Xylem cavitation and hydraulic control of stomatal conductance in Laurel (Laurus nobilis L.)

The possible link between stomatal conductance (g_L), leaf water potential ( Ψ _L) and xylem cavitation was studied in leaves and shoots of detached branches as well as of whole plants of Laurus nobilis L. (Laurel). Shoot cavitation induced complete stomatal closure in air‐dehydrated detached branches in less than 10 min. By contrast, a fine regulation of g_L in whole plants was the consequence of Ψ _L reaching the cavitation threshold ( Ψ _CAV) for shoots. A pulse of xylem cavitation in the shoots was paralleled by a decrease in g_L of about 50%, while Ψ _L stabilized at values preventing further xylem cavitation. In these experiments, no root signals were likely to be sent to the leaves from the roots in response to soil dryness because branches were either detached or whole plants were growing in constantly wet soil. The stomatal response to increasing evaporative demand appeared therefore to be the result of hydraulic signals generated during shoot cavitation. A negative feedback link is proposed between g_L and Ψ _CAV rather than with Ψ _L itself.     

Epidermal conductance, stomatal density and stomatal size among genotypes of Sorghum bicolor (L.) Moench

Abstract. The ability of a plant to survive severe water deficits depends on its ability to restrict water loss through the leaf epidermis after stomata attain minimum aperture. At this stage, the rate of water loss is regulated by the epidermal conductance (g_c). Low g_c would be a useful selection criterion to identify genotypes with enhanced survival capability. Consequently, variation in g_c among Sorghum bicolor (L.) Moench genotypes was evaluated. Since there is little conclusive evidence linking g _c with leaf waxiness, alternative hypotheses relating g _c to stomatal trails were also examined. Epidermal conductance varied from 6.3 to 17.6mmol m⁻² s⁻¹ among sorghum genotypes. It was unrelated to stomatal pore length which varied with genotype and to pore depth which was similar for all genotypes measured. However, g _c, increased with increasing stomatal density. This indicates that stomatal density plays a direct role in water loss even at very low conductances. The association of low stomatal density with low g _c is consistent with the hypothesis that at the smallest stomata aperture, water loss from the epidermis above guard cell teichodes becomes a significant source of leaf water loss. Since low g _c is directly related to crop survival under severe water deficits, it is recommended that genotypes with low g _c. be selected using the selection criterion of stomatal density.     

Limitation of transpiration by hydraulic conductance and xylem cavitation in Betula occidentalis

The extent to which stomatal conductance (g_s) was capable of responding to reduced hydraulic conductance (k)and preventing cavitation-inducing xylem pressures was evaluated in the small riparian tree, Betula occidentalis Hook. We decreased k by inducing xylem cavitation in shoots using an air-injection technique. From 1 to 18 d after shoot injection we measured midday transpiration rate (E), g_s, and xylem pressure (Ψ_p-xylem) on individual leaves of the crown. We then harvested the shoot and made direct measurements of k from the trunk (2–3 cm diameter) to the distal tip of the petioles of the same leaves measured for E and g_s. The k measurement was expressed per unit leaf area (k_l, leaf-specific conductance). Leaves measured within 2 d of shoot injection showed reduced g_s and E relative to non-injected controls, and both parameters were strongly correlated with k_l At this time, there was no difference in leaf Ψ_p-xylem between injected shoots and controls, and leaf Ψ_p-xylem was not significantly different from the highest cavitation-inducing pressure (Ψ_p-cav) in the branch xylem (-1.43 ± 0.029 MPa, n=8). Leaves measured 7–18 d after shoots were injected exhibited a partial return of g_s and E values to the control range. This was associated with a decrease in leaf Ψ_p-xylem below Ψ_p-cav and loss of foliage. The results suggest the stomata were incapable of long-term regulation of E below control values and that reversion to higher E caused dieback via cavitation.     

