Uncertainty about the effects of warming and grazing on soil nitrogen (N) availability, species composition, and aboveground net primary production (ANPP) limits our ability to predict how global carbon sequestration will vary under future warming with grazing in alpine regions. Through a controlled asymmetrical warming (1.2/1.7 degrees C during daytime/nighttime) with a grazing experiment from 2006 to 2010 in an alpine meadow, we found that warming alone and moderate grazing did not significantly affect soil net N mineralization. Although plant species richness significantly decreased by 10% due to warming after 2008, we caution that this may be due to the transient occurrence or disappearance of some rare plant species in all treatments. Warming significantly increased graminoid cover, except in 2009, and legume cover after 2008, but reduced non-legume forb cover in the community. Grazing significantly decreased cover of graminoids and legumes before 2009 but increased forb cover in 2010. Warming significantly increased ANPP regardless of grazing, whereas grazing reduced the response of ANPP to warming. N addition did not affect ANPP in both warming and grazing treatments. Our findings suggest that soil N availability does not determine ANPP under simulated warming and that heavy grazing rather than warming causes degradation of the alpine meadows.

Understanding how flowering phenology responds to warming and cooling (i.e., symmetric or asymmetric response) is needed to predict the response of flowering phenology to future climate change that will happen with the occurrence of warm and cold years superimposed upon a long-term trend. A three-year reciprocal translocation experiment was performed along an elevation gradient from 3200 m to 3800 m in the Tibetan Plateau for six alpine plants. Transplanting to lower elevation (warming) advanced the first flowering date (FFD) and transplanting to higher elevation (cooling) had the opposite effect. The FFD of early spring flowering plants (ESF) was four times less sensitive to warming than to cooling (by À2.1 d/8C and 8.4 d/8C, respectively), while midsummer flowering plants (MSF) were about twice as sensitive to warming than to cooling (À8.0 d/8C and 4.9 d/8C, respectively). Compared with pooled warming and cooling data, warming alone significantly underpredicted 3.1 d/8C for ESF and overestimated 1.7 d/8C for MSF. These results suggest that future empirical and experimental studies should consider nonlinear temperature responses that can cause such warming-cooling asymmetries as well as differing life strategies (ESF vs. MSF) among plant species.

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Organisms' life cycles consist of hierarchical stages, from a single phenological stage (for example, flowering within a season), to vegetative and reproductive phases, to the total lifespan of the individual. Yet phenological events are typically studied in isolation, limiting our understanding of life history responses to climate change. Here, we reciprocally transfer plant communities along an elevation gradient to investigate plastic changes in the duration of sequential phenological events for six alpine species. We show that prolonged flowering leads to longer reproductive phases and activity periods when plants are moved to warmer locations. In contrast, shorter post-fruiting leaf and flowering stages led to shorter vegetative and reproductive phases, respectively, which resulted in shorter activity periods when plants were moved to cooler conditions. Therefore, phenological responses to warming and cooling do not simply mirror one another in the opposite direction, and low temperature may limit reproductive allocation in the alpine region.

To clarify the response of soil organic carbon (SOC) content to season-long grazing in the semiarid typical steppes of Inner Mongolia, we examined the aboveground biomass and SOC in both grazing (G-site) and no grazing (NG-site) sites in two typical steppes dominated by Leymus chinensis and Stipa grandis, as well as one seriously degraded L. chinensis grassland dominated by Artemisia frigida. The NG-sites had been fenced for 20 years in L. chinensis and S. grandis grasslands and for 10 years in A. frigida grassland. Aboveground biomass at G-sites was 21-35% of that at NGsites in L. chinensis and S. grandis grasslands. The SOC, however, showed no significant difference between G-site and NG-site in both grasslands. In the NG-sites, aboveground biomass was significantly lower in A. frigida grassland than in the other two grasslands. The SOC in A. frigida grassland was about 70% of that in L. chinensis grassland. In A. frigida grassland, aboveground biomass in the G-site was 68-82% of that in the NGsite, whereas SOC was significantly lower in the G-site than in the NG-site. Grazing elevated the surface soil pH in L. chinensis and A. frigida communities. A spatial heterogeneity in SOC and pH in the topsoil was not detected the G-site within the minimal sampling distance of 10 m. The results suggested that compensatory growth may account for the relative stability of SOC in G-sites in typical steppes. The SOC was sensitive to heavy grazing and difficult to recover after a significant decline caused by overgrazing in semiarid steppes.