Vertical decoupling of soil nutrients and water under climate warming reduces plant cumulative nutrient uptake, water-use efficiency and productivity

Warming-induced desiccation of the fertile topsoil layer could lead to decreased nutrient diffusion, mobility, mineralization and uptake by roots. Increased vertical decoupling between nutrients in topsoil and water availability in subsoil/bedrock layers under warming could thereby reduce cumulative nutrient uptake over the growing season. We used a Mediterranean semiarid shrubland as model system to assess the impacts of warming-induced topsoil desiccation on plant water- and nutrient-use patterns. A 6 yr manipulative field experiment examined the effects of warming (2.5 degrees C), rainfall reduction (30%) and their combination on soil resource utilization by Helianthemum squamatum shrubs. A drier fertile topsoil ('growth pool') under warming led to greater proportional utilization of water from deeper, wetter, but less fertile subsoil/bedrock layers ('maintenance pool') by plants. This was linked to decreased cumulative nutrient uptake, increased nonstomatal (nutritional) limitation of photosynthesis and reduced water-use efficiency, above-ground biomass growth and drought survival. Whereas a shift to greater utilization of water stored in deep subsoil/bedrock may buffer the negative impact of warming-induced topsoil desiccation on transpiration, this plastic response cannot compensate for the associated reduction in cumulative nutrient uptake and carbon assimilation, which may compromise the capacity of plants to adjust to a warmer and drier climate.