Warming reduces both photosynthetic nutrient use efficiency and water use efficiency in Mediterranean shrubsWarming reduces nutrient use efficiency

The ratio between net photosynthetic rates and the foliar contents of essential plant macronutrients (N, P, K) is termed photosynthetic nutrient use efficiency (PNutUE). A universal trade-off exists whereby plants cannot maximize their PNutUE and their intrinsic water use efficiency (WUEi, carbon gain per unit water spent) simultaneously, because any increase in intercellular CO2 concentration (c(i)) through a greater stomatal opening would increase PNutE but also enhances transpiration and therefore decreases WUEi. Rising temperatures associated with climate change can result in large decreases in WUEi in semiarid shrubs through photosynthetic machinery impairment and enhanced stomatal conductance and transpiration, but we know remarkably little about the influence of warming and drought on PNutUE and its interplay with WUEi in dryland vegetation. Using a 6-year (2011-2017) manipulative field experiment, we examined the effects of warming (2.5?, W), rainfall reduction (30%, RR), and their combination (W+RR) on the photosynthetic use efficiency of three essential nutrients (PNUE, PPUE and PKUE) and on WUEi in three shrub species growing at two semiarid shrublands for the years 2015-2017. Across species, warming (W and W+RR) reduced PNUE by 42.9%, PPUE by 43.8% and PKUE by 41.5% on average relative to shrubs growing under ambient temperatures, whereas RR did not significantly affect their PNutUE. These drastic reductions in PNutUE with warming were mainly driven by non-stomatal and largely non-nutritional decreases in net photosynthetic rates, which were almost halved in warmed shrubs. The photosynthetic use efficiencies of N, P and K were inversely related to foliar d(13)C, a proxy for time integrated WUEi, in both ambient (control and RR) and warmed (W and W+RR) shrubs, but with significantly smaller slopes and intercepts for warmed shrubs. Thus, plants achieve a smaller gain in PNutUE for any given increase in stomatal conductance (and reduction in WUEi) under warmer climatic conditions. The strong negative impact of warming on PNutE, along with the warming-induced shift in the trade-off between PNutUE and WUEi, could be indicative of an increasing inability of native plants to cope with warmer conditions, with dire implications for dryland vegetation productivity and survival under climate change.