The Park Grass Experiment at Rothamsted in southeast England was started in 1856, making it the longest-running experiment in plant ecology anywhere in the world. Experimental inputs include a range of fertilizers (nitrogen, phosphorus, potassium, and organic manures) applied annually, with lime applied occasionally, and these have led to an increase in biomass and, where nitrogen was applied in the form of ammonium sulfate, to substantial decreases in soil pH. The number of species per plot varies from three to 44 per 200 m 2 , affording a unique opportunity to study the determinants of plant species richness and to estimate the effect sizes attributable to different factors. The response of species richness to biomass depends on the amount and type of nitrogen applied; richness declined monotonically with increasing biomass on plots receiving no nitrogen or receiving nitrogen in the form of sodium nitrate, but there was no relationship between species richness and biomass on plots acidified by ammonium sulfate application. The response to lime also depended on the type of nitrogen applied; there was no relationship between lime treatment and species richness, except in plots receiving nitrogen in the form of ammonium sulfate, where species richness increased sharply with increasing soil pH. The addition of phosphorus reduced species richness, and application of potassium along with phosphorus reduced species richness further, but the biggest negative effects were when nitrogen and phosphorus were applied together. The analysis demonstrates how multiple factors contribute to the observed diversity patterns and how environmental regulation of species pools can operate at the same spatial and temporal scale as biomass effects.
Evidence from field studies suggests that some plant species enhance their persistence by reinforcing patterns of N availability through differences in litter quality. Using mathematical models of nutrient flow, we explore whether and how recycling affects plant growth, competition, and coexistence and whether it leads to positive feedbacks. Two mechanisms are considered: the ability of plants to access two forms of soil N, complex (e.g., organic) and simple (e.g., nitrate), and the effect of density-dependent limitation of growth. Except in the trivial case of limitation by N in one form without density dependence, differences in litter quality can prevent the establishment of competitors. Feedback can, conversely, facilitate the invasion of competitors. At equilibrium, the rate of decomposition does not affect the outcome of competition. Species affect their long-term persistence if they alter the fraction of nitrogen that is returned to the soil and becomes available for plant uptake. Increasing the fraction of N that is recycled favors specialists in complex nitrogen and species that suppress the growth of others at high nitrogen availability. Increasing the rate of microbial decomposition of complex nitrogen favors specialists in simple nitrogen.
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