Shoot water relations of mature black spruce families displaying a genotype x environment interaction in growth rate. II. Temporal trends and response to varying soil water conditions

Pressure-volume curves and shoot water potentials were determined for black spruce (Picea mariana (Mill.) BSP) trees from four full-sib families at the Petawawa Research Forest, Ontario, Canada. Trees were sampled from a dry site in 1992 and from the dry site and a wet site in 1993. Modulus of elasticity (epsilon), osmotic potential at turgor loss point (Psi(tlp)) and relative water at turgor loss point (RWC(tlp)) all decreased during the growing season. Osmotic potential at saturation (Psi(sat)) and turgor displayed no general temporal trend. Across a range of environmental conditions, Female 59 progeny had equal or lower Psi(sat), and higher or similar epsilon, mean turgor pressure (P(x)) and predawn turgor pressure (P(pd)) compared with Female 63 progeny. Osmotic potential at saturation decreased as water stress increased from mild to moderate and increased as water stress increased from moderate to severe. Stable genetic differences in Psi(sat) were maintained by the same rate of osmotic adjustment from low to moderate water stress. Modulus of elasticity and RWC(tlp) decreased with decreasing water availability, whereas Psi(tlp) showed no response. The combined effects of Psi(sat) and epsilon resulted in no change in P(pd) as water stress increased from low to moderate values, but turgor declined sharply as water stress increased from moderate to high values. We conclude that drought tolerance traits strongly influence the growth of these black spruce families across sites of varying water availability.     

Diurnal and seasonal changes in stem increment and water use by yellow poplar trees in response to environmental stress

To evaluate indicators of whole-tree physiological responses to climate stress, we determined seasonal, daily and diurnal patterns of growth and water use in 10 yellow poplar (Liriodendron tulipifera L.) trees in a stand recently released from competition. Precise measurements of stem increment and sap flow made with automated electronic dendrometers and thermal dissipation probes, respectively, indicated close temporal linkages between water use and patterns of stem shrinkage and swelling during daily cycles of water depletion and recharge of extensible outer-stem tissues. These cycles also determined net daily basal area increment. Multivariate regression models based on a 123-day data series showed that daily diameter increments were related negatively to vapor pressure deficit (VPD), but positively to precipitation and temperature. The same model form with slight changes in coefficients yielded coefficients of determination of about 0.62 (0.57-0.66) across data subsets that included widely variable growth rates and VPDs. Model R2 was improved to 0.75 by using 3-day running mean daily growth data. Rapid recovery of stem diameter growth following short-term, diurnal reductions in VPD indicated that water stored in extensible stem tissues was part of a fast recharge system that limited hydration changes in the cambial zone during periods of water stress. There were substantial differences in the seasonal dynamics of growth among individual trees, and analyses indicated that faster-growing trees were more positively affected by precipitation, solar irradiance and temperature and more negatively affected by high VPD than slower-growing trees. There were no negative effects of ozone on daily growth rates in a year of low ozone concentrations.     

Modeling tree water deficit from microclimate: an approach to quantifying drought stress

Tree water deficit estimated by measuring water-related changes in stem radius (DeltaW) was compared with tree water deficit estimated from the output of a simple, physiologically reasonable model (DeltaWE), with soil water potential (Psisoil) and atmospheric vapor pressure deficit (VPD) as inputs. Values of DeltaW were determined by monitoring stem radius changes with dendrometers and detrending the results for growth. We followed changes in DeltaW and DeltaWE in Pinus sylvestris L. and Quercus pubescens Willd. over 2 years at a dry site (2001-2002; Salgesch, Wallis) and in Picea abies (L.) Karst. for 1 year at a wet site (1998; Davos, Graubuenden) in the Swiss Alps. The seasonal courses of DeltaW in deciduous species and in conifers at the same site were similar and could be largely explained by variation in DeltaWE. This finding strongly suggests that DeltaW, despite the known species-specific differences in stomatal response to microclimate, is mainly explained by a combination of atmospheric and soil conditions. Consequently, we concluded that trees are unable to maintain any particular DeltaW. Either Psisoil or VPD alone provided poorer estimates of DeltaW than a model incorporating both factors. As a first approximation of DeltaWE, Psisoil can be weighted so that the negative mean Psisoil reaches 65 to 75% of the positive mean daytime VPD over a season (Q. pubescens: approximately 65%, P. abies: approximately 70%, P. sylvestris: approximately 75%). The differences in DeltaW among species can be partially explained by a different weighting of Psisoil against VPD. The DeltaW of P. sylvestris was more dependent on Psisoil than that of Q. pubescens, but less than that of P. abies, and was less dependent on VPD than that of P. abies and Q. pubescens. The model worked well for P. abies at the wet site and for Q. pubescens and P. sylvestris at the dry site, and may be useful for estimating water deficit in other tree species.     

