Plant and Soil

, Volume 340, Issue 1–2, pp 337–345 | Cite as

Important foliar traits depend on species-grouping: analysis of a remnant temperate forest at the Keerqin Sandy Lands, China

  • Fu-Sheng Chen
  • Karl J. Niklas
  • De-Hui Zeng
Regular Article


Foliar traits are often interpreted to reflect strategies for coping with water and nutrient supply limitations. In this study, we measured several important leaf traits for 147 species sampled from a remnant, temperate deciduous broad-leaved forest in Keerqin Sandy Lands, Northeast China to test whether these traits are ‘invariant’ or dependent on water supply limitations. Our data show that average specific leaf area (SLA), nitrogen (N) and phosphorus (P) concentrations, leaf C/N, C/P and N/P were 273 cm2 g−1, 18.1 mg g−1, 1.60 mg g−1, 28.2, 343 and 12.4, respectively. However, most of these traits were significantly different (P < 0.05) for different species groupings based on growth forms, phylogenetic history, photosynthetic pathways, or habitats. SLA was positively correlated with leaf P concentration across the broad spectrum of 118 species and most species functional groupings. However, SLA was not correlated with N concentration across all species or within each species functional group. SLA and N and P concentrations in dry habitats were lower than those in wet habitats, whereas leaf C/N, C/P, and N/P had the opposite trend both across all species and within major species functional groupings (herb, monocots and C3 species). Our data indicate that SLA vs. leaf N and SLA vs. P relationships may be regulated differentially for different species functional groupings and that water limitation may have a greater influence than nutrient limitation for plant growth.


Diminishing return Leaf N/P ratio Sandy forest ecosystem Semi-arid region Species functional group Specific leaf area (SLA) 



This study was supported by grants from the National Key Technologies R&D Program of China (No. 2006BAD26B0201-1), the National Natural Science Foundation of China (Nos. 30872011 & 30960311) and National Key Basic Research Program of China (No. 2007CB106803). We thank Yi Gan, Hai-Jun Hu and Shi-Gen Wu for their field work; Lu Gan, Yi Liu and Qiu-Xiang Tian for nutrient analyses. We gratefully acknowledge two reviewers for their insightful comments on this manuscript. Dr. Hans Lambers provided suggestions on corrections of improper citing.

Supplementary material

11104_2010_606_MOESM1_ESM.doc (241 kb)
Appendix 1 Selected plant species and their characteristics in Daqinggou National Nature Reserve of Northeast China. (DOC 241 kb)


