Drought response of European beech (Fagus sylvatica L.)-A review

European beech (Fagus sylvatica) is the dominant tree species of Central Europe's natural forests and one of the continent's most important timber species. This highly competitive species is known to be drought-sensitive and thus may increasingly be threatened by climate change-related heat waves and drought in part of its distribution range. Tree responses and tolerance to drought are complex processes that are best understood by adopting a multi-level analysis spanning from the molecular physiological to the tree and stand levels. Here I review recent progress in our understanding of beech drought responses in all relevant fields of research, notably adjustments in cellular biochemistry, responses of photosynthesis, stomatal regulation and leaf water status, the vulnerability of the hydraulic and phloem systems, adaptive responses of the fine root system, leaf area adjustments, long-term declines in radial growth, and drought-induced tree mortality. A special focus is on population differences in drought tolerance as the basis for the selection of drought-hardier provenances. The evidence from the different research fields is brought together with the aim to develop a multi-disciplinary perception of beech drought response, to identify weak components in the species' drought response strategy, and to suggest future research efforts to close knowledge gaps.