Influence of irradiance on water relations and carbon flux during rooting of Shorea leprosula leafy stem cuttings

Single-node leafy stem cuttings of Shorea leprosula Miq. were subjected to a high, intermediate or low irradiance treatment for 16 weeks in an enclosed mist propagation system. Before rooting, maximum photosynthesis of the cuttings occurred at an irradiance of 400 micro mol m(-2) s(-1). Although none of the irradiance treatments affected the number of roots produced per cutting, the numbers of cuttings that formed roots were 50 and 30% in the high irradiance (diurnal range of 0-658 micro mol m(-2) s(-1)) and low irradiance (diurnal range of 0-98 micro mol m(-2) s(-1)) treatments, respectively, compared with 62% in the intermediate irradiance treatment (diurnal range of 0-360 micro mol m(-2) s(-1)). Low rooting frequency of cuttings in the high irradiance treatment was associated with water deficits (maximum leaf-to-air vapor pressure deficit (VPD) = 3.6 kPa), whereas cuttings in the low irradiance treatment had a low rooting frequency because they were below the light compensation point most of the time. In the intermediate irradiance treatment, cuttings withstood a daily maximum VPD of 1-2 kPa and recovered overnight from the previous day's deficit, as indicated by higher relative water content (RWC) and stomatal conductance (g(s)) in the morning than in the previous afternoon and evening. Higher RWC and g(s) of cuttings in all treatments on Days 14 and 21 compared with Day 8 probably indicated recovery from water deficit following severance and insertion of the cuttings in rooting medium. There were negative relationships between stem volume of cuttings and both number of cuttings that rooted and number of roots per cutting.     

Environmental control of ozone uptake in Larix decidua Mill.: a comparison between different altitudes

We studied ozone (O(3)) uptake by European larch (Larix decidua Mill.) needles under ambient conditions at both a low and a high elevation site. At a given ambient O(3) concentration, the rate of O(3) uptake by needles was effectively controlled by stomatal conductance and, hence, by factors such as light, humidity and water status, which control stomatal conductance. At both study sites, atmospheric water vapor pressure deficit (VPD) was the climatic factor most closely correlated with ambient O(3) concentration. Thus, when ambient O(3) concentrations were highest, O(3) flux into the needles tended to be restricted by narrowing of the stomata. Mitigation of potential O(3) stress by stomatal closure was most marked at the low elevation site where both soil water stress and atmospheric VPD were greater than at the high elevation site.     

Leaf gas exchange in the frankincense tree (Boswellia papyrifera) of African dry woodlands

A conceptual model was tested for explaining environmental and physiological effects on leaf gas exchange in the deciduous dry tropical woodland tree Boswellia papyrifera (Del.) Hochst. For this species we aimed at (i) understanding diurnal patterns in leaf gas exchange, (ii) exploring cause-effect relationships among external environment, internal physiology and leaf gas exchange, and (iii) exploring site differences in leaf gas exchange in response to environmental variables. Diurnal courses in gas exchange, underlying physiological traits and environmental variables were measured for 90 trees on consecutive days at two contrasting areas, one at high and the other at low altitude. Assimilation was highest in the morning and slightly decreased during the day. In contrast, transpiration increased from early morning to midday, mainly in response to an increasing vapor pressure deficit (VPD) and gradual stomatal closure. The leaf water potential varied relatively little and did not influence gas exchange during the measurement period. Our results suggest that the same cause-effect relationships function at contrasting areas. However, leaves at the higher altitude had higher photosynthetic capacity, reflecting acclimation to higher light levels. Trees at both areas nevertheless achieved similar leaf assimilation rates since assimilation was down-regulated by stomatal closure due to the higher VPD at the higher altitude, while it became more light limited at the lower altitude. Gas exchange was thus limited by a high VPD or low light levels during the wet season, despite the ability of the species to acclimate to different conditions.     

