doi:

DOI: 10.3724/SP.J.1258.2012.01205

Chinese Journal of Plant Ecology (植物生态学报) 2012/36:11 PP.1205-1216

Advances research in plant nitrogen, phosphorus and their stoichiometry in terrestrial eco-systems: a review


Abstract:
Nitrogen (N), phosphorus (P) and their stoichiometry play pivotal roles in plant structure and functions, develop-ment and ecological strategies in terrestrial ecosystems due to their coupling with each other and their irreplace-ability. Plant N and P can be influenced by biotic and abiotic factors, such as individual traits, climate change and human disturbance, and it is those factors that determine the plant community composition and structure that fi-nally affect the ecosystem processes. According to previous studies, there is an allometric relationship between N and P. Relationships between plant N and P depend on the soil nutrient condition and species plasticity in N and P. Understanding the relationships between plant N and P in major ecological gradients can further our knowledge about vegetation restoration, succession, biodiversity, ecosystem trophic structure and biogeochemical cycles. This information could help predict potential changes in terrestrial ecosystems in response to future climate change. We review recent advances in the influencing factors and mechanism of stoichiometry in order to improve understanding of plant responses to global change.

Key words:allometric relationship, ecological stoichiometry, homeostasis, N:P, plant functional group

ReleaseDate:2014-07-21 16:32:44



Aerts R (1996). Nutrient resorption from senescing leaves of perennials: Are there general patterns? Journal of Ecology, 84, 597-608.

Aerts R (1997). Nitrogen partitioning between resorption and decomposition pathways: a trade-off between nitrogen use efficiency and litter decomposability? Oikos, 80, 603-606.

Aerts R, Chapin FS III (2000). The mineral nutrition of wild plants revisited: a re-evaluation of processes and patterns. Advances in Ecological Research, 30, 1-67.

Ågren GI (2004). The C:N:P stoichiometry of autotrophs―theory and observations. Ecology Letters, 7, 185-191.

Ågren GI (2008). Stoichiometry and nutrition of plant growth in natural communities. Annual Review of Ecology, Evolution, and Systematics, 39, 153-170.

Ågren GI, Weih M (2012). Plant stoichiometry at different scales: element concentration patterns reflect environment more than genotype. New Phytologist, 194, 944-952.

Anderson TM, Ritchie ME, Mayemba E, Eby S, Grace JB, McNaughton SJ (2007). Forage nutritive quality in the Serengeti ecosystem: the roles of fire and herbivory. The American Naturalist, 170, 343-357.

Bedford BL, Walbridge MR, Aldous A (1999). Patterns in nutrient availability and plant diversity of temperate North American wetlands. Ecology, 80, 2151-2169.

Cech PG, Kuster T, Edwards PJ, Venterink HO (2008). Effects of herbivory, fire and N2-fixation on nutrient limitation in a humid African savanna. Ecosystems, 11, 991-1004.

Cheng DL, Li T, Zhong QL, Wang GX (2010). Scaling relationship between tree respiration rates and biomass. Biology Letters, 6, 715-717.

Cheng DL, Wang GX, Zhong QL (2009). Age-related relationship between annual productivity and body size of trees: testing the metabolic theory. Polish Journal of Ecology, 57, 441-449.

Cotrufo MF, Ineson P, Scott A (1998). Elevated CO2 reduces nitrogen concentration of plant tissues. Global Change Biology, 4, 43-54.

Cui Q, Lü XT, Wang QB, Han XG (2010). Nitrogen fertilization and fire act independently on foliar stoichiometry in a temperate steppe. Plant and Soil, 334, 209-219.

Demars BOL, Edwards AC (2007). Tissue nutrient concentrations in freshwater aquatic macrophytes: high intertaxon differences and low phenotypic response to nutrient supply. Freshwater Biology, 52, 2073-2086.

Dickman EM, Newell JM, González MJ, Vanni MJ (2008). Light, nutrients, and food-chain length constrain planktonic energy transfer efficiency across multiple trophic levels. Proceedings of the National Academy of Sciences of the United States of America, 105, 18408-18412.

