Regulation of soil nitrogen cycling by shrubs in grasslands

Globally, grasslands are experiencing shrub expansion, but the effects of such plant community shifts on soil N cycling are seldom investigated. We explored the spatial heterogeneity of total soil N content by grid sampling down to a soil depth of 1.2 m, and analyzed net ammonification and nitrification rates in the top 0.2 m, during the growing season, in response to the patchiness of grasses and four shrub species in a grazed meadow on the Tibetan Plateau. The delta N-15 analyses of plant, soil, and mineral N (NH4+ and NO3-) revealed differences in N absorption strategies between shrubs and grasses. Heterogeneity in the total soil N distribution was common to at least a depth of 1.2 m. The lower delta N-15 values of the 1.2 m soil profile under shrubs were consistent with the lower delta N-15 values of leaves, aboveground litter, and roots of shrubs compared to grasses, reflecting the imprint of long-term N cycling in the soil as affected by shrubs. Increases in total soil N stock in the entire 1.2 m profile by 28%-32% under non-leguminous shrubs relative to grasses were ascribed to N accumulation in the lower 0.8 m. Consequently, deep sampling is essential for accurate estimation of soil N storage in shrubby meadows. The total N stock in soil under the leguminous shrub species was similar to that under grasses, which showed that the effect of shrubification on soil N storage depends on the shrub species. Nitrification rates in the topsoil under all shrubs were greater than under grasses, producing higher N-15-depleted NO3- contents under shrubs. Therefore, the lower delta N-15 signatures of shrub tissues compared to grasses reflected a preferential or adaptive uptake of NO3- by shrubs. The expansion of shrubs in grasslands creates high heterogeneity in N cycling and storage down to a considerable soil depth.