Drought-induced decline of productivity in the dominant grassland species Lolium perenne L. depends on soil type and prevailing climatic conditions

Severe constraints on grasslands productivity, ecosystem functions, goods and services are expected to result from projected warming and drought scenarios under climate change. Negative effects on vegetation can be mediated via soil fertility and water holding capacity, though specific mechanisms are fairly complex to generalise. In field drought experiments, it can be difficult to disentangle a drought effect per se from potential confounding effects related to vegetation or soil type, both varying along with climate. Furthermore, there is the need to distinguish the long-term responses of vegetation and soil to gradual climate shift from responses to extreme and stochastic climatic events. Here we address these limitations by means of a factorial experiment using a single dominant grassland species (the perennial ryegrass Lolium perenne L.) grown as a phytometer on two soils types with contrasted physicochemical characteristics, placed at two elevation sites along a climatic gradient, and exposed to early or late-season drought during the plant growing season. Warmer site conditions and reduced precipitation along the elevational gradient affected biogeochemistry and plant productivity more than the drought treatments alone, despite the similar magnitude in volumetric soil moisture reduction. Soil type, as defined here by its organic matter content (SOM), modulated the drought response in relation to local site climatic conditions and, through changes in microbial biomass and activity, determined the seasonal above and belowground productivity of L. perenne. More specifically, our combined uni-and multivariate analyses demonstrate that microbes in a loamy soil with low SOM are strongly responsive to change in climate, as indicated by a simultaneous increase in their C,N,P pools at high elevation with cooler temperatures and wetter soils. Contrastingly, microbes in a clay-loam soil with high SOM are mainly sensitive to temperature, as indicated by a strong increase in microbial biomass under warmer temperatures at low elevation and a concomitant increase in C:N, C:P and N:P ratios. High SOM promoted a better annual yield of the phytometer grass under warmer climate and the effect of drought on productivity was transient. In contrast, low SOM reduced cumulative yield under warmer conditions and root production strongly decreased, enduring a lasting drought effect. Microbes in soils with high organic matter remained more active during warmer and drier conditions, ensuring soil fertility and stimulating a higher overall plant nutrient availability and productivity. Our study highlights the important role of soil type for grassland responses to both stochastic climatic extremes and long-term climate change. Management practices enhancing SOM accumulation via organic residue incorporation seem a promising way to mitigate the effects of increased temperature and drought on plants and soil microbes alike promoting thereby a sustainable ecosystem functioning.