Abstract
Nitrogen (N) and carbon–nitrogen (C:N) ratio are key foliar traits with great ecological importance, but their patterns across biomes have only recently been explored. We conducted a systematic census of foliar C, N and C:N ratio for 213 species, from 41 families over 199 research sites across the grassland biomes of China following the same protocol, to explore how different environmental conditions and species composition affect leaf N and C:N stoichiometry. Leaf C:N stoichiometry is stable in three distinct climatic regions in Inner Mongolia, the Tibetan Plateau, and Xinjiang Autonomous Region, despite considerable variations among co-existing species and among different vegetation types. Our results also show that life form and genus identity explain more than 70% of total variations of foliar N and C:N ratio, while mean growing season temperature and growing season precipitation explained only less than 3%. This suggests that, at the biome scale, temperature affects leaf N mainly through a change in plant species composition rather than via temperature itself. When our data were pooled with a global dataset, the previously observed positive correlation between leaf N and mean annual temperature (MAT) at very low MATs, disappeared. Thus, our data do not support the previously proposed biogeochemical hypothesis that low temperature limitations on mineralization of organic matter and N availability in soils lead to low leaf N in cold environments.
Similar content being viewed by others
References
Ackerly DD (2004) Adaptation, niche conservatism, and convergence: comparative studies of leaf evolution in the California chaparral. Am Nat 163:654–671
Aerts R, Chapin FS III (2000) The mineral nutrition of wild plants revisited: a re-evaluation of processes and patterns. Adv Ecol Res 30:1–67
Ågren GI (2004) The C:N:P stoichiometry of autotrophs—theory and observations. Ecol Lett 7:185–191
Brown JH, Lomolino MV (1998) Biogeography. Sinauer, Sunderland
Chadwick O, Derry L, Vitousek P, Huebert B, Hedin L (1999) Changing sources of nutrients during four million years of ecosystem development. Nature 397:491–497
Chapin FS III (1980) The mineral nutrition of wild plants. Annu Rev Ecol Syst 11:233–260
Chen ZZ, Wang SP (2000) Typical steppe ecosystems of China. Science Press, Beijing
Chown SL, Gaston KJ, Robinson D (2004) Macrophysiology: large-scale patterns in physiological traits and their ecological implications. Funct Ecol 18:159–167
Elser JJ, Fagan WF, Denno RF, Dobberfuhl DR, Folarin A, Huberty A, Interlandi S, Kilham SS, McCauley E, Schulz KL, Siemann EH, Sterner RW (2000) Nutritional constraints in terrestrial and freshwater food webs. Nature 408:578–580
Fang J, Piao S, Tang Z, Peng C, Ji W (2001) Interannual variability in net primary production and precipitation. Science 293:1723a
Güsewell S (2004) N:P ratios in terrestrial plants: variation and functional significance. New Phytol 164:243–266
Han W, Fang JY, Guo D, Zhang Y (2005) Leaf nitrogen and phosphorus stoichiometry across 753 terrestrial plant species in China. New Phytol 168:377–385
Hou H-Y (1982) Vegetation map of the People’s Republic of China (1:4M). Chinese Map Publisher, Beijing
Kay AD, Ashton IW, Gorokhova EC, Kerhoff AJ, Liess A, Litchman E (2005) Toward a stoichiometric framework for evolutionary biology. Oikos 109:6–17
Körner C (1989) The nutrient status of plants from high altitudes: a worldwide comparison. Oecologia 81:379–391
Körner C (1999) Alpine plant life: functional plant ecology of high mountain ecosystems. Springer, Berlin Heidelberg New York
Li WH, Zhou XM (eds) (1998) Ecosystems of Qinghai-Xizang (Tibetan) Plateau and approaches for their sustainable management. Guangdong Science and Technology Press, Guangzhou
