The Effect of Leaf Water Potential, Leaf Temperature and Light Intensity on Leaf Diffusion Resistance and the Transpiration of Leaves of Malus sylvestris

The relationship between leaf resistance to water vapour diffusion and each of the factors leaf water potential, light intensity and leaf temperature was determined for leaves on seedling apple trees (Malus sylvestris Mill. cv. Granny Smith) in the laboratory. Leaf cuticular resistance was also determined and transpiration was measured on attached leaves for a range of conditions. Leaf resistance was shown to be independent of water potential until potential fell below — 19 bars after which leaf resistance increased rapidly. Exposure of leaves to CO₂-free air extended the range for which resistance was independent of water potential to — 30 bars. The light requirement for minimum leaf resistance was 10 to 20 W m^?2 and at light intensities exceeding these, leaf resistance was unaffected by light intensity. Optimum leaf temperature for minimum diffusion resistance was 23 ± 2°C. The rate of change measured in leaf resistance in leaves given a sudden change in leaf temperature increased as the magnitude of the temperature change increased. For a sudden change of 1°C in leaf temperature, diffusion resistance changed at a rate of 0.01 s cm^?1 min^?1 whilst for a 9°C leaf temperature change, diffusion resistance changed at a rate of 0.1 s cm^?1 min^?1. Cuticular resistance of these leaves was 125 s cm^?1 which is very high compared with resistances for open stomata of 1.5 to 4 s cm^?1 and 30 to 35 s cm^?1 for stomata closed in the dark. Transpiration was measured in attached apple leaves enclosed in a leaf chamber and exposed to a range of conditions of leaf temperature and ambient water vapour density. Peak transpiration of approximately 5 × 10^?6 g cm^?2 s^?1 occurred at a vapour density gradient from the leaf to the air of 12 to 14 g m^?3 after which transpiration declined due presumably to increased stomatal resistance. Leaves in CO₂-free air attained a peak transpiration of 11 × 10^?6 g cm^?2 s^?1 due to lower values of leaf resistance in CO₂ free air. Transpiration then declined in these leaves due to development of an internal leaf resistance (of up to 2 s cm^?1). The internal resistance was masked in leaves at normal CO₂ concentrations by the increase in stomatal resistance.     

The responses of stomata and leaf gas exchange to vapour pressure deficits and soil water content

Summary The responses of photosynthesis, transpiration and leaf conductance to changes in vapour pressure deficit were followed in well-watered plants of the herbaceous species, Helianthus annuus, Helianthus nuttallii, Pisum sativum and Vigna unguiculata, and in the woody species having either sclerophyllous leaves, Arbutus unedo, Nerium oleander and Pistacia vera, or mesomorphic leaves, Corylus avellana, Gossypium hirsutum and Prunus dulcis. When the vapour pressure deficit of the air around a single leaf in a cuvette was varied from 10 to 30 Pa kPa^-1 in 5 Pa kPa^-1 steps, while holding the remainder of the plant at a vapour presure deficit of 10 Pa kPa^-1, the leaf conductance and net photosynthetic rate of the leaf decreased in all species. The rate of transpiration increased initially with increase in vapour pressure deficit in all species, but in several species a maximum transpiration rate was observed at 20 to 25 Pa kPa^-1. Concurrent measurements of the leaf water potential by in situ psychrometry showed that an increase in the vapour pressure deficit decreased the leaf water potential in all species. The decrease was greatest in woody species, and least in herbaceous species. When the vapour pressure deficit around the remainder of the plant was increased while the leaf in the cuvette was exposed to a low and constant vapour pressure deficit, similar responses in both degree and magnitude in the rates of transpiration and leaf conductance were observed in the remainder of the plant as those occurring when the vapour pressure deficit around the single leaf was varied. Increasing the external vapour pressure deficit lowered the water potential of the leaf in the cuvette in the woody species and induced a decrease in leaf conductance in some, but not all, speies. The decrease in leaf conductance with decreasing water potential was greater in the woody species when the vapour pressure deficit was increased than when it remained low and constant, indicating that changing the leaf-to-air vapour pressure difference had a direct effect on the stomata in these species. The low hydraulic resistance and maintenance of a high leaf water potential precluded such an analysis in the herbaceous species. We conclude that at least in the woody species studied, an increase in the vapour pressure deficit around a leaf will decrease leaf gas exchange through a direct effect on the leaf epidermis and sometimes additionally through a lowering of the mesophyll water potential.     

