The seasonal patterns of xylem embolism and xylem transport properties in Quercus pubescens Willd. and Quercus ilex L. trees growing in a natural mixed coppice stand in conditions of severe water stress were investigated. Xylem embolism was evaluated in both dehydrating branches and in apical twigs during a whole year. Measurements of xylem water potential were conducted from predawn to sunset on selected sunny days. On the same days, diurnal courses of leaf conductance were monitored. Measurements of half‐hourly sap flow were made by the heat‐pulse technique throughout the summer. At the onset of summer, a sharp decrease in water potential was observed in both species. Full recovery of water potentials was observed for both species after the first major rainfall event in September. Both experienced serious embolism throughout the year, ranging between minima of c. 60% (expressed as percentage loss of hydraulic conductivity) after the rains in autumn and after bud burst in spring, and maxima of c. 80% during summer and after freezing‐thawing events during the winter season. A significant negative linear relationship was found between water potential and xylem embolism in branches dehydrating in air for Q. pubescens and Q. ilex. Q. pubescens had greater efficiency in hydraulic transport (higher specific conductivity and leaf specific conductivity) by the xylem than Q. ilex. In June, leaf conductance was high early in the morning and decreased gradually during the day. Midday depression of leaf conductance, as a result of high evaporative demand combined with water deficit, was observed in both species. In August, leaf conductance of both species was greatly reduced, as water potential dropped to extremely low values, and the stomata were almost completely closed during the afternoon. No hysteresis resulting from plant capacitance was observed in the relationship between shoot water potential and sap flow. Q. pubescens exhibited very high values of whole‐tree hydraulic resistance between July and September, whereas Q. ilex generally showed lower values. The effect of soil moisture depletion on the relationship between sap flow and shoot water potential appears as a lowering of water potential at zero flow. A significant decrease of whole‐tree hydraulic resistance in both species was observed with the onset of the autumn, preceding the partial recovery of twig hydraulic conductivity. The results demonstrate that both Q. pubescens and Q. ilex, although highly tolerant of severe water stress and tissue dehydration, operate at the limits of safety which are surpassed under severe droughts, and prolonged climatic stress might predispose these Quercus species to decline.
Variations in the water relations and stomatal response of Quercus ilex were analysed under field conditions by comparing trees at two locations in a Mediterranean environment during two consecutive summers (1993 and 1994). We used the heat-pulse velocity technique to estimate transpirational water use of trees during a 5 month period from June to November 1994. At the end of sap flow measurements, the trees were harvested, and the foliage and sapwood area measured. A distinct environmental gradient exists between the two sites with higher atmospheric CO 2 concentrations in the proximity of a natural CO 2 spring. Trees at the spring site have been growing for generations in elevated atmospheric CO 2 concentrations. At both sites, maximum leaf conductance was related to predawn shoot water potential. The effects of water deficits on water relations and whole-plant transpiration during the summer drought were severe. Leaf conductance and water potential recovered after major rainfall in September to predrought values. Sap flow, leaf conductance and predawn water potential decreased in parallel with increases in hydraulic resistance, reaching a minimum in mid-summer. These relationships are in agreement with the hypothesis of the stomatal control of transpiration to prevent desiccation damage but also to avoid 'runaway embolism'. Trees at the CO 2 spring underwent less reduction in hydraulic resistance for a given value of predawn water potential. The decrease in leaf conductance caused by elevated CO 2 was limited and tended to be less at high than at low atmospheric vapour pressure deficit. Mean (and diurnal) sap flux were consistently higher in the control site trees than in the CO 2 spring trees. The degree of reduction in water use between the two sites varied among the summer periods. The control site trees had consistently higher sap flow at corresponding values of either sapwood cross-sectional area or foliage area. Larger trees displayed smaller differences than smaller trees, between the control and the CO 2 spring trees. A strong association between foliage area and sapwood cross-sectional area was found in both the control and the CO 2 spring trees, the latter supporting a smaller foliage area at the corresponding sapwood stem cross-sectional area. The specific leaf area (SLA) of the foliage was not influenced by site. The results are discussed in terms of the effects of elevated CO 2 on plant water use at the organ and whole-tree scale.
Variations in water relations and stomatal response of Quercus pubescens Willd. were analyzed under Mediterranean field conditions during two consecutive summers (1993 and 1994) at two locations characterized by different atmospheric CO(2) concentrations because of the presence at one of them of a CO(2) spring. Trees at the CO(2) spring site have been growing for generations in elevated atmospheric CO(2) concentrations. The heat-pulse velocity technique was used to estimate water use of trees during a 5-month period from June to November 1994. At the end of the sap flow measurements, the trees were harvested and foliage and sapwood area measured. At both sites, maximum leaf conductance was related to predawn shoot water potential. Effects of summer drought on plant water relations, including whole-plant transpiration, were severe, but leaf conductance and water potential recovered to predrought values after major rainfall in September. Leaf conductance, predawn water potential, and sometimes sap flow, decreased in parallel with increases in hydraulic resistance, reaching a minimum in midsummer. Hydraulic resistance was higher in trees at the control site than in trees at the CO(2) spring site. The effect of elevated CO(2) concentration on leaf conductance was less at high leaf-to-air water vapor pressure difference than at low leaf-to-air water vapor pressure difference. Mean and diurnal sap fluxes were consistently higher in trees at the control site than in trees at the CO(2) spring site. During the summer period, plant water use varied between the two sites. Trees at the control site had consistently higher sap flow at corresponding values of sapwood cross-sectional area than trees at the CO(2) spring site. Because trees at the CO(2) spring site supported a smaller foliage area for a corresponding sapwood cross-sectional area than trees at the control site, the overall mean sap flux/foliage area ratio did not differ between sites. The results are discussed in terms of effects of elevated CO(2) concentration on plant water use at the organ and whole-tree scale.
Summary — The effect of elevated atmospheric carbon dioxide on water relations was examined on downy oak (Quercus pubescens) and holm oak (Q ilex)
We investigated how proximity to natural CO(2) springs affected the seasonal patterns of xylem embolism in Quercus ilex L., Quercus pubescens Willd., Fraxinus ornus L., Populus tremula L. and Arbutus unedo L., which differ in leaf phenology and wood anatomy. Xylem embolism was evaluated in both artificially dehydrated branches and in hydrated apical branches collected at monthly intervals during a 20-month sampling period. Initial specific hydraulic conductivity was also evaluated. We found species-dependent differences in xylem hydraulic properties in response to elevated CO(2) concentration. Populus tremula was the most embolized and A. unedo was the least embolized of the species examined. Effects of elevated CO(2) were significant in Q. pubescens, P. tremula and A. unedo, whereas the overall response to elevated CO(2) was less evident in F. ornus and Q. ilex. Specific hydraulic conductivity differed among species but not between sites, although the interaction between species and site was significant. Differences in xylem vulnerability between trees growing near to the CO(2) spring and those growing in control areas were small. Although differences in hydraulic properties in response to elevated CO(2) concentration were small, they may be of great importance in determining future community composition in Mediterranean-type forest ecosystems. The possible causes and ecological significance of such differences are discussed in relation to elevated CO(2) concentration and other environmental conditions.
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