Integration of hydraulic and chemical signalling in the control of stomatal conductance and water status of droughted plants

We describe here an integration of hydraulic and chemical signals which control stomatal conductance of plants in drying soil, and suggest that such a system is more likely than control based on chemical signals or water relations alone. The determination of xylem [ABA] and the stomatal response to xylem [ABA] are likely to involve the water flux through the plant. (1) If, as seems likely, the production of a chemical message depends on the root water status (Ψ_r), it will not depend solely on the soil water potential (Ψ_s) but also on the flux of water through the soil-plant-atmosphere continuum, to which are linked the difference between Ψ_r and Ψ_s. (2) The water flux will also dilute the concentration of the message in the xylem sap. (3) The stomatal sensitivity to the message is increased as leaf water potential falls. Stomatal conductance, which controls the water flux, therefore would be controlled by a water-flux-dependent message, with a water-flux-dependent sensitivity. In such a system, we have to consider a common regulation for stomatal conductance, leaf and root water potentials, water flux and concentration of ABA in the xylem. In order to test this possibility, we have combined equations which describe the generation and effects of chemical signals and classical equations of water flux. When the simulation was run for a variety of conditions, the solution suggested that such common regulation can operate. Simulations suggest that, as well as providing control of stomatal conductance, integration of chemical and hydraulic signalling may also provide a control of leaf water potential and of xylem [ABA], features which are apparent from our experimental data. We conclude that the root message would provide the plant with a means to sense the conditions of water extraction (soil water status and resisance to water flux) on a daily timescale, while the short-term plant response to this message would depend on the evaporative demand.     

Roles of leaf water potential and soil-to-leaf hydraulic conductance in water use by understorey woody plants

Diurnal changes of leaf water potential and stomatal conductance were measured for 12 deciduous shrubs and tree saplings in the understorey of a temperate forest. Sunflecks raised the leaf temperature by 4°C, and vapor pressure deficit to 2 kPa. Although the duration of the sunflecks was only 17% of daytime, the photon flux density (PFD) of sunflecks was 52% of total PFD on a sunny summer day. Leaf osmotic potential at full turgor decreased in summer, except in some species that have low osmotic potential in the spring. Plants that endured low leaf water potential had rigid cell walls and low osmotic potential at full turgor. These plants did not have lower relative water content and turgor potential than plants with higher leaf water potential. There were three different responses to an increase in transpiration rate: (i) plants had low leaf water potential and slightly increased soil-to-leaf hydraulic conductance; (ii) plants decreased leaf water potential and increased the hydraulic conductance; and (iii) plants had high leaf water potential and largely increased the hydraulic conductance.     

Water movement through Quercus rubra I. Leaf water potential and conductance during polycyclic growth

Two experiments examined simultaneous changes in leaf area (A_L), root length (L_r), stomatal conductance (g_s), leaf water potential (Ψ_L), transpiration and hydraulic plant conductance per unit leaf area (G) during the first three shoot cycles of northern red oak (Quercus rubra L.) grown under favourable and controlled conditions. Each shoot cycle consisted of bud swell, stem elongation, leaf expansion and rest; roots grew almost continuously. The g_s of all leaves decreased substantially while leaves of the newest flush were expanding and increased modestly when seedling leaf area remained constant. Overall, g_s decreased. The Ψ_L of mature leaves decreased during leaf expansion and increased by an equivalent amount during intervening periods. Possible explanations for the paired changes in g_s and Ψ_L are considered. Changes in G closely paralleled those of canopy g_s. These parallel changes during polycyclic seedling growth should act to keep seedling Ψ_L relatively constant as plant size increases and thereby help prevent Ψ_L from dropping to levels that would cause runaway embolism.     

A coupled model of stomatal conductance, photosynthesis and transpiration

A model that couples stomatal conductance, photosynthesis, leaf energy balance and transport of water through the soil–plant–atmosphere continuum is presented. Stomatal conductance in the model depends on light, temperature and intercellular CO₂ concentration via photosynthesis and on leaf water potential, which in turn is a function of soil water potential, the rate of water flow through the soil and plant, and on xylem hydraulic resistance. Water transport from soil to roots is simulated through solution of Richards’ equation. The model captures the observed hysteresis in diurnal variations in stomatal conductance, assimilation rate and transpiration for plant canopies. Hysteresis arises because atmospheric demand for water from the leaves typically peaks in mid‐afternoon and because of uneven distribution of soil matric potentials with distance from the roots. Potentials at the root surfaces are lower than in the bulk soil, and once soil water supply starts to limit transpiration, root potentials are substantially less negative in the morning than in the afternoon. This leads to higher stomatal conductances, CO₂ assimilation and transpiration in the morning compared to later in the day. Stomatal conductance is sensitive to soil and plant hydraulic properties and to root length density only after approximately 10 d of soil drying, when supply of water by the soil to the roots becomes limiting. High atmospheric demand causes transpiration rates, LE, to decline at a slightly higher soil water content, θ_s, than at low atmospheric demand, but all curves of LE versus θ_s fall on the same line when soil water supply limits transpiration. Stomatal conductance cannot be modelled in isolation, but must be fully coupled with models of photosynthesis/respiration and the transport of water from soil, through roots, stems and leaves to the atmosphere.     