Dong XY (董晓玉), Fu H (傅华), Li XD (李旭东), Niu DC (牛得草), Guo D (郭丁), Li XD (李晓东) (2010). Effects on plant biomass and CNP contents of plants in grazed and fenced steppe grasslands of the Loess Plateau. Acta Pra-taculturae Sinica (草业学报), 19, 175-182. (in Chinese with English abstract)

Drenovsky RE, Richards JH (2006). Low leaf N and P resorption contributes to nutrient limitation in two desert shrubs. Plant Ecology, 183, 305-314.

El-Kahloun M, Boeye D, Verhagen B, van Haesebroeck V (2000). A comparison of the nutrient status of Molinia caerulea and neighbouring vegetation in a rich fen. Belgian Journal of Botany, 133, 91-102.

Elser JJ, Acharya K, Kyle M, Cotner J, Makino W, Markow T, Watts T, Hobbie S, Fagan W, Schade J, Hood J, Sterner RW (2003). Growth rate-stoichiometry couplings in diverse biota. Ecology Letters, 6, 936-943.

Elser JJ, Fagan WF, Kerkoff AJ, Swenson NG, Enquist BJ (2010). Biological stoichiometry of plant production: metabolism, scaling and ecological response to global change. New Phytologist, 186, 593-608.

Elser JJ, Hamilton A (2007). Stoichiometry and the new biology: the future is now. PLoS Biology, 5, 1403-1405.

Elser JJ, Sterner RW, Gorokhova E, Fagan WF, Markow TA, Cotner JB, Harrison JF, Hobbie SE, Odell GM, Weider LJ (2000). Biological stoichiometry from genes to ecosystems. Ecology Letters, 3, 540-550.

Falkowski P, Scholes RJ, Boyle E, Canadell J, Canfield D, Elser J, Gruber N, Hibbard K, Hogberg P, Linder S, Mackenzie FT, Moore B III, Pedersen T, Rosenthal Y, Seitzinger S, Smetacek V, Steffen W (2000). The global carbon cycle: a test of our knowledge of earth as a system. Science, 290, 291-296.

Field CB, Mooney HA (1986). The photosynthesis-nitrogen relationship in wild plants. In: Givnish TJ ed. On the Economy of Plant Form and Function. Cambridge University Press, London. 25-55

Finzi AC, Delucia EH, Shlesinger WH (2004). Canopy N and P dynamics of a south-eastern US pine forest under elevated CO2. Biogeochemistry, 69, 363-378.

Frank DA (2008). Ungulate and topographic control of nitrogen: phosphorus stoichiometry in a temperate grassland; soils, plants and mineralization rates. Oikos, 117, 591-601.

Fujita Y, Robroek BJM, de Ruiter PC, Heil GW, Wassen MJ (2010). Increased N affects P uptake of eight grassland species: the role of root surface phosphatase activity. Oikos, 119, 1665-1673.

Gao SP (高三平), Li JX (李俊祥), Xu MC (徐明策), Chen X (陈熙), Dai J (戴洁) (2007). Leaf N and P stoichiometry of common species in successional stages of the evergreen broad-leaved forest in Tiantong National Forest Park, Zhejiang Province, China. Acta Ecologica Sinica (生态学报), 27, 947-952. (in Chinese with English abstract)

Granath G, Strengbom J, Breeuwer A, Heijmans MMPD, Berendse F, Rydin H (2009). Photosynthetic performance in Sphagnum transplanted along a latitudinal nitrogen deposition gradient. Oecologia, 159, 705-715.

Gressel N, McColl JG (2003). Phosphorus mineralisation and organic matter decomposition: a critical review. In: Cadisch G, Giller KE eds. Driven by Nature: Plant Litter Quality and Decomposition. CAB International, Walling-ford, UK. 297-309.

Gruber N, Galloway JN (2008). An earth-system perspective of the global nitrogen cycle. Nature, 451, 293-296.

Güsewell S (2004). N:P ratios in terrestrial plants: variation and functional significance. New Phytologist, 164, 243-266.

Güsewell S, Bollens U (2003). Composition of plant species mixtures grown at various N:P ratios and levels of nutrient supply. Basic and Applied Ecology, 4, 453-466.

Güsewell S, Koerselman W (2002). Variation in nitrogen and phosphorus concentrations of wetland plants. Perspectives in Plant Ecology, Evolution and Systematics, 5, 37-61.

Güsewell S, Koerselman W, Verhoeven JTA (2003). Biomass N:P ratios as indicators of nutrient limitation for plant populations in wetlands. Ecological Applications, 13, 372-384.

