Acclimation of mature spruce and beech to five years of repeated summer drought - The role of stomatal conductance and leaf area adjustment for water use.

Forests globally are experiencing severe droughts, leading to significant reductions in growth, crown dieback and even tree mortality. The ability of forest ecosystems to acclimate to prolonged and repeated droughts is critical for their survival with ongoing climate change. In a five-year throughfall exclusion experiment, we investigated the long-term physiological and morphological acclimation of mature Norway spruce (Picea abies [L.] KARST.) and European beech (Fagus sylvatica L.) to repeated summer drought at the leaf, shoot and whole tree level. Throughout the drought period, spruce reduced their total water use by 70 % to only 4-9 L per day and tree, while beech was less affected with about 30 % reduction of water use. During the first two summers, spruce achieved this by closing their stomata by up to 80 %. Additionally, from the second drought summer onwards, spruce produced shorter shoots and needles, resulting in a stepwise reduction of total leaf area of over 50 % by the end of the experiment. Surprisingly, no premature leaf loss was observed. This reduction in leaf area allowed a gradual increase in stomatal conductance. After the five-year drought experiment, water consumption per leaf area was the same as in the controls, while the total water consumption of spruce was still reduced. In contrast, beech showed no significant reduction in whole-tree leaf area, but nevertheless reduced water use by up to 50 % by stomatal closure. If the restriction of transpiration by stomatal closure is sufficient to ensure survival of Norway spruce during the first drought summers, then the slow but steady reduction in leaf area will ensure successful acclimation of water use, leading to reduced physiological drought stress and long-term survival. Neighboring beech appeared to benefit from the water-saving strategy of spruce by using the excess water.