Variation in shoot water status of Picea abies (L.) Karst. trees with different life histories

Daily dynamics of the shoot water potential (ψ) in Norway spruce [Picea abies (L.) Karst.] depending on soil water availability and atmospheric evaporative demand was studied on originally freely-grown trees and suppressed trees exposed to full sunlight after clear-cutting. The base water potential (ψ_b) was significantly correlated (R² = 0.73) with the available soil water storage (W_tr) at a depth of 20–40 cm. ψ_b was relatively constant at high soil water status and decreased sharply if W_tr dropped below 14 mm. Among the atmospheric factors observed, the vapour pressure deficit (VPD) proved to be the most relevant variable in predicting diurnal changes in ψ. The leaf water status was more sensitive to VPD at sufficient soil water storage. With soil drying, the daily amplitude of ψ diminished, testifying to the more efficient stomatal control under soil drought. For predicting the daily course of ψ from VPD and ψ_b (taken as a measure of the soil water availability) an empirical model was derived, which described 78–80% of the total variatin in the shoot water potential. In contrast to values of ψ_b, the daily values of ψ differed significantly (P < 0.05) between the trees with different life histories. The newly exposed trees demonstrated substantially lower values of ψ with increasing atmospheric evaporative demand at the same soil water availability. Dynamic water stress is considered one of the main causes of the continuing growth retardation in suppressed spruce trees after their release from overstorey.     

Capacity of Stem Water Conductivity for Two Eucalyptus (Eucalyptus urophylla) Plantations in South China

The sap flux density (SFD) was used as a measure of water capacity through stems of eucalyptus in this paper. It was found that daily SFD increased with daily vapor pressure deficit (VPD) in nonlinear regression Y = AX³ + BX² + CX + D (R² = 0.702 1, n = 135, p = 0.01) at both sites (Hetou and Jijia) in the Leizhou Peninsula, Guangdong Province, China, where Y was daily SFD, X was mean daily VPD, A, B, C, D were constants. But extremely high VPD might limit stem water conductivity. The environmental factors, such as air vapor pressure deficit (VPD), solar radiation (RAD), etc., were the main determinants of SFD for E. urophylla plantations. The upper threshold of diurnal SFD was 51.55–55.65 mL&#8226;cm^–2&#8226;h^–1 under the selected extremely high environmental conditions.     

Growth CO2 concentration modifies the transpiration response of Populus deltoides to drought and vapor pressure deficit

Cottonwood (Populus deltoides Bartr. ex Marsh.) trees grown for 9 months in elevated carbon dioxide concentration ([CO2]) showed significant increases in height, leaf area and basal diameter relative to trees in a near-ambient [CO2] control treatment. Sample trees in the CO2 treatments were subjected to high and low atmospheric vapor pressure deficits (VPD) over a 5-week period at both high and low soil water contents (SWC). During these periods, transpiration rates at both the leaf and canopy levels were calculated based on sap flow measurements and leaf-to-sapwood area ratios. Leaf-level transpiration rates were approximately equivalent across [CO2] treatments when soil water was not limiting. In contrast, during drought stress, canopy-level transpiration rates were approximately equivalent across [CO2] treatments, indicating that leaf-level fluxes during drought stress were reduced in elevated [CO2] by a factor equal to the leaf area ratio of the two canopies. The shift from equivalent leaf-level transpiration to equivalent canopy-level transpiration with increasing drought stress suggests maximum water use rates were controlled primarily by atmospheric demand at high SWC and by soil water availability at low SWC. Changes in VPD had less effect on transpiration than changes in SWC for trees in both CO2 treatments. Transpiration rates of trees in both CO2 treatments reached maximum values at a VPD of about 2.0 kPa at high SWC, but leveled off and decreased slightly in both canopies as VPD increased above this value. At low SWC, increasing VPD from approximately 1.4 to 2.5 kPa caused transpiration rates to decline slightly in the canopies of trees in both treatments, with significant (P = 0.004) decreases occurring in trees in the near-ambient [CO2] treatment. The transpiration responses at high VPD in the presence of high SWC and throughout the low SWC treatment suggest some hydraulic limitations to water use occurred. Comparisons of midday leaf water potentials of trees in both CO2 treatments support this conclusion.     