Han WX, Fang JY, Guo DL, Zhang Y (2005). Leaf nitrogen and phosphorus stoichiometry across 753 terrestrial plant species in China. New Phytologist, 168, 377-385.

Harpole SW, Ngai JT, Cleland EE, Seabloom EW, Borer ET, Bracken MES, Elser JJ, Gruner DS, Hillebrand H, Shurin JB, Smith JE (2011). Nutrient co-limitation of primary producer communities. Ecology Letters, 14, 852-862.

He JS (贺金生), Han XG (韩兴国) (2010). Ecological stoichiometry: searching for unifying principles from individuals to ecosystems. Chinese Journal of Plant Ecology (植物生态学报), 34, 2-6. (in Chinese with English abstract)

He JS, Fang JY, Wang ZH, Guo DL, Flynn DFB, Geng Z (2006). Stoichiometry and large-scale patterns of leaf carbon and nitrogen in the grassland biomes of China. Oecologia, 149, 115-122.

He JS, Wang L, Flynn DFB, Wang XP, Ma WH, Fang JY (2008). Leaf nitrogen: phosphorus stoichiometry across Chinese grassland biomes. Oecologia, 155, 301-310.

He NP, Han XG, Yu GR, Chen QS (2011). Divergent changes in plant community composition under 3-decade grazing exclusion in continental steppe. PLoS One, 6(11), e26506.

Hessen DO, Ågren GI, Anderson TR, Elser JJ, de Ruiter PC (2004). Carbon sequestration in ecosystems: the role of stoichiometry. Ecology, 85, 1179-1192.

Hobbie SE, Gough L (2002). Foliar and soil nutrients in tundra on glacial landscapes of contrasting ages in northern Alaska. Oecologia, 131, 453-462.

Hogan EJ, Minnullina G, Smith RI, Crittenden PD (2010). Effects of nitrogen enrichment on phosphatase activity and nitrogen: phosphorus relationships in Cladonia porten-tosa. New Phytologist, 186, 911-925.

Huang JJ (黄建军), Wang XH (王希华) (2003). Leaf nutrient and structural characteristics of 32 evergreen broad-leaved species. Journal of East China Normal University (Natural Science Edition) (华东师范大学学报(自然科学版)), (1), 92-97. (in Chinese with English abstract)

Huang JY, Yu HL, Li LH, Yuan ZY, Bartels S (2009). Water supply changes N and P conservation in a perennial grass Leymus chinensis. Journal of Integrative Plant Biology, 51, 1050-1056.

Huang WJ, Zhou GY, Liu JX, Zhang DQ, Xu ZH, Liu SZ (2012). Effects of elevated carbon dioxide and nitrogen addition on foliar stoichiometry of nitrogen and phosphorus of five tree species in subtropical model forest ecosystems. Environmental Pollution, 168, 113-120.

Johnson DW, Cheng W, Joslin JD, Norby RJ, Edwards NT, Todd JDE (2004). Effects of elevated CO2 on nutrient cycling in a sweetgum plantation. Biogeochemistry, 69, 379-403.

Kerkhoff AJ, Enquist BJ (2006). Ecosystem allometry: the scaling of nutrient stocks and primary productivity across plant communities. Ecology Letters, 9, 419-427.

Kerkhoff AJ, Enquist BJ, Elser JJ, Fagan WF (2005). Plant allometry, stoichiometry and the temperature-dependence of primary productivity. Global Ecology and Biogeography, 14, 585-598.

Kerkhoff AJ, Fagan WF, Elser JJ, Enquist JB (2006). Phylogenetic and growth form variation in the scaling of nitrogen and phosphorus in the seed plants. The American Naturalist, 168, 103-122.

Koerselman W, Meuleman AFM (1996). The vegetation N:P ratio: a new tool to detect the nature of nutrient limitation. Journal of Applied Ecology, 33, 1441-1450.

Körner C, Asshoff R, Bignucolo O, Hattenschwiler S, Keel SG, Peláez-Riedl S, Pepin S, Siegwolf RTW, Zotz G (2005). Carbon flux and growth in mature deciduous forest trees exposed to elevated CO2. Science, 309, 1360-1362.

Ladanai S, Ågren GI, Olsson BA (2010). Relationships between tree and soil properties in Picea abies and Pinus sylvestris forests in Sweden. Ecosystems, 13, 302-316.