McGroddy ME, Daufresne T, Hedin LO (2004) Scaling of C:N:P stoichiometry in forests worldwide: implications of terrestrial Redfield-type ratios. Ecology 85:2390–2401
Piao SL, Fang JY, Zhou LM, Guo QH, Henderson M, Ji W, Li Y, Tao S (2003) Interannual variations of monthly and seasonal normalized difference vegetation index (NDVI) in China from 1982 to 1999. J Geophys Res Atmos 108(D14):4401. DOI 4410.1029/2002JD002848
Redfield AC (1958) The biological control of chemical factors in the environment. Am Sci 46:205–221
Reich PB, Oleksyn J (2004) Global patterns of plant leaf N and P in relation to temperature and latitude. Proc Natl Acad Sci USA 101:11001–11006
Reich PB, Walters MB, Ellsworth DS (1997) From tropics to tundra: global convergence in plant functioning. Proc Natl Acad Sci USA 94:13730–13734
Reich PB, Wright IJ, Cavender-Bares J, Craine JM, Oleksyn J, Westoby M, Walters MB (2003) The evolution of plant functional variation: traits, spectra, and strategies. Int J Plant Sci 164:S143–S164
SAS (1999) SAS/STAT User’s guide, 8.01 edn. SAS Institute, Cary
Sterner RW, Elser JJ (2002) Ecological stoichiometry: the biology of elements from molecules to the biosphere. Princeton University Press, Princeton
Vitousek PM, Howarth RW (1991) Nitrogen limitation on land and in the sea: how can it occur? Biogeochemistry 13:87–115
Westoby M, Falster DS, Moles AT, Vesk PA, Wright IJ (2002) Plant ecological strategies: some leading dimensions of variation between species. Annu Rev Ecol Syst 33:125–159
Woods HA, Makino W, Cotner JB, Hobbie SE, Harrison JF, Acharya K, Elser JJ (2003) Temperature and the chemical composition of poikilothermic organisms. Funct Ecol 17:237–245
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, Hikosaka K, Lamont BB, Lee T, Lee WJ, Lusk C, Midgley JJ, Navas M-L, Niinemets U, Oleksyn J, Osada N, Poorter H, Poot P, Prior L, Pyankov VI, Roumet C, Thomas SC, Tjoelker MG, Veneklaas EJ, Villar R (2004) The worldwide leaf economics spectrum. Nature 428:821–827
Wright IJ, Reich PB, Cornelissen JHC, Falster DS, Garnier E, Hikosaka K, Lamont BB, Lee W, Oleksyn J, Osada N, Poorter H, Villar R, Warton DI, Westoby M (2005) Assessing the generality of global leaf trait relationships. New Phytol 166:485–496
Wu ZY (ed) (1980) Vegetation of China. Science Press, Beijing
Xu P (ed) (1993) Grassland resources and utilization of Xinjiang. Xinjiang Science, Technology and Public Health Publishing House, Urumqi
Zhang J, Wang JT, Chen W, Li B, Zhao K (1988) Vegetation of Xizang (Tibet). Science Press, Beijing
Acknowledgements
The authors thank members of Peking University Expedition Teams to the Tibetan Plateau (2003, 2004), Xinjiang Autonomous Region (2004) and Inner Mongolia (2004) for assistance with field data collection. Ch. Körner, B. Schmid, H. Heilmeier, Y.H. Tang, X.P. Wang and two anonymous reviewers helped with data interpretation and discussion. This research was supported by the National Natural Science Foundation of China (Grant 90411004, 40021101 and 90211016) to J.S.H. and J.Y.F., the State Key Basic Research and Development Plan (Project 2002CB412502) to J.S.H. and Peking University Research Fund (Project 211-II and 985-II) to J.Y.F. We declare that the work reported here complies with the current laws of the countries in which it was performed.
Author information
Authors and Affiliations
Corresponding author
Additional information
Communicated by Hermann Heilmeier
Rights and permissions
About this article
Cite this article
He, JS., Fang, J., Wang, Z. et al. Stoichiometry and large-scale patterns of leaf carbon and nitrogen in the grassland biomes of China. Oecologia 149, 115–122 (2006). https://doi.org/10.1007/s00442-006-0425-0
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s00442-006-0425-0