Shoot water relations of mature black spruce families displaying a genotype x environment interaction in growth rate. I. Family and site effects over three growing seasons

Pressure-volume curves were determined for black spruce (Picea mariana (Mill.) BSP) trees from four full-sib families. During the first two years, trees were measured from a plantation on a dry site. In the third year, trees were sampled from the dry site and a wet site. Diurnal measurements of shoot water potential allowed in situ shoot turgor to be estimated in addition to standard water relations traits. Over all years, Female 59 progeny displayed lower osmotic potentials at saturation (Psi(sat)) than Female 63 progeny. Genetic differences in Psi(sat) were similar on both the dry and wet sites. Modulus of elasticity (epsilon) was greater for Female 59 progeny than for Female 63 progeny, producing a compensatory effect resulting in no genetic or site differences in osmotic potential at turgor loss point (Psi(tlp)) or relative water content at turgor loss point (RWC(tlp)). Mean and predawn shoot turgor pressures (P(x) and P(pd)) were higher for Female 59 progeny than for Female 63 progeny and higher at the wet site than the dry site. Genotype x environment trends were observed; compared to Female 63 progeny, Female 59 progeny displayed 9.8 and 5.1% higher P(pd) on the dry and wet sites, respectively, and 3.4 and 9.8% greater P(pd) values in wet and dry years, respectively. Tree volume growth showed no relationship to Psi(tlp) or RWC(tlp), but was correlated with Psi(sat) and P(x); however, the strongest correlation was with P(pd) (r = 0.90).     

Canopy conductance and stand transpiration of <Emphasis Type="Italic">Populus simonii Carr</Emphasis> in response to soil and atmospheric water deficits in farmland shelterbelt,Northwest China

Populus simonii Carr trees are the most abundant species in Xinjiang farmland shelterbelt. They played an important role in protecting farmland ecosystem. Their stand transpiration rates and canopy conductances ranged from 0.14 to 1.02 cm d^?1 and from 1.19*10^?3 to 8.99*10^?5 m s^?1, respectively, during the 2014 growing season. Transpiration rate was showed quadratic polynomial regression with vapor pressure deficit (VPD) and air temperature (T), and exponential relationship with solar radiation (R_n). Furthermore, transpiration peaked at VPD, T, and Rn were 2.1 kPa, 23 °C, and 200 W m^?2, respectively. Canopy conductance increased exponentially with the increasing R_n, whereas decreased logarithmically with the increasing VPD. Besides, transpiration increased with the increasing T and VPD ,and decreased sharply with the increasing soil water deficit. Finally, canopy conductance increased with the decreasing atmospheric water deficit. These results are not only providing the basis for more detailed analyses of water physiology and growth of Populus simonii Carr trees for the later application of a climate-driven process model, but also might have implications for farmland shelterbelt management.     

Water relations and gas exchange of Acer saccharum seedlings in contrasting natural light and water regimes

Field measurements were made of leaf photosynthesis (A), stomatal conductance (g) and leaf water relations for sugar maple (Acer saccharum Marsh.) seedlings growing in a forest understory, small gap or large clearing habitat in southwestern Wisconsin, USA. Predawn water status, leaf gas exchange and plasticity in field and laboratory water relations characteristics were compared among contrasting light environments in a wet year (1987) and a dry year (1988) to evaluate possible interactions between light and water availability in these habitats. Leaf water potentials (Psi(leaf)) at predawn and midday were lower for clearing than gap or understory seedlings. Acclimation of tissue osmotic potentials to light environment was observed among habitats but did not occur within any of the habitats in response to prolonged drought. During a summer drought in 1988, decreases in daily maximum g (g(max)) and maximum A (A(max)) in clearing seedlings were correlated with predawn Psi(leaf), which reached a seasonal minimum of -2.0 MPa. Under well-watered conditions, diurnal fluctuations in Psi(leaf) of up to 2.0 MPa in clearing seedlings occurred along with large midday depressions of A and g. In a wet year, strong stomatal responses to leaf-to-air vapor pressure difference (VPD) in sunny habitats were observed over nine diurnal courses of gas exchange measurements on seedlings in a gap and a clearing. Increasing stomatal limitations to photosynthesis appeared to be responsible for the reduction in A at high VPD for clearing seedlings. In understory seedlings, however, low water-use efficiency and development of leaf water deficits in sunflecks was related to reduced stomatal limitations to photosynthesis relative to seedlings in sunny habitats. Predawn Psi(leaf) and VPD appear to be important factors limiting carbon assimilation in sugar maple seedlings in light-saturating irradiances, primarily through stomatal closure. The overall results are consistent with the idea that sugar maple seedlings exhibit "conservative" water use patterns and have low drought tolerance. Leaf water relations and patterns of water use should be considered in studies of acclimation and species photosynthetic performance in contrasting light environments.     

