Regulation of soil phosphorus cycling in grasslands by shrubs

The globally expanding colonization of grasslands by shrubs increases soil organic carbon and nitrogen, but the effects of shrubs on phosphorus (P) cycling have been rarely studied. We compared P contents in roots and soil fractions, phosphatase activity in the 1 m profile, and in situ net P mineralization between shrubby Potentilla fruticosa patches and grassy interspaces in grazed shrubby meadows at three representative sites on the eastern Tibetan Plateau. The P uptake of P. fruticosa exceeded 1 m soil depth, whereas grasses acquired P mainly within the upper 0.6 m. The P contents in shoots, aboveground litter and roots under P. fruticosa were greater than those under grasses. Litter P stock under the shrubs was 4-8 times higher than that under grasses and the root P stock doubled compared to that in grass areas. P. fruticosa generally increased the organic P (OP) content in the topsoil but decreased inorganic P (IP) in the subsoil. Phosphorus availability increased in the topsoil but decreased in the subsoil under the shrubs compared to grasses. Microbial biomass P (MBP), the activities of acid and alkaline phosphatases, and OP lability were all greater in the 1 m soil under P. fruticosa than grasses, leading to faster P mineralization and the P turnover under the shrubs. In the 1 m soil, P. fruticosa increased MBP and OP stocks but decreased IP and available P stocks. The larger and deeply distributed root system of P. fruticosa improved its P uptake ability especially from the subsoil. The subsequent greater organic matter input through litter fall and root turnover under P. fruticosa fed a larger microbial biomass that synthesized more microbial-derived OP in the topsoil. We concluded that shrubs increase the biological (plant and microbial) P transformation in the soil, the P uplift in the profile, and P cycling in shrubby grassland ecosystems. Such mechanisms structuring spatial heterogeneity of P content, transformation, turnover and fluxes are common in shrubby grasslands worldwide.