Lagomarsino A, Moscatelli MC, Hoosbeek MR, de Angelis P, Grego S (2008). Assessment of soil nitrogen and phos-phorous availability under elevated CO2 and N-fertilization in a short rotation poplar plantation. Plant and Soil, 308, 131-147.

Lal R (2004). Soil carbon sequestration impacts on global cli-mate change and food security. Science, 304, 1623-1627.

Lambers H, Raven JA, Shaver GR, Smith SE (2008). Plant nutrient-acquisition strategies change with soil age. Trends in Ecology and Evolution, 23, 95-103.

Larsen KS, Andresen LC, Beier C, Jonasson S, Albert KR, Ambus P, Arndal MF, Carter MS, Christensen S, Holm-strup M, Ibrom A, Kongstad J, van der Linden L, Maraldo K, Michelsen A, Mikkelsen TN, Pilegaard K, Priemé A, RoPoulsen H, Schmidt IK, Selsted MB, Stevnbak K (2011). Reduced N cycling in response to elevated CO2, warming, and drought in a Danish heathland: synthesizing results of the climaite project after two years of treat-ments. Global Change Biology, 17, 1884-1899.

LeBauer DS, Treseder KK (2008). Nitrogen limitation of net primary productivity in terrestrial ecosystems is globally distributed. Ecology, 89, 371-379.

Li YL (李玉霖), Mao W (毛伟), Zhao XY (赵学勇), Zhang TH (张铜会) (2010). Leaf nitrogen and phosphorus stoichiometry in typical desert and desertified regions, North China. Environmental Science (环境科学), 31, 1716-1725. (in Chinese with English abstract)

Limpens J, Berendse F, Klees H (2003). N deposition affects N availability in interstitial water, growth of Sphagnum and invasion of vascular plants in bog vegetation. New Phy-tologist, 157, 339-347.

Liu P, Huang JH, Han XG, Sun OJ, Zhou ZY (2006). Differential responses of litter decomposition to increased soil nutrients and water between two contrasting grassland plant species of Inner Mongolia, China. Applied Soil Ecology, 34, 266-275.

Liu WD (刘万德), Su JR (苏建荣), Li SF (李帅锋), Zhang ZJ (张志钧), Li ZW (李忠文) (2010). Stoichiometry study of C, N and P in plant and soil at different successional stages of monsoon evergreen broad-leaved forest in Pu’er Yunnan Province. Acta Ecologica Sinica (生态学报), 30, 6581-6590. (in Chinese with English abstract)

Liu XZ (刘兴诏), Zhou GY (周国逸), Zhang DQ (张德强), Liu SZ (刘世忠), Chu GW (褚国伟), Yan JH (闫俊华) (2010). N and P stoichiometry of plant and soil in lower subtropical forest successional series in southern China. Chinese Journal of Plant Ecology (植物生态学报), 34, 64-71. (in Chinese with English abstract)

Loladze I (2002). Rising atmospheric CO2 and human nutrition: Toward globally imbalanced plant stoichiometry? Trends in Ecology and Evolution, 17, 457-461.

Lu J (鲁静), Zhou HX (周虹霞), Tian GY (田广宇), Liu GH (刘贵华) (2011). Nitrogen and phosphorus contents in 44 wetland species from the Lake Erhai Basin. Acta Ecologica Sinica (生态学报), 31, 709-715. (in Chinese with English abstract)

Lü XT, Kong DL, Pan QM, Simmons ME, Han XG (2012a). Nitrogen and water availability interact to affect leaf stoichiometry in a semi-arid grassland. Oecologia, 168, 301-310.

Lü XT, Lü FM, Zhou LS, Han X, Han XG (2012b). Stoichiometric response of dominant grasses to fire and mowing in a semi-arid grassland. Journal of Arid Envi-ronments, 78, 154-160.

Lukac M, Calfapietra C, Lagomarsino A, Loreto F (2010). Global climate change and tree nutrient: effects of elevated CO2 and temperature. Tree Physiology, 30, 1209-1220.

Makarieva AM, Gorshkov VG, Li BL, Chown SL, Reich PB, Gavrilov VM (2008). Mean mass-specific metabolic rates are strikingly similar across life’s major domains: evidence for life’s metabolic optimum. Proceedings of the National Academy of Sciences of the United States of America, 105, 16994-16999.