Leaf water relations of Eucalyptus cloeziana and Eucalyptus argophloia in response to water deficit

Leaf water relations responses to limited water supply were determined in 7-month-old plants of a dry inland provenance of Eucalyptus argophloia Blakely and in a humid coastal provenance (Gympie) and a dry inland provenance (Hungry Hills) of Eucalyptus cloeziana F. Muell. Each provenance of E. cloeziana exhibited a lower relative water content at the turgor loss point, a lower apoplastic water content, a smaller ratio of dry mass to turgid mass and a lower bulk modulus of elasticity than the single provenance of E. argophloia. Osmotic potential at full turgor and water potential at the turgor loss point were significantly lower in E. argophloia and the inland provenance of E. cloeziana than in the coastal provenance of E. cloeziana. There was limited osmotic adjustment in response to soil drying in E. cloeziana, but not in E. argophloia. Between-species differences in water relations parameters were larger than those between the E. cloeziana provenances. Both E. cloeziana provenances maintained turgor under moderate water stress through a combination of osmotic and elastic adjustments. Eucalyptus argophloia had more rigid cell walls and reached lower water potentials with less reduction in relative water content than either of the E. cloeziana provenances, thereby enabling it to extract water from dryer soils.     

Transpiration sensitivity of urban trees in a semi-arid climate is constrained by xylem vulnerability to cavitation

Establishing quantitative links between plant hydraulic properties and the response of transpiration to environmental factors such as atmospheric vapor pressure deficit (D) is essential for improving our ability to understand plant water relations across a wide range of species and environmental conditions. We studied stomatal responses to D in irrigated trees in the urban landscape of Los Angeles, California. We found a strong linear relationship between the sensitivity of tree-level transpiration estimated from sap flux (m(T); slope of the relationship between tree transpiration and ln D) and transpiration at D=1 kPa (E(Tref)) that was similar to previous surveys of stomatal behavior in natural environments. In addition, m(T) was significantly related to vulnerability to cavitation of branches (P(50)). While m(T) did not appear to differ between ring- and diffuse-porous species, the relationship between m(T) and P(50) was distinct by wood anatomy. Therefore, our study confirms systematic differences in water relations in ring- versus diffuse-porous species, but these differences appear to be more strongly related to the relationship between stomatal sensitivity to D and vulnerability to cavitation rather than to stomatal sensitivity per se.     

Planting stress in newly planted jack pine and white spruce. 2. Changes in tissue water potential components

Water relations of bare-root jack pine (Pinus banksiana Lamb.) and white spruce (Picea glauca (Moench) Voss) planted in a greenhouse and on a boreal cut-over site were examined during the first growing season. In field-planted trees, maximum stomatal conductances (g(wv)) were initially low (< 0.10 cm s(-1)). Base and minimum xylem pressure potentials (Psi(x(base)) and Psi(x(min))) were less than -1.5 and -1.7 MPa for jack pine and -2.0 and -2.6 MPa for white spruce, respectively. During the growing season, maximum g(wv) increased in both species to around 0.2 cm s(-1). Base and minimum xylem pressure potentials also increased in both species to around -0.5 and -1.0 MPa in jack pine and -1.0 and -1.5 MPa in white spruce, respectively. Minimum xylem pressure potentials in white spruce fell below the turgor loss point during the first half of the growing season. Osmotic potential at the turgor loss point Psi(pi(TLP)) decreased after field planting to around -2.7 and -2.3 MPa in jack pine and white spruce, respectively. In the greenhouse, minimum values of Psi(pi(TLP)) were -2.2 and -2.3 MPa in jack pine and white spruce, respectively. Maximum bulk modulus of elasticity was greater in white spruce and underwent greater seasonal change than in jack pine. Relative water content (RWC) at turgor loss ranged between 71 and 74% in jack pine and 80 and 87% in white spruce. Available turgor (T(avail)), defined as the integral of turgor over the range of RWC between Psi(x(base)) and xylem pressure potential at the turgor loss point, was similar in jack pine and white spruce just after field planting. For the rest of the growing season, however, T(avail) in jack pine was two to three times that in white spruce. Diurnal turgor (T(diurnal)), defined as the integral of turgor over the range of RWC between Psi(x(base)) and Psi(x(min)), as a percent of T(avail) was higher in field-planted white spruce than jack pine until the end of the season. Dynamics of tissue water potential components are discussed in relation to plantation establishment.     

