Competitive strategies for water availability in two Mediterranean Quercus species

Competition for water availability was studied in a mixed natural stand of Quercus suber L. and Quercus cerris L. growing in Sicily by measuring diurnal changes of leaf conductance to water vapour ( g _L), water potential ( Ψ _L) and relative water content ( RWC ) in April, July and October 1997 as well as the seasonal changes in root hydraulic conductance per unit leaf surface area ( K _RL). Quercus cerris behaved as a drought-tolerant species, with strong reductions of K _RL, Ψ _L, and RWC in the summer. By contrast, Q. suber appeared to withstand summer drought by an avoidance strategy based on reducing g _L, maintaining Ψ _L and RWC high and K _RL at the same level as that measured in the spring. A 'conductance ratio' ( CR ) was calculated in terms of the ratio of g _L to K _RL. Seasonal changes of this ratio contrasted in the two species, thus suggesting that Q. suber and Q. cerris did not really compete for available water. In the summer, when Q. suber was extracting water from the soil to maintain high leaf hydration, Q. cerris had restricted water absorption, thus suffering drought but tolerating its effects. The possibility that cohabitation of drought-tolerant with drought-avoiding species can be generalized is also discussed.     

Changes in leaf hydraulics and stomatal conductance following drought stress and irrigation in Ceratonia siliqua (Carob tree)

Changes in leaf hydraulic conductance (K) were measured using the vacuum chamber technique during dehydration and rehydration of potted plants of Ceratonia siliqua . K of whole, compound leaves as well as that of rachides and leaflets decreased by 20–30% at leaf water potentials (Ψ_L) of −1.5 and −2.0 MPa, i.e. at Ψ_L values commonly recorded in field-growing plants of the species. Higher K losses (up to 50%) were measured for leaves at Ψ_L of −2.5 and −3.0 MPa, i.e. near or beyond the leaf turgor loss point. Leaves of plants rehydrated while in the dark for 30 min, 90 min and 12 h recovered from K loss with characteristic times and to extents inversely proportional to the initial water stress applied. Leaf conductance to water vapour of plants dehydrated to decreasing Ψ_L and rehydrated at low transpiration was inversely related to loss of K, thus suggesting that leaf vein embolism and refilling (and related changes in leaf hydraulics) may play a significant role in the stomatal response.     

Effect of water availability on leaf water isotopic enrichment in beech seedlings shows limitations of current fractionation models

Current models of leaf water enrichment predict that the differences between isotopic enrichment of water at the site of evaporation (Δ_e) and mean lamina leaf water enrichment (Δ_L) depend on transpiration rates ( E ), modulated by the scaled effective length ( L ) of water isotope movement in the leaf. However, variations in leaf parameters in response to changing environmental conditions might cause changes in the water path and thus L . We measured the diel course of Δ_L for ¹⁸O and ²H in beech seedlings under well-watered and water-limited conditions. We applied evaporative enrichment models of increasing complexity to predict Δ_e and Δ_L, and estimated L from model fits. Water-limited plants showed moderate drought stress, with lower stomatal conductance, E and stem water potential than the control. Despite having double E , the divergence between Δ_e and Δ_L was lower in well-watered than in water-limited plants, and thus, L should have changed to counteract differences in E . Indeed, L was about threefold higher in water-limited plants, regardless of the models used. We conclude that L changes with plant water status far beyond the variations explained by water content and other measured variables, thus limiting the use of current evaporative models under changing environmental conditions.     

Physiological effects of long-term exposure to low and moderate concentrations of atmospheric NH3 on poplar leaves