  1. Aerts R, Chapin FS III (2000) The mineral nutrition of wild plants revisited: a re-evaluation of processes and patterns. Adv Ecol Res 30:1–67CrossRefGoogle Scholar
  2. Allen SE (1989) Chemical analysis of ecological materials. Blackwell Scientific Publications, OxfordGoogle Scholar
  3. Chen FS, Zeng DH, He XY (2006) Small-scale spatial variability of soil nutrients and vegetation properties in a semi-arid site of northern China. Pedosphere 16:778–787CrossRefGoogle Scholar
  4. Chen FS, Zeng DH, Fahey TJ, Yao CY, Yu ZY (2010) Response of leaf anatomy of Chenopodium acuminatum to soil resource availability in a semi-arid grassland. Plant Ecol 209:375–382CrossRefGoogle Scholar
  5. Cunningham SA, Summerhayes B, Westoby M (1999) Evolutionary divergences in leaf structure and chemistry, comparing rainfall and soil nutrient gradients. Ecol Monogr 69:569–588CrossRefGoogle Scholar
  6. 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 408:578–580CrossRefPubMedGoogle Scholar
  7. Garnier E, Laurent G (1994) Leaf anatomy, specific leaf mass and water content in congeneric annual and perennial grass species. New Phytol 128:725–736CrossRefGoogle Scholar
  8. Garnier E, Shipley B, Roumet C, Laurent G (2001) A standardized protocol for the determination of specific leaf area and leaf dry matter content. Funct Ecol 15:688–695CrossRefGoogle Scholar
  9. Güsewell S (2004) N:P ratios in terrestrial plants: variation and functional significance. New Phytol 164:243–266CrossRefGoogle Scholar
  10. Han WX, Fang JX, Guo DL, Zhang Y (2005) Leaf nitrogen and phosphorus stoichiometry across 753 terrestrial plant species in China. New Phytol 168:377–385CrossRefPubMedGoogle Scholar
  11. Koerselman W, Meuleman AFM (1996) The vegetation N/P ratio – a new tool to detect the nature of nutrient limitation. J Appl Ecol 33:1441–1450CrossRefGoogle Scholar
  12. Lambers H, Poorter H (1992) Inherent variation in growth rate between higher plants: a search for physiological causes and ecological consequences. Adv Ecol Res 23:187–261CrossRefGoogle Scholar
  13. Lambers H, Brundrett MC, Raven JA, Hopper SD (2010) Plant mineral nutrition in ancient landscapes: high plant species diversity on infertile soils is linked to functional diversity for nutritional strategies. Plant Soil 334:11–31CrossRefGoogle Scholar
  14. Leigh RA, Storey R (1991) Differential distribution of nutrients between the epidermis and mesophyll of barley leaves: an X-ray microanalytical study. J Exp Bot 42(suppl):24–25Google Scholar
  15. McGroddy ME, Daufresne T, Hedin LO (2004) Scaling of C:N:P stoichiometry in forests worldwide: implications of terrestrial Redfield type ratios. Ecology 85:2390–2401CrossRefGoogle Scholar
  16. Niinemets Ü, Kull K (2003) Leaf structure vs. nutrient relationships vary with soil conditions in temperate shrubs and trees. Acta Oecol 24:209–219CrossRefGoogle Scholar
  17. Niinemets Ü, Ellsworth DS, Lukjanova A, Tobias M (2001) Site fertility and the morphological and photosynthetic acclimation of Pinus sylvestris needles to light. Tree Physiol 21:1231–1244PubMedGoogle Scholar
  18. Niklas KJ, Owens T, Reich PB, Cobb ED (2005) Nitrogen/phosphorus leaf stoichiometry and the scaling of plant growth. Ecol Lett 8:636–642CrossRefGoogle Scholar
  19. Niklas KJ, Cobb ED, Niinemets Ü, Reich PB, Sellin A, Shipley AB, Wright IJ (2007) “Diminishing returns” in the scaling of functional leaf traits across and within species groups. PNAS 104:8891–8896CrossRefPubMedGoogle Scholar
  20. Niklas KJ, Cobb ED, Spatz HC (2009) Predicting the allometry of leaf surface area and dry mass. Am J Bot 96:531–536CrossRefGoogle Scholar
  21. Peterson AG, Field CB, Ball JT, Amthor JS, Drake B, Emanuel WR, Johnson DW, Hanson PJ, Luo YQ, McMurtrie RE, Norby RJ, Oechel WC, Owensby CE, Parton WJ, Peterson AG, Pierce LL, Rastetter EB, Ruimy A, Running SW, Zak DR (1999) Reconciling the apparent difference between mass- and area-based expressions of the photosynthesis-nitrogen relationship. Oecologia 118:144–150CrossRefGoogle Scholar
  22. Poorter H, Bergkotte M (1992) Chemical composition of 24 wild species differing in relative growth rate. Plant Cell Environ 15:221–229CrossRefGoogle Scholar
  23. Poorter H, Niinemets U, Poorter L, Wright IJ, Villar R (2009) Causes and consequences of variation in leaf mass per area (LMA): a meta-analysis. New Phytol 183:565–588CrossRefGoogle Scholar
  24. Reich PB, Oleksyn J (2004) Global patterns of plant leaf N and P in relation to temperature and latitude. PNAS 101:11001–11006CrossRefPubMedGoogle Scholar
  25. Reich PB, Walters MB, Ellsworth DS (1992) Leaf life-span in relation to leaf, plant, and stand characteristics among diverse ecosystems. Ecol Monogr 62:365–392CrossRefGoogle Scholar
  26. Reich PB, Ellsworth DS, Walters MB, Vose JM, Gresham C, Volin JC, Bowman WD (1999) Generality of leaf trait relationships: a test across six biomes. Ecology 80:1955–1969CrossRefGoogle Scholar
  27. Sobrado MA, Medina E (1980) General morphology, anatomical structure, and nutrient content of sclerophyllous leaves of the ‘Bana’ vegetation of Amazonas. Oecologia 45:341–345CrossRefGoogle Scholar
  28. SPSS Inc. (2001) SPSS (Statistical Product and Service Solutions) for Windows (10.0). Chicago, ILGoogle Scholar
  29. Thompson WA, Huang LK, Kriedemann PE (1992) Photosynthetic response to light and nutrients in sun-tolerant and shade-tolerant rainforest trees. II. Leaf gas exchange and component processes of photosynthesis. Aust J Plant Physiol 19:19–42CrossRefGoogle Scholar
  30. Thompson K, Parkinson JA, Band SR, Spencer RE (1997) A comparative study of leaf nutrient concentrations in a regional herbaceous flora. New Phytol 136:679–689CrossRefGoogle Scholar
  31. Westoby M, Falster DS, Moles AT, Vesk PA, Wright IJ (2002) Plant ecological strategies: some leading dimensions of variation between species. Ann Rev Ecolog Syst 33:125–159CrossRefGoogle Scholar
  32. Wright IJ, Reich PB, Westoby M (2001) Strategy-shifts in leaf physiology, structure and nutrient content between species of high and low rainfall, and high and low nutrient habitats. Funct Ecol 15:423–434CrossRefGoogle Scholar
  33. 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 W, Lusk C, Midgley JJ, Navas M-L, 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 428:821–827CrossRefPubMedGoogle Scholar
  34. 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 D, Westoby M (2005) Assessing the generality of global leaf trait relationships. New Phytol 166:485–496CrossRefPubMedGoogle Scholar
  35. Zeng DH, Hu YL, Chang SX, Fan ZP (2009) Land cover change effects on soil chemical and biological properties after planting Mongolian pine (Pinus sylvestris var. mongolica) in sandy lands in Keerqin, northeastern China. Plant Soil 317:121–133CrossRefGoogle Scholar
  36. Zhang N-N, Guan W-B, Xie J, Yu M-T, Ye M-S, Su F-X (2007) Temporal and spatial distribution of soil moisture of Daqinggou Nantrue Reserve in the southeastern margin of Horqin sandy land, Inner Mongolia. China Acta Ecol Sin 27:3960–3876 (in Chinese with English abstract)Google Scholar
  37. Zheng YR (1998) The origin of plant community in Daqinggou. Sci Silvae Sin 34(6):22–28 (in Chinese with English abstract)Google Scholar
  38. Zheng YR (1999) Stability of Daqinggou forest communities. Acta Ecol Sin 19:579–581 (in Chinese with English abstract)Google Scholar

Copyright information

© Springer Science+Business Media B.V. 2010

Authors and Affiliations

  1. 1.Daqinggou Ecological Station, Institute of Applied EcologyChinese Academy of SciencesShenyangPeople’s Republic of China
  2. 2.College of Life SciencesNanchang UniversityNanchangPeople’s Republic of China
  3. 3.Department of Plant BiologyCornell UniversityIthacaUSA

Personalised recommendations