Vulnerability of low temperature induced needle retention in balsam fir (Abies balsamea L.) to vapor pressure deficits

Low temperature (LT) exposure has been shown to delay postharvest needle abscission and senescence (NAS) in balsam fir and changes in vapor pressure deficit (VPD) are expected to alter these processes. Two and half year-old seedlings were exposed to a LT of 5°C for 15 days while the control (CT) group was maintained at 22°C. Seedlings were then exposed to four different VPD (0.22, 0.87, 1.3 and 1.86?kPa) and observed for relative water content (RWC), xylem pressure potential (XPP), membrane injury index (MII), stem capacitance (SC) and NAS. An interactive effect of LT and VPD was observed in RWC, XPP, MII and NAS. Low VPD (0.22 and 0.87?kPa) resulted in less negative XPP, lower electrolyte leakage, higher SC and ultimately higher NAS than those at high VPD. Maximum NAS was recorded at 0.22?kPa. At 1.86?kPa, LT had 5× lower RWC (13%), 3× more negative XPP (?1.1?MPa), 1.8× higher membrane damage and 35% lower NAS (47 days) than CT. The SC declined with an increase in VPD with no effect of LT. The XPP and RWC of LT seedlings showed a positive relationship with NAS with R² values of 0.54 and 0.59, respectively. LT offered no benefits to NAS at high VPD environments.     

Differences in leaf gas exchange and water relations among species and tree sizes in an Arizona pine-oak forest

We compared leaf gas exchange and water potential among the dominant tree species and major size classes of trees in an upland, pine-oak forest in northern Arizona. The study included old-growth Gambel oak (Quercus gambelii Nutt.), and sapling, pole, and old-growth ponderosa pines (Pinus ponderosa var. scopulorum Dougl. ex Laws.). Old-growth oak had higher predawn leaf water potential (Psi(leaf)) than old-growth pine, indicating greater avoidance of soil water stress by oak. Old-growth oak had higher stomatal conductance (G(w)), net photosynthetic rate (P(n)), and leaf nitrogen concentration, and lower daytime Psi(leaf) than old-growth pine. Stomatal closure started at a daytime Psi(leaf) of about -1.9 MPa for pine, whereas old-growth oak showed no obvious reduction in G(w) at Psi(leaf) values greater than -2.5 MPa. In ponderosa pine, P(n) and G(w) were highly sensitive to seasonal and diurnal variations in vapor pressure deficit (VPD), with similar sensitivity for sapling, pole, and old-growth trees. In contrast, P(n) and G(w) were less sensitive to VPD in Gambel oak than in ponderosa pine, suggesting greater tolerance of oak to atmospheric water stress. Compared with sapling pine, old-growth pine had lower morning and afternoon P(n) and G(w), predawn Psi(leaf), daytime Psi(leaf), and soil-to-leaf hydraulic conductance (K(l)), and higher foliar nitrogen concentration. Pole pine values were intermediate between sapling and old-growth pine values for morning G(w) and daytime Psi(leaf), similar to sapling pine for predawn Psi(leaf), and similar to old-growth pine for morning and afternoon P(n), afternoon G(w), K(l), and foliar nitrogen concentration. For the pines, low predawn Psi(leaf), daytime Psi(leaf), and K(l) were associated with low P(n) and G(w). Our data suggest that hydraulic limitations are important in reducing P(n) in old-growth ponderosa pine in northern Arizona, and indicate greater avoidance of soil water stress and greater tolerance of atmospheric water stress by old-growth Gambel oak than by old-growth ponderosa pine.     

