Two important threats to the sustainable functioning of seminatural grasslands in temperate zones are (1) nutrient loading due to agricultural fertilization and pollution, and (2) the increase of extreme drought events due to climate change. These threats may cause substantial shifts in species diversity and abundance and considerably affect the carbon and water balance of ecosystems. The synergistic effects between those two threats, however, can be complex and are poorly understood. Here, we experimentally investigated the effects of nitrogen addition and extreme drought (separately and in combination) on a seminatural temperate grassland, located in Freiburg (South Germany). To study the grassland response, we combined eddy-covariance techniques with open gas exchange systems. Open gas exchange chambers were connected to an infrared gas analyzer and water isotope spectrometer, which allowed the partitioning of net ecosystem exchange and evapotranspiration. Vegetation parameters were described by species richness, species abundance, and leaf area index. Our results suggest that grassland communities, strongly weakened in their stress response by nitrogen loading, can substantially lose their carbon sink function during drought. While nitrogen addition caused a significant loss in forb species (−25%), precipitation reduction promoted a strong dominance of grass species at season start. Consequently, the grass-dominated and species-poor community suffered from a strong above-ground dieback during the dry summer months, likely caused by lower water use efficiency and weaker drought adaptations of the species community. Over the growing season (April-September), the carbon sequestration of the studied grassland was reduced by more than 60% as a consequence of nitrogen addition. Nitrogen addition in combination with precipitation reduction decreased carbon sequestration by 73%. Eutrophication can severely threaten the resilient functioning of grasslands, in particular when drought periods will increase as predicted by future climate scenarios. Our findings emphasize the importance of preserving high diversity of grasslands to strengthen their resistance against extreme events such as droughts.
Nitrogen (N) loading and extreme drought strongly alter biomass production, species composition and carbon and water fluxes of temperate grasslands. Such changes at the community level are often attributed to species‐ and functional group‐specific responses in phenology and/or physiology. In a multifactorial field experiment, we studied the responses of three abundant grassland species (forb Centaurea jacea, grasses Arrhenatherum elatius and Dactylis glomerata) to N loading and extreme drought, focusing on responses of carbon and water relations at the leaf level. We analysed (1) changes in bulk leaf N (uptake efficiency of additional N), (2) adaptation of plant water status (leaf water potential) and (3) impact on leaf carbon and water fluxes. We observed more efficient N utilization in the two grasses compared to C. jacea. Naturally occurring summer drought significantly impacted the plant water status of all species, while extreme drought treatment only further affected water status during and after summer drought. C. jacea was able to maintain much lower leaf water potentials compared to the grasses during drought. Despite these clear species‐specific responses to N loading and drought, the species were able to maintain homeostasis of leaf carbon and water fluxes. Thus, strong declines in the (community) carbon sequestration observed at this site during the (natural) summer drought were not related to leaf physiological responses in assimilation, but were driven by phenological adaptions of the species community: the drought‐sensitive grasses, even though exhibiting higher N uptake efficiency, responded with a shortened life cycle to severe summer drought.
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