Matson PA, McDowell WH, Townsend AR, Vitousek PM (1999). The globalization of N deposition: ecosystem consequences in tropical environments. Biogeochemistry, 46, 67-83.

Matzek V, Vitousek PM (2009). N:P stoichiometry and protein: RNA ratios in vascular plants: an evaluation of the growth-rate hypothesis. Ecology Letters, 12, 765-771.

McGroddy ME, Daufresne T, Hedin LO (2004). Scaling of C:N:P stoichiometry in forest worldwide: implications of terrestrial Redfield-type ratios. Ecology, 85, 2390-2401.

Nguyen NT, Mohapatra PK, Fujita K (2006). Elevated CO2 alleviates the effects of low P on the growth of N2-fixing Acacia auriculiformis and Acacia mangium. Plant and Soil, 285, 369-379.

Niklas KJ (2006). Plant allometry, leaf nitrogen and phosphorus stoichiometry, and interspecific trends in annual growth rates. Annals of Botany, 97, 155-163.

Niklas KJ, Owens T, Reich PB, Cobb ED (2005). Nitrogen/ phosphorus leaf stoichiometry and the scaling of plant growth. Ecology Letters, 8, 636-642.

Novotny AM Schade JD, Hobbie SE, Kay AD, Kyle M, Reich PB, Elser JJ (2007). Stoichiometric response of nitrogen-fixing and non-fixing dicots to manipulations of CO2, ni-trogen, and diversity. Oecologia, 151, 687-696.

Olsen JK, Bell LC (1990). A glasshouse evaluation of ‘critical’ N and P concentrations and N:P ratios in various plant parts of six Eucalypt species. Australian Journal of Botany, 38, 291-298.

Ordoñez JC, van Bodegom PM, Witte JPM, Wright IJ, Reich PB, Aerts R (2009). A global study of relationships between leaf traits, climate and soil measures of nutrient fertility. Global Ecology and Biogeography, 18, 137-149.

Orgeas J, Ourcival JM, Bonin G (2002). Seasonal and spatial patterns of foliar nutrients in cork oak (Quercus suber L.) growing on siliceous soils in Provence (France). Plant Ecology, 164, 201-211.

Ostertag R (2001). Effects of nitrogen and phosphorus avail-ability on fine-root dynamics in Hawaiian montane forests. Ecology, 82, 485-499.

Perring MP, Hedin LO, Levin SA, McGroddy M, de Mazan-court C (2008). Increased plant growth from nitrogen ad-dition should conserve phosphorus in terrestrial ecosystems. Proceedings of the National Academy of Sciences of the United States of America, 105, 1971-1976.

Polley HW, Fay PA, Jin VL, Combs GF Jr (2011). CO2 enrichment increases element concentrations in grass mix-tures by changing species abundances. Plant Ecology, 212, 945-957.

Qin H (秦海), Li JX (李俊祥), Gao SP (高三平), Li C (李铖), Li R (李蓉), Shen XH (沈兴华) (2010). Characteristics of leaf element contents for eight nutrients across 660 terres-trial plant species in China. Acta Ecologica Sinica (生态学报), 30, 1247-1257. (in Chinese with English abstract)

Reich PB, Hobbie SE, Ellsworth DS, West JB, Tilman D, Knops JMH, Naeem S, Trost J (2006a). Nitrogen limitation constrains sustainability of ecosystem response to CO2. Nature, 440, 922-925.

Reich PB, Oleksyn J (2004). Global patterns of plant leaf N and P in relation to temperature and latitude. Proceedings of the National Academy of Sciences of the United States of America, 101, 11001-11006.

Reich PB, Oleksyn J, Wright IJ, Niklas KJ, Hedin L, Elser JJ (2010). Evidence of a general 2/3-power law of scaling leaf nitrogen to phosphorus among major plant groups and biomes. Proceedings of the Royal Society B: Biological Sciences, 277, 877-883.

Reich PB, Tjoelker MG, Machado JL, Oleksyn J (2006b). Universal scaling of respiratory metabolism, size and nitrogen in plants. Nature, 439, 457-461.

Rejmánková E (2005). Nutrient resorption in wetland macro-phytes: comparison across several regions of different nutrient status. New Phytologist, 167, 471-482.