A coupled model of photosynthesis,stomatal conductance and transpiration for a rose leaf (Rosa hybrida L.)

The following three models were combined to predict simultaneously photosynthesis, stomatal conductance, transpiration and leaf temperature of a rose leaf: the biochemical model of photosynthesis of Farquhar, von Caemmerer and Berry (1980, Planta 149: 78-90), the stomatal conductance model of Ball, Woodrow and Berry (In: Biggens J, ed. Progress in photosynthesis research. The Netherlands: Martinus Nijhoff Publishers), and an energy balance model. The photosynthetic parameters: maximum carboxylation rate, potential rate of electron transport and rate of triose phosphate utilization, and their temperature dependence were determined using gas exchange data of fully expanded, young, sunlit leaves. The stomatal conductance model was calibrated independently. Prediction of net photosynthesis by the coupled model agreed well with the validation data, but the model tended to underestimate rates of stomatal conductance and transpiration. The coupled model developed in this study can be used to assist growers making environmental control decisions in glasshouse production.     

Sensitivity of infrared water vapor analyzers to oxygen concentration and errors in stomatal conductance

Use of infrared analyzers to measure water vapor concentrations in photosynthesis systems is becoming common. It is known that sensitivity of infrared carbon dioxide and water vapor analyzers is affected by the oxygen concentration in the background gas, particularly for absolute analyzers, but the potential for large errors in estimates of stomatal conductance due to effects of oxygen concentration on the sensitivity of infrared water vapor analyzers is not widely recognized. This work tested three types of infrared water vapor analyzers for changes in sensitivity of infrared water vapor analyzers depending on the oxygen content of the background gas. It was found that changing from either 0 or 2% to 21% oxygen in nitrogen decreased the sensitivity to water vapor for all three types of infrared water vapor analyzers by about 4%. The change in sensitivity was linear with oxygen mole fraction. The resulting error in calculated stomatal conductance would depend strongly on the leaf to air vapor pressure difference and leaf temperature, and also on whether leaf temperature was directly measured or calculated from energy balance. Examples of measurements of gas exchange on soybean leaves under glasshouse conditions indicated that changing from 21% to 2% oxygen produced an artifactual apparent increase in stomatal conductance which averaged about 30%. Similar errors occurred for `conductances' of wet filter paper. Such errors could affect inferences about the carbon dioxide dependence of the sensitivity of photosynthesis to oxygen. This revised version was published online in June 2006 with corrections to the Cover Date.     

ABA xylem concentrations determine maximum daily leaf conductance of field-grown Vitis vinifera L. plants

Differences in maximum leaf conductance in grapevine plants growing in soils with contrasting water availabilities during mid-summer in Portugal could be accounted for by differences in the concentration of ABA in xylem sap. This conclusion is reinforced by the observation that the relationship between leaf conductance and endogenous ABA concentration can be mimicked by the application of exogenous ABA to leaves detached from irrigated plants. During the day, leaf conductance decreased after a morning peak, even when the leaves remained in a constant environment at a moderate temperature and leaf-to-air vapour pressure difference. This decline in leaf conductance was not a consequence of an increase in the xylem ABA concentration or the rate of delivery of this compound by the transpiratory stream. The afternoon depression in leaf conductance was associated with an apparent limitation in stomatal opening potential, which persisted even when detached leaves were fed with water and rehydrated. The reason for this inhibition has still to be identified.     

Maize stomatal conductance in the field: its relationship with soil and plant water potentials, mechanical constraints and ABA concentration in the xylem sap

Abstract. Stomatal conductance, leaf water potential, soil water potential and concentration of abscisic acid (ABA) in the xylem sap were measured on maize plants growing in the field, in two treatments with contrasting soil structures. Soil compaction affected the stomatal conductance, but this effect was no longer observed if the soil water potential was increased by irrigation. Differences in leaf water potential did not account for the differences in conductance between treatments. Conversely, the relationship between stomatal conductance and concentration of ABA in the xylem sap was consistent during the experiment. The proposed interpretation is that stomatal conductance was controlled by the root water potential via an ABA message. Control of the stomatal conductance by the leaf water potential or by an effect of mechanical stress on the roots is unlikely.     

