The low-temperature limited alpine treeline is one of the most obvious boundaries in mountain landscapes. The question of whether resource limitation is the physiological mechanism for the formation of the alpine treeline is still waiting for conclusive evidence and answers. We therefore examined non-structural carbohydrates (NSC) and nitrogen (N) in treeline trees (TATs) and low-elevation trees (LETs) in both summer and winter in 11 alpine treeline cases ranging from subtropical monsoon to temperate continental climates across Eurasia. We found that tissue N concentration did not decrease with increasing elevation at the individual treeline level, but the mean root N concentration was lower in TATs than in LETs across treelines in summer. The TATs did not have lower tissue NSC concentrations than LETs in summer. However, the present study with multiple tree species across a large geographical scale, for the first time, revealed a common phenomenon that TATs had significantly lower NSC concentration in roots but not in the aboveground tissues than LETs in winter. Compared with LETs, TATs exhibited both a passive NSC storage in aboveground tissues in excess of carbon demand and an active starch storage in roots at the expense of growth reduction during the growing season. This starch accumulation disappeared in winter. Our results highlight some important aspects of the N and carbon physiology in relation to season in trees at their upper limits. Whether or to what extent the disadvantages of winter root NSC and summer root N level of TATs affect the growth of treeline trees and the alpine treeline formation needs to be further studied.

Soil stoichiometry of carbon (C), nitrogen (N), and phosphorus (P) are indicators for nutrient balance. Shrub encroachment into grasslands could change nutrient concentrations and stoichiometry in soils, but the general patterns remain unclear. With a meta-analysis of a global dataset covering 344 observations from 68 studies, we examined the responses of grassland soil C:N:P stoichiometry to shrub encroachment under various environmental conditions. Our results show that: 1) Shrub encroachment significantly increased the concentrations of soil C (+29 %), N (+25 %), P (+20 %), C:N (+5 %), C:P (+12 %), and N:P (+6 %). The magnitude of such effects varied with climate, soil texture, and soil layer. 2) Increases in SOC and TN concentrations mainly occurred in Mediterranean and very humid climate zones. Soil C:P and N:P decreased in semi-humid climate zone after shrub encroachment. 3) The increases in SOC and TN concentrations and in the C:N, C:P, and N:P ratios after shrub encroachment were greater in the topsoil than in deeper soil layers. 4) Both finest-textured soil (clay) and coarsest-textured soil (sand) are beneficial for increase of soil nutrient concentrations following shrub encroachment. 5) The magnitude of the change in soil C:N was negatively correlated with the duration of shrub encroachment, due to greater increases in soil TN than in SOC concentrations with longer durations of encroachment. Our results indicate that soil stoichiometric shifts in shrub-encroached grasslands are relatively sensitive to environmental factors, including soil texture, soil pH, and climate. These findings help us to better understand the effects of shrub encroachment on biogeochemical cycling, functioning, and services in grasslands across a broad range of spatio-temporal scales.

Plant-plant interaction plays a key role in regulating the composition and structure of communities and ecosystems. Studies of plant-plant interactions in forest ecosystems have mainly concentrated on growth effects of neighboring plants on target trees. Physiological effects of neighboring plants on target trees, in particular understorey effects on physiology of overstorey trees, have received less attention. It is still unclear what is the physiological mechanisms underlying positive growth effects of understorey removal, although understorey removal has been applied to improve the wood production for hundreds of years worldwide. Only 17.5% of published works dealt with understorey-overstorey interactions and only a few of those researched the understorey effects on the physiology of overstorey trees. Case studies indicated that overstorey Abies faxoniana trees grown with different understorey shrubs showed significantly different levels of tissue nitrogen and mobile carbohydrates. Removal experiment showed that nitrogen and mobile carbohydrates concentrations in Cunninghamia lanceolata trees grown in the absence of understorey shrubs differed significantly (pure stand > mixture) with those in trees grown in the presence of understorey shrubs, in particular during the dry season. This review highlighted that the neighboring woody plants affect Cand N-physiology in overstorey trees. These effects may be mainly resulted from underground competition for soil water rather than for other resources as the effects were more pronounced during the dry season. The present review suggests that positive effects of neighboring removal (e.g., understorey removal, thinning) on overstorey trees can be expected more rapidly and strongly in stressful area (e.g., low rainfall, nutrient-poor site) than in areas with optimal growth conditions. Hence, ecophysiology-based management strategies for dealing with neighboring plants in forest ecosystems should take into account: 1) site conditions, 2) timing, duration and frequency of management practices, and 3) species-specific properties and other aspects such as biodiversity conservation and soil erosion