Abstract. Poplar shoots ( Populus euramericana L.) obtained from cuttings were exposed for 6 or 8 weeks to NH₃ concentrations of 50 and 100 μgm⁻³ or filtered air in fumigation chambers. After this exposure the rates of NH₃ uptake, transpiration, CO₂ assimilation and respiration of leaves were measured using a leaf chamber. During the long-term exposure also modulated chlorophyll fluorescence measurements were carried out to obtain information about the photosynthetic performance of individual leaves. Both fluorescence and leaf chamber measurements showed a higher photosynthetic activity of leaves exposed to 100 μg NH₃ m⁻³. These leaves showed also a larger leaf conductance and a larger uptake rate of NH₃ than leaves exposed to 50 μg m⁻³ NH₃ or filtered air. The long-term NH₃ exposure did not induce an internal resistance against NH₃ transport in the leaf, nor did it affect the leaf cuticle. So, not only at a short time exposure, but also at a long-term exposure NH₃ uptake into leaves can be calculated from data on the boundary layer and stomatal resistance for H₂O and ambient NH₃-concentration. Furthermore, the NH₃ exposure had no effect on the relation between CO₂-assimilation and stomatal conductance, indicating that NH₃ in concentrations up to 100 μg m⁻³ has no direct effect on stomatal behaviour; for example, by affecting the guard or contiguous cells of the stomata.     

Effects of boundary layer conductance on substomatal pressures of carbon dioxide

Abstract. Gas exchange measurements were made on single leaves of three C₃ and one C₄ species at air speeds of 0.4 and 4.0 m s⁻¹ to determine if boundary layer conductance substantially affected the substomatal pressure of carbon dioxide. Boundary layer conductances to water vapour were 0.4 to 0.5 mol m⁻² s⁻¹ at the lower air speed, and 1.2 to 1.5 mol m⁻² s⁻¹ at the higher air speed. Substomatal carbon dioxide pressures were about 5 Pa lower at low boundary layer conductance in the C₃ species, and about 3 Pa lower in the C₄ species when measurements were made at high and moderate photosynthetic photon flux densities. No evidence of stomatal adjustment to altered boundary layer conductance was found. Photosynthetic rates at high photon flux densities were reduced by about 20% at the low air speed in the C₃ species. The commonly reported values of substomatal carbon dioxide pressure for C₃ and C₄ species were found to occur only when measurements were made at the higher air speed.     

Nitrogen/phosphorus leaf stoichiometry and the scaling of plant growth

We adopted previous N : P stoichiometric models for zooplankton relative growth to predict the relative growth rates of the leaves μ _L of vascular plants assuming that annual leaf growth in dry mass is dictated by how leaf nitrogen N _L is allocated to leaf proteins and how leaf phosphorus P _L is allocated to rRNA. This model is simplified provided that N _L scales as some power function of P _L across the leaves of different species. This approach successfully predicted the μ _L of 131 species of vascular plants based on the observation that, across these species, N _L scaled, on average, as the 3/4 power of P _L, i.e. N _L ∝  P     . When juxtaposed with prior allometric theory and observations, our findings suggest that a transformation in N : P stoichiometry occurs when the plant body undergoes a transition from primary to secondary growth.     

Phosphorylation and Palmitoylation of the Human D2L Dopamine Receptor in Sf9 Cells

Abstract: We have expressed and biochemically characterized the human D2_long (D2_L) dopamine receptor isoform using the baculovirus/Sf9 cell system. The expressed receptor bound ligands with a pharmacological profile similar to that reported for neuronal and cloned D2_L receptors expressed in mammalian cell lines. Dopamine binding to D2_L receptor was sensitive to guanine nucleotides, indicating receptor coupling to endogenous G proteins. A D2_L receptor-specific antibody identified two major protein species at ∼44 kDa and at ∼93 kDa in immunoblots, suggesting the presence of D2_L receptor monomers and dimers. Both species were purified by immunoprecipitation from digitonin-solubilized preparation of cells expressing D2_L receptor prelabeled with ³²P_i or [³H]-palmitate. These results constitute the first direct evidence for D2_L receptor phosphorylation and palmitoylation.     

Photosynthesis, nitrogen allocation and specific leaf area in invasive Eupatorium adenophorum and native Eupatorium japonicum grown at different irradiances