Effects of water vapour pressure deficit and stomatal conductance on photosynthesis,internal pressurization and convective flow in three emergent wetland plants

Internal pressurization and convective gas flow in emergent wetland plants is a function of the water vapour pressure deficit (WPD) and stomatal conductance (G _s) separating the external atmosphere from the internal aerenchyma. We have compared the effects of WPD and G _s under a range of light intensities on static pressures and convective flows in Phragmites australis, Typha orientalis and Baumea articulata. The capacity of the three species to generate flows per unit leaf area differed, being greatest in P. australisand lowest in B. articulata. In all three species, decreasing light intensity from full sunlight (2200 mol m^–2 s^–1 photosynthetically active photon flux density (PPFD)) to < 200 and < 10 mol m^–2 s^–1PPFD caused immediate decreases in photosynthetic assimilation, followed by more gradual decreases in transpiration and G _s. However, internal pressures and flows in the two low light intensities remained similar to values recorded in full sunlight. WPD was more significantly related to pressures and flows in P. australis and T. orientalis than G _s. In B. articulata, pressures increased at low G _s values but flow rates were unaffected, as predicted by earlier models describing pore size effects on pressures and flows. The data suggest that emergent macrophytes can maintain significant internal convection even at low light intensities, and this may be beneficial for nocturnal aeration, particularly in arid climates where the atmospheric humidity at night is low.     

The relationship between stomatal conductance, transpiration rate and tracheid structure in Pinus radiata clones grown at different water vapour saturation deficits

Abstract. Stomatal conductance and needle water potential of P. radiata clones were measured after 2, 5 and 8 months on plants grown in controlled environment rooms with markedly different water vapour saturation deficits (D). Conductance was significantly lower at high D, but water potential differences between treatments were not significant. When trees were moved between treatments most of the changes in conductances occurred within 2 h, with residual changes after 24 h. Water potentials were not different 24 h after the trees were moved. The effects were completely reversible.
Transpiration rates of individual trees were highest in the high D treatment and lowest in the low D treatment. They were not linearly related to D because of decreasing conductance with increasing D.
Height growth, diameter growth and foliage areas were not significantly different between treatments. Tracheid lumen diameters tended to be larger in trees grown at higher D although treatment differences were not significant.
There were significant clonal differences in shoot conductance and tracheid dimensions.     

Influence of leaf water status on stomatal response to humidity,hydraulic conductance,and soil drought in Betula occidentalis

Whole-canopy measurements of water flux were used to calculate stomatal conductance (g _s ) and transpiration (E) for seedlings of western water birch (Betula occidentalis Hook.) under various soil-plant hydraulic conductances (k), evaporative driving forces (ΔN; difference in leaf-to-air molar fraction of water vapor), and soil water potentials (Ψ_s). As expected, g _s dropped in response to decreased k or Ψ_S, or increased ΔN(> 0.025). Field data showed a decrease in mid-day g _s with decreasing k from soil-to-petiole, with sapling and adult plants having lower values of both parameters than juveniles. Stomatal closure prevented E and Ψ from inducing xylem cavitation except during extreme soil drought when cavitation occurred in the main stem and probably roots as well. Although all decreases in g _s were associated with approximately constant bulk leaf water potential (ψ_l), this does not logically exclude a feedback response between Ψ_L and g _s . To test the influence of leaf versus root water status on g _s , we manipulated water status of the leaf independently of the root by using a pressure chamber enclosing the seedling root system; pressurizing the chamber alters cell turgor and volume only in the shoot cells outside the chamber. Stomatal closure in response to increased ΔN, decreased k, and decreased Ψ_S was fully or partially reversed within 5 min of pressurizing the soil. Bulk Ψ_L remained constant before and after soil pressurizing because of the increase in E associated with stomatal opening. When ΔN was low (i.e., < 0.025), pressurizing the soil either had no effect on g _s , or caused it to decline; and bulk Ψ_L increased. Increased Ψ_l may have caused stomatal closure via increased backpressure on the stomatal apparatus from elevated epidermal turgor. The stomatal response to soil pressurizing indicated a central role of leaf cells in sensing water stress caused by high ΔN, low k, and low Ψ_S. Invoking a prominent role for feedforward signalling in short-term stomatal control may be premature.     

