Forest transpiration estimates are frequently based on xylem sap flux measurements in the outer sections of the hydro-active stem sapwood. We used Granier's constant-heating technique with heating probes at various xylem depths to analyze radial patterns of sap flux density in the sapwood of seven broad-leaved tree species differing in wood density and xylem structure. Study aims were to (1) compare radial sap flux density profiles between diffuse- and ring-porous trees and (2) analyze the relationship between hydro-active sapwood area and stem diameter. In all investigated species except the diffuse-porous beech (Fagus sylvatica L.) and ring-porous ash (Fraxinus excelsior L.), sap flux density peaked at a depth of 1 to 4 cm beneath the cambium, revealing a hump-shaped curve with species-specific slopes. Beech and ash reached maximum sap flux densities immediately beneath the cambium in the youngest annual growth rings. Experiments with dyes showed that the hydro-active sapwood occupied 70 to 90% of the stem cross-sectional area in mature trees of diffuse-porous species, whereas it occupied only about 21% in ring-porous ash. Dendrochronological analyses indicated that vessels in the older sapwood may remain functional for 100 years or more in diffuse-porous species and for up to 27 years in ring-porous ash. We conclude that radial sap flux density patterns are largely dependent on tree species, which may introduce serious bias in sap-flux-derived forest transpiration estimates, if non-specific sap flux profiles are assumed.
The functional role of internal water storage is increasingly well understood in tropical trees and conifers, while temperate broad-leaved trees have only rarely been studied. We examined the magnitude and dynamics of the use of stem water reserves for transpiration in five coexisting temperate broad-leaved trees with largely different morphology and physiology (genera Fagus, Fraxinus, Tilia, Carpinus and Acer). We expected that differences in water storage patterns would mostly reflect species differences in wood anatomy (ring vs. diffuse-porous) and wood density. Sap flux density was recorded synchronously at five positions along the root-to-branch flow path of mature trees (roots, three stem positions and branches) with high temporal resolution (2 min) and related to stem radius changes recorded with electronic point dendrometers. The daily amount of stored stem water withdrawn for transpiration was estimated by comparing the integrated flow at stem base and stem top. The temporal coincidence of flows at different positions and apparent time lags were examined by cross-correlation analysis. Our results confirm that internal water stores play an important role in the four diffuse-porous species with estimated 5–12 kg day−1 being withdrawn on average in 25–28 m tall trees representing 10–22% of daily transpiration; in contrast, only 0.5–2.0 kg day−1 was withdrawn in ring-porous Fraxinus. Wood density had a large influence on storage; sapwood area (diffuse- vs. ring-porous) may be another influential factor but its effect was not significant. Across the five species, the length of the time lag in flow at stem top and stem base was positively related to the size of stem storage. The stem stores were mostly exhausted when the soil matrix potential dropped below −0.1 MPa and daily mean vapor pressure deficit exceeded 3–5 hPa. We conclude that stem storage is an important factor improving the water balance of diffuse-porous temperate broad-leaved trees in moist periods, while it may be of low relevance in dry periods and in ring-porous species.
Plants across diverse biomes tend to produce smaller leaves and a reduced total leaf area when exposed to drought. For mature trees of a single species, however, the leaf area–water supply relationship is not well understood. We tested the paradigm of leaf area reduction upon drought by a transect study with 14 mature Fagus sylvatica forests along a steep precipitation gradient (970–520 mm y−1) by applying two independent methods of leaf size determination. Contrary to expectation, average leaf size in dry stands (520–550 mm y−1) was about 40% larger and SLA was higher than in moist stands (910–970 mm y−1). As a result of increased leaf sizes, leaf area index significantly increased from the high- to the low-precipitation stands. Multiple regression analyses suggested that average leaf size was primarily controlled by temperature, whereas the influence of soil moisture and soil C/N ratio was low. Summer rainfall of the preceding year was the most significant predictor of total leaf number. We assume that leaf expansion of beech was independent of water supply, because it takes place in May with ample soil water reserves along the entire transect. In contrast, bud formation, which determines total leaf number, occurs in mid-summer, when droughts are severest. We conclude that leaf expansion and stand leaf area of beech along this precipitation gradient are not a simple function of water availability, but are controlled by several abiotic factors including spring temperature and possibly also nitrogen supply, which both tend to increase toward drier sites, thus overlaying any negative effect of water shortage on leaf development.