Internal water reserves are depleted and replenished daily, not only in succulent plants, but also in trees. The significance of these changes in tissue water storage for tree water relations was investigated by monitoring diurnal fluctuations in stem radius. In 6-year-old potted Norway spruce (Picea abies (L.) Karst.) trees, whole-tree transpiration rate (T), sap flow at the stem base and fluctuations in stem radius were measured at 10-min intervals over eight successive weeks. The dynamics of diurnal water storage in relation to the daily course of water movement was simulated and the contribution of stored water to T quantified. The finding that, in P. abies, the course of bark water content is linearly coupled to stem radius fluctuations provided the basis for linking stem radius changes to a functional flow and storage model for tree water relations. This model, which consists of physical functions only and is driven by a single input variable (T), accurately simulates the diurnal course of changes in stem radius and water storage of the tree crown and stem. It was concluded that fluctuations were mainly determined by the course of transpiration. The availability of soil water and the degree to which storage tissues were saturated were also factors affecting the diurnal course of stem radius changes. Internally stored water contributed to daily transpiration even in well-watered trees, indicating that stored water plays an important role not only during periods of drought, but whenever water transport occurs within the tree. Needle and bark water reserves were most heavily depleted during transpiration. Together they supplied approximately 10% of daily T on sunny days, and up to 65% on cloudy days. On a daily basis, the crown (mainly needles) contributed approximately eight times more water to T than the stem (mainly bark). The depletion of the two storage pools and the water movements observed in the trees always occurred in the same sequence. In the morning, T first caused a depletion of the water stored in the crown. It then caused depletion of bark storage tissues at ever increasing distances from the needles. Up to 75% of the transpired water could be withdrawn from storage tissues when the increase in T reached a maximum.

Changes in the stem radius of young Norway spruce [Picea abies (L.) Karst.] were related to changes in stem water content in order to investigate the relationship between diurnal stem size fluctuations and internally stored water. Experiments were performed on living trees and on cut stem segments. The defoliated stem segments were dried under room conditions and weight (W), volume (V), and xylem water potential (Ψ s ) were continuously monitored for 95 h. Additionally, photos of crosssections of fresh and air-dried stem segments were taken. For stem segments we found that the change in V was linearly correlated to the change in W as long as Ψ s was >-2.3±0.3 MPa (phase transition point). Stem contraction occurred almost solely in the elastic tissues of the bark (cambium, phloem, and parenchyma), and the stem radius changes were closely coupled to bark water content. For living trees, it is therefore possible to estimate the daily contribution of "bark water" to transpiration from knowledge of the stem size and continuous measurements of the stem radius fluctuations. When Ψ s reaches the phase-transition point, water is also withdrawn from the inelastic tissue of the stem (xylem), which -in the experiment with stem segments -was indicated by an increasing ratio between ∆V and ∆W. We assume that for Ψ s below the transition point, air is sucked into the tracheids (cavitation) and water is also withdrawn from the xylem. Due to the fact that in living P. abies Ψ s rarely falls below -2.3±0.3 MPa and the xylem size is almost unaffected by radius fluctuations, dendrometers are useful instruments with which to derive the diurnal changes in the bark water contents of Norway spruce trees.

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Internal water reserves in bark and foliage of trees contribute to transpiration (T) and play an essential role in optimizing water transport by buffering extreme peaks of water consumption. We examined patterns of stem shrinkage and their relationship to tree water dynamics. We measured fluctuations in root radius and stem radius at different stem heights, T of twigs at the top of the crown and sap flow velocities in stem sections of mature subalpine Norway spruce (Picea abies (L.) Karst.) trees over 2 years. The output of each sensor was coupled by physical functions to a mechanistic flow and storage model of tree water relations. The data verified the model-predicted lag in water storage depletion in response to the onset of transpiration and the lag increased with increasing distance from the crown periphery. Between the crown and stem base, the delay ranged from a few minutes to several hours, depending on microclimatic conditions and tree water status. Stem volume changes were proportional to the amount of water exchanged between the elastic tissues of the bark and the rigid xylem, indicating that the "peristaltic" wave of stem contraction along the flow path represented depletion of water stored in bark. On a daily basis, stems lost between 0.2 and 0.5% of their volume as a result of bark dehydration, corresponding to about 2 to 5 l of water. This water contributed directly to T. According to the model based on hydraulic principles, there are three main components underlying the dynamics of water storage depletion: flow resistance, storage capacities of needles and bark, and T of each tree section. The resistances and capacities were proportional to the response delay, whereas T in the lower parts of the tree was inversely proportional. The pattern of T within the crown depended on water intercepted by the branches. Because of these weather-dependent factors, there was no time constant for the response delay along the flow path. Nevertheless, the upper crown and the root section tended to have longer response delays per meter of flow path than the stem. The diurnal course of stem radius fluctuations represents the sum of all external and internal conditions affecting tree water relations; stem radius fluctuations, therefore, provide a sensitive measure of tree water status.

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