Control of stomatal conductance by leaf water potential in Hymenoclea salsola (T. & G.), a desert subshrub

The role of leaf water potential in controlling stomatal conductance ( g _s) was examined in the desert subshrub Hymenoclea salsola . For plants operating at high irradiance, stomatal closure in response to high leaf-air humidity gradient ( D ) was largely reversed by soil pressurization. Stomatal re-opening eliminated, on average, 89% of the closure normally induced by high D . Transpiration rates ( E ) reached under these conditions were far higher than maximal rates normally observed at any point of the D response. In situ stem psychrometry indicated that water flux at all times conformed to a simple Ohm's-law analogy. Under conditions of high D, E increased substantially in response to soil pressurization. Stomatal regulation did not constrain E during this treatment, but did result in nearly constant minimum leaf water potentials.     

Correlations of stomatal conductance with hydraulic and chemical factors in several deciduous tree species in a natural habitat

Recent research in whole-plant stomatal physiology, conducted largely with potted plants in controlled environments, suggests that stomatal conductance ( g _s) might be more closely linked to plant chemical variables than to hydraulic variables. To test this in a field situation, seasonal g _s was examined in relation to a number of plant and environmental variables in 11 temperate, deciduous forest tree species. Stomatal conductance was generally better correlated with environmental variables (air temperature, vapor pressure deficit, PPFD) than with plant variables, and slightly better correlated with plant hydraulic variables (shoot water and osmotic potentials) than with plant chemical variables (xylem sap ABA concentration, xylem sap pH). We examined a model, developed previously for maize, which describes regulation of g _s by xylem sap ABA concentration with leaf water status acting to modify stomatal sensitivity to the ABA signal. This model explained slightly more variation in seasonal g _s in the forest trees than did single plant variables but not more variation than most single environmental variables. Response surface models, especially those incorporating environmental variables, were more consistently successful at explaining g _s across species.     

High yields in advanced lines of Pima cotton are associated with higher stomatal conductance, reduced leaf area and lower leaf temperature

Advanced lines of Pima cotton ( Gossypium barbadense L.) bred for higher yield potential and heat resistance have higher stomata conductance and smaller leaf areas than those of obsolete lines. In controlled experiments, five commercial lines of Pima cotton having increasing lint yield and heat resistance showed a gradient of increasing stomatal conductance and decreasing leaf size. In field experiments, heat-sensitive, low yield Pima lines showed a lower stomatal conductance than high yielding, advanced lines. This indicates that selection for high yield potential and heat resistance has imposed a selection pressure for higher stomatal conductance and smaller leaf areas. The higher stomatal conductance and smaller leaf area in the advanced lines resulted in a lower leaf temperature in both controlled environments and in the field. The largest leaf temperature differences between obsolete and advanced lines were observed in the afternoon. These differences coincided with the largest differences in stomatal conductance and the highest air temperatures. Measurements of stomatal conductance and leaf temperature in field-grown progeny from a cross between the advanced line, Pima S-6. and the obsolete line, Pima 32, showed that genetically determined differences in stomatal conductance resulted in corresponding differences in leaf temperature. None of the altered physiological traits were selected for in the breeding program, indicating that selection for the desired agronomic traits imposed selection pressures on the altered physiological traits. The increases in stomatal conductance and decreases in leaf area could represent an integrated response to selection pressures on enhanced evaporative cooling, ensuing from selection for heat resistance.     

Effect of elevated CO2 on the stomatal distribution and leaf physiology of Alnus glutinosa

Variation in stomatal development and physiology of mature leaves from Alnus glutinosa plants grown under reference (current ambient, 360 μmol mol⁻¹ CO₂) and double ambient (720 μmol mol⁻¹ CO₂) carbon dioxide (CO₂) mole fractions is assessed in terms of relative plant growth, stomatal characters (i.e. stomatal index and density) and leaf photosynthetic characters. This is the first study to consider the effects of elevated CO₂ concentration on the distribution of stomata and epidermal cells across the whole leaf and to try to ascertain the cause of intraleaf variation. In general, a doubling of the atmospheric CO₂ concentration enhanced plant growth and significantly increased stomatal index. However, there was no significant change in relative stomatal density. Under elevated CO₂ concentration there was a significant decrease in stomatal conductance and an increase in assimilation rate. However, no significant differences were found for the maximum rate of carboxylation ( V _cmax) and the light saturated rate of electron transport ( J _max) between the control and elevated CO₂ treatment.     

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.