Ren SJ (任书杰), Yu GR (于贵瑞), Jiang CM (姜春明), Fang HJ (方华军), Sun XM (孙晓敏) (2012). Stoichiometric characteristics of leaf carbon, nitrogen, and phosphorus of 102 dominant species in forest ecosystems along the North-South Transect of East China. Chinese Journal of Applied Ecology (应用生态学报), 23, 581-586. (in Chinese with English abstract)

Ren SJ (任书杰), Yu GR (于贵瑞), Tao B (陶波), Guan LL (官丽莉), Fang HJ (方华军), Jiang CM (姜春明) (2009). Spatial patterns for variations in leaf nutrient contents of Dahurian larch (Larix gmelinii Rupr.). Acta Ecologica Sinica (生态学报), 29, 1899-1906. (in Chinese with Eng-lish abstract)

Ren SJ (任书杰), Yu GR (于贵瑞), Tao B (陶波), Wang SQ (王绍强) (2007). Leaf nitrogen and phosphorus stoichiometry across 654 terrestrial plant species in NSTEC. Envi-ronmental Science (环境科学), 28, 2665-2673. (in Chinese with English abstract)

Richardson SJ, Allen RB, Doherty JE (2008). Shifts in leaf N:P ratio during resorption reflect soil P in temperate rainforest. Functional Ecology, 22, 738-745.

Sardans J, Peñuelas J (2010). Soil enzyme activity in a Mediterranean forest after six years of drought. Soil Science Society of America Journal, 74, 838-851.

Sardans J, Rivas-Ubach A, Peñuelas J (2012). The C:N:P stoichiometry of organisms and ecosystems in a changing world: a review and perspectives. Perspectives in Plant Ecology, Evolution and Systematics, 14, 33-47.

Schachtman DP, Reid RJ, Ayling SM (1998). Phosphorus up-take by plants: from soil to cell. Plant Physiology, 116, 447-453.

Schipper LA, Schipper LA, Percival HJ, Sparling GP (2004). An approach for estimating when soils will reach maximum nitrogen storage. Soil Use and Management, 20, 281-286.

Shi JY (施家月), Wang XH (王希华), Yan ER (阎恩荣), Cheng MH (程铭华) (2006). Saplings nutrient characteristics of common plants in Tiantong National Forest Park. Journal of East China Normal University (Natural Science Edition) (华东师范大学学报(自然科学版)), (2), 121- 129. (in Chinese with English abstract)

Sims L, Pastor J, Lee T, Dewey B (2012). Nitrogen, phosphorus and light effects on growth and allocation of biomass and nutrients in wild rice. Oecologia, 170, 65-76.

Song YT (宋彦涛), Zhou DW (周道玮), Li Q (李强), Wang P (王平), Huang YX (黄迎新) (2012). Leaf nitrogen and phosphorus stoichiometry in 80 herbaceous plant species of Songnen grassland in Northeast China. Chinese Journal of Plant Ecology (植物生态学报), 36, 222-230. (in Chinese with English abstract)

Sterner RW, Elser JJ (2002). Ecological Stoichiometry: the Biology of Elements from Molecules to the Biosphere. Princeton University Press, Princeton. 906-907.

Stock WD, Verboom GA (2012). Phylogenetic ecology of foliar N and P concentrations and N:P ratios across mediterranean-type ecosystems. Global Ecology and Biogeography.

Striebel M, Sporl G, Stibo H (2008). Light-induced changes of plankton growth and stoichiometry: experiments with natural phytoplankton communities. Limnology and Oceanography, 53, 513-522.

Taub DR, Wang XZ (2008). Why are nitrogen concentrations in plant tissues lower under elevated CO2? A critical ex-amination of the hypotheses. Journal of Integrative Plant Biology, 50, 1365-1374.

Tessier JT, Raynal DY (2003). Use of nitrogen to phosphorus ratios in plant tissue as an indicator of nutrient limitation and nitrogen saturation. Journal of Applied Ecology, 40, 523-534.

Tjoelker MG, Oleksyn J, Reich PB, Żytkowiak R (2008). Cou-pling of respiration, nitrogen, and sugars underlies convergent temperature acclimation in Pinus banksiana across wide-ranging sites and populations. Global Change Biology, 14, 782-797.

Townsend AR, Cleveland CC, Asner GP, Bustamante MMC (2007). Controls over foliar N:P ratios in tropical rain forests. Ecology, 88, 107-118.