The mechanisms underlying biological invasions are still not well elucidated. In this study, ecophysiological traits of invasive Eupatorium adenophorum and native E.   japonicum were compared at 10 irradiances in field. I hypothesized that the invader may allocate a higher fraction of leaf nitrogen (N) to photosynthesis and have higher light-saturated photosynthetic rate ( P _max) and specific leaf area (SLA) than E.   japonicum . The invader had a significantly higher ability to acclimate to high irradiance than E.   japonicum , while it showed a similar shade-tolerant ability. The invader indeed allocated a higher fraction of leaf N to photosynthesis than E.   japonicum , which, with its high leaf N content ( N _A), resulted in a higher N content in photosynthesis ( N _P), contributing to its higher biochemical capacity for photosynthesis and P _max. However, the invader had a significantly lower SLA than E.   japonicum , contributing to its higher P _max but increasing its area-based leaf construction cost. The abilities to acclimate to a wider range of irradiance and to allocate a higher fraction of leaf N to photosynthesis, and the higher P _max, N _A, N _P and leaf area ratio may contribute to the invasion of the invader. High SLA is not always necessary for invasive species.     

Responses of apple leaf stomata to environmental factors

Abstract. Stomatal conductances ( g _s) were measured on the leaves of 3–4 year old Golden Delicious trees and of seedlings of two other cultivars. Measurements were made on container grown trees in the field with a diffusion porometer in 1975 and 1976, and in controlled conditions in a leaf chamber in the laboratory in 1976. Stomatal densities in the Golden Delicious leaves were assessed from scanning electron micrographs. Stomatal density on extension shoot leaves was higher than on other leaf types after June.
The response to irradiance shown by both the porometer and the leaf chamber results could be described by a rectangular hyperbola: where g _max is maximum conductance and β indicates the sensitivity of g_s to photon influx density ( Q _p). The values of β were in the range 60–90 μmol m⁻² s⁻¹.
There was no evidence that apple stomata are sensitive to temperature per se, but g _s was reduced by increasing leaf to air vapour pressure deficits ( D ). There was a linear relationship between g _s and D which was not attributable to feed-back to leaf water potential (ψ_L) as the latter did not affect g _s until a threshold of about −2.0 to −2.5 MPa was reached. Conductance generally declined with increasing ambient CO₂ concentration.     

Limitation of plant water use by rhizosphere and xylem conductance: results from a model

Hydraulic conductivity ( K ) in the soil and xylem declines as water potential ( Ψ ) declines. This results in a maximum rate of steady-state transpiration ( E _crit) and corresponding minimum leaf Ψ ( Ψ _crit) at which K has approached zero somewhere in the soil–leaf continuum. Exceeding these limits causes water transport to cease. A model determined whether the point of hydraulic failure (where K = 0) occurred in the rhizosphere or xylem components of the continuum. Below a threshold of root:leaf area ( A _R: A _L), the loss of rhizosphere K limited E _crit and Ψ _crit. Above the threshold, loss of xylem K from cavitation was limiting. The A _R: A _L threshold ranged from > 40 for coarse soils and/or cavitation-resistant xylem to < 0·20 in fine soils and/or cavitation-susceptible xylem. Comparison of model results with drought experiments in sunflower and water birch indicated that stomatal regulation of E reflected the species' hydraulic potential for extracting soil water, and that the more sensitive stomatal response of water birch to drought was necessary to avoid hydraulic failure. The results suggest that plants should be xylem-limited and near their A _R: A _L threshold. Corollary predictions are (1) within a soil type the A _R: A _L should increase with increasing cavitation resistance and drought tolerance, and (2) across soil types from fine to coarse the A _R: A _L should increase and maximum cavitation resistance should decrease.     

Leaf xylem embolism, detected acoustically and by cryo-SEM, corresponds to decreases in leaf hydraulic conductance in four evergreen species

Hydraulic conductance of leaves ( K _leaf) typically decreases with increasing water stress. However, the extent to which the decrease in K _leaf is due to xylem cavitation, conduit deformation or changes in the extra-xylary pathway is unclear. We measured K _leaf concurrently with ultrasonic acoustic emission (UAE) in dehydrating leaves of two vessel-bearing and two tracheid-bearing species to determine whether declining K _leaf was associated with an accumulation of cavitation events. In addition, images of leaf internal structure were captured using cryo-scanning electron microscopy, which allowed detection of empty versus full and also deformed conduits. Overall, K _leaf decreased as leaf water potentials ( Ψ _L) became more negative. Values of K _leaf corresponding to bulk leaf turgor loss points ranged from 13 to 45% of their maximum. Additionally, Ψ _L corresponding to a 50% loss in conductivity and 50% accumulated UAE ranged from −1.5 to −2.4 MPa and from −1.1 to −2.8 MPa, respectively, across species. Decreases in K _leaf were closely associated with accumulated UAE and the percentage of empty conduits. The mean amplitude of UAEs was tightly correlated with mean conduit diameter ( R ² = 0.94, P  = 0.018). These results suggest that water stress-induced decreases in K _leaf in these species are directly related to xylem embolism.     

