	-0.89^***	-0.87^***	0.75^***	0.36	-0.75^**	-0.56^*	-0.10	-0.81^***	-0.60^*	0.48^*	1	0.76^***	-0.53^*	-0.81^***	-0.23	-0.82^***
LMA	-0.16	-0.29	-0.27	0.31	0.33	-0.29	-0.24	0.40	-0.47	0	-0.37	0.14	1	-0.22	-0.77^***	0.16	-0.75^**
A_a	0.16	0	-0.04	-0.39	-0.42	0.69^**	0.22	0.58^*	0.61^**	0.54^*	0.35	-0.20	-0.28	1	0.77^***	0.76^***	0.44
A_m	0.14	0.10	0.10	-0.37	-0.37	0.60^*	0.27	0.18	0.62^**	0.40	0.44	-0.18	-0.68^**	0.80^***	1	0.49^*	0.87^***
R	0.20	-0.08	-0.14	-0.41	-0.56^*	0.53^*	0.10	0.70^**	0.38	0.52^*	0.48^*	-0.18	-0.24	0.59^**	0.52^*	1	-0.62^**
R_m	0.16	0.12	0.12	-0.36	-0.40	0.50^*	0.25	0.04	0.56^*	0.30	0.49^*	-0.15	-0.86^***	0.58^*	0.92^***	-0.45	1