Usuda H (1995). Phosphate deficiency in maize. V. Mobilization of nitrogen and phosphorus within shoots of young plants and its relationship to senescence. Plant and Cell Physiology, 36, 1041-1049.

Valladares F, Martinez-Ferri E, Balaguer L, Perez-Corona E, Manrique E (2000). Low leaf-level response to light and nutrients in Mediterranean evergreen oaks: a conservative resource-use strategy? New Phytologist, 148, 79-91.

Venterink HO, Wassen MJ, Verkroost AWM, de Ruiter PC (2003). Species richness-productivity patterns differ between N-, P-, and K-limited wetlands. Ecology, 84, 2191-2199.

Vitousek PM (1982). Nutrient cycling and nutrient use effi-ciency. The American Naturalist, 119, 553-572.

Vitousek PM (1984). Litterfall, nutrient cycling, and nutrient limitation in tropical forests. Ecology, 65, 285-298.

Vitousek PM, Aber JD, Howarth RW, Likens GE, Matson PA, Schindler DW, Schlesinger WH, Tilman DG (1997). Hu-man alteration of the global nitrogen cycle: sources and consequences. Ecological Applications, 7, 737-750.

Vitousek PM, Hobbie S (2000). Heterotrophic nitrogen fixation in decomposing litter: patterns and regulation. Ecology, 81, 2366-2376.

Vitousek PM, Porder S, Houlton BZ, Chadwick OA (2010). Terrestrial phosphorus limitation: mechanisms, implications, and nitrogen-phosphorus interactions. Ecological Applications, 20, 5-15.

Vrede T, Dobberfuhl DR, KooijmanS ALM, Elser JJ (2004). Fundamental connections among organism C:N:P stoichiometry, macromolecular composition, and growth. Ecology, 85, 1217-1229.

Wang JY (王晶苑), Wang SQ (王绍强), Li RL (李纫兰), Yan JH (闫俊华), Sha LQ (沙丽清), Han SJ (韩士杰) (2011). C:N:P stoichiometric characteristics of four forest types’ dominant tree species in China. Chinese Journal of Plant Ecology (植物生态学报), 35, 587-595. (in Chinese with English abstract)

Wang N, Xu SS, Jia X, Gao J, Zhang WP, Qiu YP, Wang GX (2012). Variations in foliar stable carbon isotopes among functional groups and along environmental gradients in China—a meta-analysis. Plant Biology, doi: 10.1111/ j.1438-8677.2012.00605.x.

Wang SQ (王绍强), Yu GR (于贵瑞) (2008). Ecological stoichiometry characteristics of ecosystem carbon, nitro-gen and phosphorus elements. Acta Ecologica Sinica (生态学报), 28, 3937-3947.

Wang T (汪涛), Yang YH (杨元合), Ma WH (马文红) (2008). Storage, patterns and environmental controls of soil phos-phorus in China. Acta Scientiarum Naturalium Universita-tis Pekinensis (北京大学学报(自然科学版)), 44, 945- 952. (in Chinese with English abstract)

Wang YP, Law RM, Pak B (2010). A global model of carbon, nitrogen and phosphorus cycles for the terrestrial bio-sphere. Biogeosciences, 7, 2261-2282.

Wassen MJ, Olde Venterink HGM, de Swart EOAM (1995). Nutrient concentrations in mire vegetation as a measure of nutrient limitation in mire ecosystems. Journal of Vegetation Science, 6, 5-16.

Watanabe T, Broadley MR, Jansen S, White PJ, Takada J, Sa-take K, Takamatsu T, Tuah SJ, Osaki M (2007). Evolutionary control of leaf element composition in plants. New Phytologist, 174, 516-523.

Wright IJ, Reich PB, Cornelissen JHC, Falster DS, Groom PK, Hikosaka K, Lee W, Lusk CH, Niinemets Ü, Oleksyn J, Osada N, Poorter H, Warton DI, Westoby M (2005). Modulation of leaf economic traits and trait relationships by climate. Global Ecology and Biogeography, 14, 411- 421.

Wright IJ, Reich PB, Westoby M, Ackerly DD, Baruch Z, Bongers F, Cavender-Bares J, Chapin T, Cornelissen JHC, Diemer M, Flexas J, Garnier E, Groom PK, Gulias J, Hi-kosaka K, Lamont BB, Lee T, Lee W, Lusk C, Midgley JJ, Navas ML, Niinemets Ü, Oleksyn J, Osada N, Poorter H, Poot P, Prior L, Pyankov VI, Roumet C, Thomas SC, Tjoelker MG, Veneklaas EJ, Villar R (2004). The world-wide leaf economics spectrum. Nature, 428, 821-827.