Measurements of electrical leaf surface conductance reveal re-condensation of transpired water vapour on leaf surfaces

Electrical conductance ( λ ) was measured continuously and in vivo on leaf surfaces of Vicia faba and Aegopodium podagraria . λ increased with rise and decreased with fall in humidity, exhibiting a hysteresis during an applied humidity cycle [90–20–-90% relative humidity (r.h.)]. After treatment with NaNO₃ aerosols, a sudden increase in λ was observed at 73% r.h., which is close to the deliquescence point of the salt. Transpiration and electrical conductance of untreated leaves were measured simultaneously under conditions of constant r.h., while the photosynthetic photon flux density and CO₂ concentration of the air were varied to induce changes of stomatal aperture. At 35% r.h., changes of light and CO₂ level revealed a strong correlation between stomatal conductance ( g _S) and λ for Vicia faba leaves. This was also found at 90, 75, 60, 45 and 25% r.h. on the lower but not on the astomatous, upper surface of Aegopodium podagraria . The correlation between g _S and λ for stomata-bearing leaf surfaces indicates that an equilibrium exists between the ambient water vapour phase and the liquid water phase on and within the cuticle. This is modified by transpired water vapour influencing the air humidity inside the boundary layer. Our results imply re-condensation of transpired water vapour to salts on the leaf surface and its sorption to the cuticle.     

Low carbon dioxide concentrations can reverse stomatal closure during water stress

Leaf water potentials below threshold values result in reduced stomatal conductance (g_s). Stomatal closure at low leaf water potentials may serve to protect against cavitation of xylem. Possible control of g_s by leaf water potential or hydraulic conductance was tested by drying the rooting medium in four herbaceous annual species until g_s was reduced and then lowering the [CO₂] to determine whether g_s and transpiration rate could be increased and leaf water potential decreased and whether hydraulic conductance was reduced at the resulting lower leaf water potential. In all species, low [CO₂] could reverse the stomatal closure because of drying despite further reductions in leaf water potential, and the resulting lower leaf water potentials did not result in reductions in hydraulic conductance. The relative sensitivity of g_s to internal [CO₂] in the leaves of dry plants of each species averaged three to four times higher than in leaves of wet plants. Two species in which g_s was reputed to be insensitive to [CO₂] were examined to determine whether high leaf to air water vapor pressure differences (D) resulted in increased stomatal sensitivity to [CO₂]. In both species, stomatal sensitivity to [CO₂] was indeed negligible at low D, but increased with D, and low [CO₂] partly or fully reversed closure caused by high D. In no case did low leaf water potential or low hydraulic conductance during drying of the air or the rooting medium prevent low [CO₂] from increasing g_s and transpiration rate.     

Effects of tree height on branch hydraulics, leaf structure and gas exchange in California redwoods

We examined changes in branch hydraulic, leaf structure and gas exchange properties in coast redwood ( Sequoia sempervirens ) and giant sequoia ( Sequoiadendron giganteum ) trees of different sizes. Leaf-specific hydraulic conductivity ( k _L) increased with height in S. sempervirens but not in S. giganteum , while xylem cavitation resistance increased with height in both species. Despite hydraulic adjustments, leaf mass per unit area (LMA) and leaf carbon isotope ratios ( δ ¹³C) increased, and maximum mass-based stomatal conductance ( g _mass) and photosynthesis ( A _mass) decreased with height in both species. As a result, both A _mass and g _mass were negatively correlated with branch hydraulic properties in S. sempervirens and uncorrelated in S. giganteum . In addition, A _mass and g _mass were negatively correlated with LMA in both species, which we attributed to the effects of decreasing leaf internal CO₂ conductance ( g _i). Species-level differences in wood density, LMA and area-based gas exchange capacity constrained other structural and physiological properties, with S. sempervirens exhibiting increased branch water transport efficiency and S. giganteum exhibiting increased leaf-level water-use efficiency with increasing height. Our results reveal different adaptive strategies for the two redwoods that help them compensate for constraints associated with growing taller, and reflect contrasting environmental conditions each species faces in its native habitat.     