Diurnal variation in leaf extension of Salix viminalis at two nitrogen supply rates

To investigate how nitrogen supply might affect the biophysical factors controlling diurnal variation in leaf extension, pot-grown Salix viminalis L. were supplied with nitrogen at a low relative addition rate of 0.05 g N g(-1) N day(-1) (low N) or were given free access to all nutrients (high N). Leaf extension, turgor pressure, turgor after stress relaxation and the plastic extensibility of leaf tissue were determined for growing leaves every 4 h during two days of clear skies in August. Plants in the high-N treatment had a significantly higher relative growth rate, dry weight, shoot/root ratio, leaf nitrogen concentration, total leaf area, final area of single leaves and epidermal cell size than plants in the low-N treatment. The periodicity of leaf extension was similar in both treatments with high values during the afternoon and early evening, and negligible values during the night and in the early morning. The maximum rate of leaf extension was higher in high-N than in low-N plants. Leaf water potential and leaf osmotic potential decreased in the morning and increased in the afternoon with highest values during the night. Calculated values of turgor pressure showed no consistent diurnal trend and did not correlate with the rate of leaf extension. There was no consistent difference in turgor between treatments. Turgor after stress relaxation varied diurnally. The difference between turgors before and after stress relaxation also varied diurnally and was largely in phase with the diurnal pattern of leaf extension. These data are consistent with either a causal role for growth turgor (difference between turgors before and after stress relaxation) in the regulation of cell expansion, or a diurnal variation in turgors after relaxation, attributable to different capacities for cell wall loosening at different times of day. Plastic extensibility of leaf tissue showed no diurnal pattern but consistently higher values were found in high-N than in low-N plants. We conclude that the effects of nitrogen supply on leaf water relations did not limit leaf extension, but that nitrogen supply did affect processes associated with cell wall loosening and enlargement. Nitrogen supply did not affect final values of turgor after relaxation, but it presumably affected the rate at which relaxation proceeded.     

Response of stomatal conductance to drought in ponderosa pine: implications for carbon and ozone uptake

To gain insight into the limitations imposed by a typical Mediterranean-climate summer drought on the uptake of carbon and ozone in the ponderosa pine (Pinus ponderosa Dougl. ex Laws.) ecosystem, we compared diurnal trends in leaf physiology of young trees in a watered and a control plot located in the Sierra Nevada Mountains, CA, USA (Blodgett Forest, 38 degrees 53' N, 120 degrees 37' W, 1315 m elevation). Predawn water potential of trees in the watered plot remained above -0.3 MPa throughout the growing season, whereas it dropped in the control plot from -0.24 to -0.52 MPa between late May and mid-August. Photosynthesis and stomatal conductance of trees in the watered plot were relatively insensitive to atmospheric vapor pressure deficit (VPD), whereas gas exchange of trees in the control plot varied with changes in soil water, VPD and temperature. Although the 1998 growing season was abnormally wet, we saw a pronounced drought effect at the control site. Over the 2 months following the onset of watering, carbon and ozone uptake were measured on three days at widely spaced intervals. Carbon uptake per unit leaf area by 1-year-old foliage of trees in the control plot was 39, 35 and 30% less, respectively, than in the watered plot, and estimated ozone deposition per unit leaf area (ozone concentration times stomatal conductance) was 36, 46 and 41% less.     

Growth of European beech (Fagus sylvatica L.) saplings is limited by elevated atmospheric vapour pressure deficits

There is an ongoing debate about how European beech might be affected by a future drier climate. While numerous studies have examined the effects of soil drought on beech growth and development, studies investigating the effects of elevated atmospheric water vapour pressure deficit (VPD) are lacking. By increasing VPD in climate and open-top chamber experiments, with moisture in the rooting medium near optimum, we tested the hypothesis that increased VPD negatively affects the growth and development of European beech saplings. In the climate chambers, a reduction in relative air humidity by 40% resulted in a 68% reduction in productivity. Similarly, in the open-top chamber experiment conducted on the forest floor, biomass declined by 30% when relative air humidity was 15% lower. The reduction in biomass was mainly a consequence of a dramatically reduced leaf growth of beech in the elevated VPD treatments. Our results show that growth and development of beech saplings strongly depend not only on soil moisture but also on the prevailing VPD level. We conclude that the vapour pressure deficit is a widely ignored factor which influences the growth and vitality, and possibly also the distribution of European beech. Future forest management schemes under an altered climate should take this factor into account.