Aims Carbon and nutrient physiology of trees at their upper limits have been extensively studied, but those of shrubs at their upper limits have received much less attention. The aim of this study is to examine the general patterns of nonstructural carbohydrates (NSCs), nitrogen (N) and phosphorous (P) in shrubs at the upper limits, and to assess whether such patterns are similar to those in trees at the upper limits.
Methods Across Eurasia, we measured the concentrations of soluble sugars, starch, total NSCs, N and P in leaves, branches and fine roots (<0.5 cm in diameter) of five shrub species growing at both the upper limits and lower elevations in both summer (peak growing season) and winter (dormancy season).
Important Findings Neither elevation nor season had significant effects on tissue N and P concentrations, except for lower P concentrations in fine roots in winter than in summer. Total NSCs and soluble sugars in branches were significantly higher in winter than in summer. There were significant interactive effects between elevation and season for total NSCs, starch, soluble sugars and the ratio of soluble sugar to starch in fine roots, showing lower soluble sugars and starch in fine roots at the upper limits than at the lower elevations in winter but not in summer. These results suggest that the carbon physiology of roots in winter may play an important role in determining the upward distribution of shrubs, like that in the alpine tree-line trees.