Elevated CO2 enhances stomatal responses to osmotic stress and abscisic acid in Arabidopsis thaliana

A , carbon assimilation rate
ABA, abscisic acid
Ci , intercellular space CO₂ concentration
g , leaf conductance
WUE, water use efficiency

Carbon dioxide and abscisic acid (ABA) are two major signals triggering stomatal closure. Their putative interaction in stomatal regulation was investigated in well-watered air-grown or double CO₂-grown Arabidopsis thaliana plants, using gas exchange and epidermal strip experiments. With plants grown in normal air, a doubling of the CO₂ concentration resulted in a rapid and transient drop in leaf conductance followed by recovery to the pre-treatment level after about two photoperiods. Despite the fact that plants placed in air or in double CO₂ for 2 d exhibited similar levels of leaf conductance, their stomatal responses to an osmotic stress (0·16–0·24 MPa) were different. The decrease in leaf conductance in response to the osmotic stress was strongly enhanced at elevated CO₂. Similarly, the drop in leaf conductance triggered by 1 μ M ABA applied at the root level was stronger at double CO₂. Identical experiments were performed with plants fully grown at double CO₂. Levels of leaf conductance and carbon assimilation rate measured at double CO₂ were similar for air-grown and elevated CO₂-grown plants. An enhanced response to ABA was still observed at high CO₂ in pre-conditioned plants. It is concluded that: (i) in the absence of stress, elevated CO₂ slightly affects leaf conductance in A. thaliana ; (ii) there is a strong interaction in stomatal responses to CO₂ and ABA which is not modified by growth at elevated CO₂.     

The influence of leaf thickness on the CO2 transfer conductance and leaf stable carbon isotope ratio for some evergreen tree species in Japanese warm-temperate forests

1. The influence of leaf thickness on internal conductance for CO₂ transfer from substomatal cavity to chloroplast stroma ( g _i) and carbon isotope ratio (δ¹³C) of leaf dry matter was investigated for some evergreen tree species from Japanese temperate forests. g _i was estimated based on the combined measurements of gas exchange and concurrent carbon isotope discrimination.
2. Leaves with thicker mesophyll tended to have larger leaf dry mass per area (LMA), larger surface area of mesophyll cells exposed to intercellular air spaces per unit leaf area ( S _mes) and smaller volume ratio of intercellular spaces to the whole mesophyll (mesophyll porosity).
3. g _i of these leaves was correlated positively to S _mes but negatively to mesophyll porosity. The variation in g _i among these species would be therefore primarily determined by variation of the conductance in liquid phase rather than that in gas phase.
4. δ¹³C was positively correlated to mesophyll thickness and leaf nitrogen content on an area basis. However, g _i values did not correlate to δ¹³C. These results suggest that difference in δ¹³C among the species was not caused by the variation in g _i, but mainly by the difference in long-term photosynthetic capacity.
5. Comparison of our results with those of previous studies showed that the correlation between leaf thickness and g _i differed depending on leaf functional types (evergreen, deciduous or annual). Differences in leaf properties among these functional types were discussed.     

Effect of boundary layer conductance on the response of stomata to humidity

Abstract. Leaf conductance responses to leaf to air water vapour partial pressure difference (VPD) have been measured at air speeds of 0.5 and 3.0 ms⁻¹ in single attached leaves of three species in order to test the hypothesis that leaf conductance response to VPD is controlled by evaporation from the outer surface of the epidermis, rather than by evaporation through stomata. Total conductance decreased linearly with increassing VPD at both air speeds, but was decreased 1.6 3.0 times as much as by a given incrase in VPD at high than at low air speed. depending on species. In all species the relationship between leaf conductance and the gradient for evaporation from the epidermis was the same at both values of boundary layer conductance, supporting the hypothesis that direct epidermal evaporation controls stomatal guard cell behaviour in responses of stomata to VPD in these species.     

