Earthworm effects on native grassland root system dynamics under natural and increased rainfall

Earthworms (EWs) can modify soil structure and nutrient availability, and hence alter conditions for plant growth through their burrowing and casting activities. However, few studies have specifically quantified EW effects by experimentally manipulating earthworm densities (EWDs). In an earlier field study in native grassland ecosystems exposed to ambient and experimentally elevated rainfall (+280 mm year(-1), projected under some climate change scenarios), we found no effects of EWDs (37, 114, 169 EW m(-2)) and corresponding EW activity on aboveground net primary productivity (ANPP), even though soil nutrient availability likely increased with increasing EWDs. The lack of effects of EWDs on ANPP suggested that EWs may have adversely affected root systems in that study in some way. The objective of the present study was to quantify responses of root length density (RLD), using data collected from the same grassland plots during the earlier study. RLDs were highest in plots with low EWDs and decreased in plots with higher EWDs. Elevated rainfall primarily increased RLDs in the low EWD treatment (by almost +40%). Reductions in RLDs resulting from increased EWDs did not affect ANPP. Our results indicate that elevating EWDs above ambient levels may limit root growth through large increases in soil bioturbation, but concurrent increases in cast production and nutrient availability may compensate for the suppression of root nutrient absorbing surface area leaving ANPP unchanged, but with shifts in growth (biomass) allocation toward shoots. Similarly, reductions in EWDs appeared to promote higher RLDs that increased soil nutrient foraging in soil with lower amounts of nutrients because of reduced casting activity. Amplified responses observed when rainfall during the growing season was increased suggest that EWDs may mainly affect RLDs and above- vs. belowground growth (biomass) allocation under climate changes that include more frequent wetter-than-average growing seasons.