Wu GL, Li W, Shi ZH, Shangguan ZP (2011). Aboveground dominant functional group predicts belowground proper-ties in an alpine grassland community of western China. Journal of Soils and Sediments, 11, 1011-1019.

Wu TG (吴统贵), Wu M (吴明), Li L (刘丽), Xiao JH (萧江华) (2010). Seasonal variations of leaf nitrogen and phos-phorus stoichiometry of three herbaceous species in Hangzhou Bay coastal wetlands, China. Chinese Journal of Plant Ecology (植物生态学报), 34, 23-28. (in Chinese with English abstract)

Xia JY, Wan SQ (2008). Global response patterns of terrestrial plant species to nitrogen addition. New Phytologist, 179, 428-439.

Xu B (徐冰), Cheng YX (程雨曦), Gan HJ (甘慧洁), Zhou WJ (周文嘉), He JS (贺金生) (2010). Correlations between leaf and fine root traits among and within species of typi-cal temperate grassland in Xilin River Basin, Inner Mon-golia, China. Chinese Journal of Plant Ecology (植物生态学报), 34, 29-38. (in Chinese with English abstract)

Yan ER (阎恩荣), Wang XH (王希华), Guo M (郭明), Zhong Q (仲强), Zhou W (周武) (2010). C:N:P stoichiometry across evergreen broad-leaved forests, evergreen conifer-ous forests and deciduous broad-leaved forests in the Tiantong region, Zhejiang Province, eastern China. Chi-nese Journal of Plant Ecology (植物生态学报), 34, 48- 57. (in Chinese with English abstract)

Yan ER (阎恩荣), Wang XH (王希华), Zhou W (周武) (2008). N:P stoichiometry in secondary succession in evergreen broad-leaved forest, Tiantong, Eastern China. Journal of Plant Ecology (Chinese Version) (植物生态学报), 32, 13-22. (in Chinese with English abstract)

Yang HM (杨慧敏), Wang DM (王冬梅) (2011). Advances in the study on ecological stoichiometry in grass-environ- ment system and its response to environmental factors. Acta Prataculturae Sinica (草业学报), 20, 244-252. (in Chinese with English abstract)

Yin XR (银晓瑞), Liang CZ (梁存柱), Wang LX (王立新), Wang W (王炜), Liu ZL (刘钟龄), Liu XP (刘小平) (2010). Ecological stoichiometry of plant nutrients at dif-ferent restoration succession stages in typical steppe of Inner Mongolia, China. Chinese Journal of Plant Ecology (植物生态学报), 34, 39-47. (in Chinese with English ab-stract)

Yuan ZY, Chen HYH, Reich PB (2011). Global-scale latitu-dinal patterns of plant fine-root nitrogen and phosphorus. Nature Communications, 2, 344.

Zeng DH (曾德慧), Chen GX (陈广生) (2005). Ecological stoichiometry: a science to explore the complexity of liv-ing systems. Acta Phytoecologica Sinica (植物生态学报), 29, 1007-1019. (in Chinese with English abstract)

Zhang LX, Bai YF, Han XG (2004). Differential responses of N:P stoichiometry of Leymus chinensis and Carex kor-shinskyi to N additions in a steppe ecosystem in Nei Mongol. Acta Botanica Sinica, 46, 259-270.

Zhang SB, Zhang JL, Ferry Slik JW, Cao KF (2012). Leaf ele-ment concentrations of terrestrial plants across China are influenced by taxonomy and the environment. Global Ecology and Biogeography, 21, 809-818.

Zheng SX, Shangguan ZP (2007). Spatial patterns of leaf nu-trient traits of the plants in the Loess Plateau of China. Trees-Structure and Function, 21, 357-370.

Zhou P (周鹏), Geng Y (耿燕), Ma WH (马文红), He JS (贺金生) (2010). Linkages of functional traits among plant or-gans in the dominant species of the Inner Mongolia grass-land, China. Chinese Journal of Plant Ecology (植物生态学报), 34, 7-16. (in Chinese with English abstract)

PDF