Gas exchange studies in two Portuguese grapevine cultivars

Gas exchange characteristics of leaves of Vitis vinifera L. cvs Tinta Amarela and Periquita, two grapevine cultivars grown in distinct climatic regions of Portugal, were studied under natural and controlled conditions. Daily time courses of gas exchange were measured on both a hot, sunny day and a cooler, partly cloudy day. Responses of net photosynthesis to irradiance and internal partial pressure of CO₂, were also obtained. A strong correlation between net photosynthesis (P_N) and leaf conductance (g_s) was found during the diurnal time courses of gas exchange, as well as a relatively constant internal partial pressure of CO₂ (P_i), even under non-steady-state conditions. On the cloudless day, both P_N and g_s were lower in the afternoon than in the morning, despite similar conditions of leaf temperature, air to leaf water vapor deficit and irradiance. The response curves of net photosynthesis to internal CO₂ showed linearity up to p_i values of 50 Pa, possibly indicating a substantial excess of photosynthetic capacity. When measured at low partial pressures of O₂ (1 kPa), P_N became inhibited at high CO₂ levels. Inhibition of P_N at high CO₂ was absent under normal levels of O₂ (21 kPa). Significant differences in gas exchange characteristics were found between the two cultivars, with T. Amarela having higher rates under similar measurement conditions. In particular, the superior performance of T. Amarela at high temperatures may represent adaptation to the warmer conditions at its place of origin.     

Transport of gases into leaves

Abstract. Transport of gases between the intercellular spaces of plant leaves and the surrounding air is analysed in terms of multicomponent collision processes through an isothermal, porous septum. Interaction of diffusing species with each other and with the pore walls is described using a modified Stefan–Maxwell equation and an equation relating the pressure gradient to the sum of the diffusive fluxes, weighted by their appropriate Knudsen diffusivities. Viscous How arising from an excess pressure within the leaf is also considered.
Equations are derived which describe the flux densities of water vapour and CO₂ through the stomata. The analysis is general and is applicable to trace gases other than CO₂. A simple conductance is defined for water vapour to relate the flux and mol fraction difference across the stomata, viz. N_w =− g_w , δ x_w / x_a . A simple conductance cannot be defined for CO₂ because the flux of water vapour has a significant influence on the CO₂ gradient. The equation derived for the intercellular mol fraction of CO₂ is in terms of the fluxes of CO₂ and water vapour and represents a 'large-pore' ( d > μm) approximation which requires no information about stomalal geometry. Analogous equations are developed for transfer of gases through the leaf boundary layer. Sample calculations are presented to illustrate the effect of neglecting the interaction of water vapour and CO₂ on the calculated intercellular and surface concentrations of CO₂. Equations for computing water vapour and CO₂ flux densities from leaf chamber measurements are also presented.     

Salinity effects on water relations in Lycopersicon esculentum and its wild salt-tolerant relative species L. pennellii

Cultivated tomato Lycopersicon esculentum (L.) Mill. cv. P-73 and its wild salt tolerant relative L. pennellii (Correll) D'Arcy accession PE-47, were grown during spring-summer 1989 under unheated plastic greenhouse conditions. Plants were submitted to two different salt treatments using 0 and 140 mM NaCI irrigation water. In both tomato species, salinity caused a proportionally larger reduction in leaf area than in leaf weight and, in L. esculentum , a proportionally larger decrease in stem weight than in leaf weight. Daily variations in leaf water potential (Ψ₁) were fundamentally due to changes in the evaporative demand of the atmosphere. Reductions in Ψ₁ due to salinity were consistent only in L. esculentum . In all the conditions studied, leaf turgor was maintained. Leaf conductance (g₁)was higher in L. esculentum than in L. pennellii .Salinity induced a clear reduction in g₁ levels in L. esculentum whereas, in L. pennellii , this reduction was noted only in May. In both species the Ψ^o_s (leaf osmotic potential at full turgor) levels were reduced by salinity. The bulk modulus of elasticity (E) and relative water content at turgor loss point (RWC_tlp) were not affected by salinity. The RWC_tlp values in L. pennellii seem to be controlled by E values.