Stomatal control of transpiration from a developing sugarcane canopy

Abstract. Stomatal conductance of single leaves and transpiration from an entire sugarcane (Saccharum spp. hybrid) canopy were measured simultaneously using independent techniques. Stomatal and environmental controls of transpiration were assessed at three stages of canopy development, corresponding to leaf area indices (L) of 2.2, 3.6 and 5.6. Leaf and canopy boundary layers impeded transport of transpired water vapour away from the canopy, causing humidity around the leaves to find its own value through local equilibration rather than a value determined by the humidity of the bulk air mass above the canopy. This tended to uncouple transpiration from direct stomatal control, so that transpiration predicted from measurement of stomatal conductance and leaf-to-air vapour pressure differences was increasingly overestimated as the reference point for ambient vapour pressure measurement was moved farther from the leaf and into the bulk air. The partitioning of control between net radiation and stomata was expressed as a dimensionless decoupling coefficent ranging from zero to 1.0. When the stomatal aperture was near its maximum this coefficient was approximately 0.9, indicating that small reductions in stomatal aperture would have had little effect on canopy transpiration. Maximum rates of transpiration were, however, limited by large adjustments in maximum stomatal conductance during canopy development. The product of maximum stomatal conductance and L. a potential total canopy conductance in the absence of boundary layer effects, remained constant as L increased. Similarly, maximum canopy conductance, derived from independent micrometeorological measurements, also remained constant over this period. Calculations indicated that combined leaf and canopy boundary layer conductance decreased with increasing L such that the ratio of boundary layer conductance to maximum stomatal conductance remained nearly constant at approximately 0.5. These observations indicated that stomata adjusted to maintain both transpiration and the degree of stomatal control of transpiration constant as canopy development proceeded.     

Stomatal control of transpiration in the canopy of a clonal Eucalyptus grandis plantation

 Predawn leaf water potential, stomatal conductance and microclimatic variables were measured on 13 sampling days from November 1995 through August 1996 to determine how environmental and physiological factors affect water use at the canopy scale in a plantation of mature clonal Eucalyptus grandis Hill ex-Maiden hybrids in the State of Espirito Santo, Brazil. The simple ”big leaf” Penman-Monteith model was used to estimate canopy transpiration. During the study period the predawn leaf water potential varied from –0.4 to –1.3 MPa, with the minimum values observed in the winter months (June and August 1996), while the average estimated values for canopy conductance and canopy transpiration fell from 17.3 to 5.8 mm s^–1 and from 0.54 to 0.18 mm h^–1, respectively. On the basis of all measurements, the average value of the decoupling coefficient was 0.25. During continuous soil water shortage a proportional reduction was observed in predawn leaf water potential and in daily maximum values of stomatal conductance, canopy transpiration and decoupling coefficient. The results showed that water vapour exchange in this canopy is strongly dominated by the regional vapour pressure deficit and that canopy transpiration is controlled mainly by stomatal conductance. On a seasonal basis, stomatal conductance and canopy transpiration were mainly related to predawn leaf water potential and, thus, to soil moisture and rainfall. Good results were obtained with a multiplicative empirical model that uses values of photosynthetically active radiation, vapour pressure deficit and predawn leaf water potential to estimate stomatal conductance. Received: 10 June 1998 / Accepted: 20 July 1998     

Diurnal courses and factorial dependencies of leaf conductance and transpiration of differently potassium fertilized and watered field grown barley plants

During the grain filling period we followed diurnal courses in leaf water potential (ψ₁), leaf osmotic potential (ψ_π), transpiration (E), leaf conductance to water vapour transfer (g) and microclimatic parameters in field-grown spring barley (Hordeum distichum L. cv. Gunnar). The barley crop was grown on a coarse textured sandy soil at low (50 kg ha⁻¹) or high (200 kg ha⁻¹) levels of potassium applied as KCl. The investigation was undertaken at full irrigation or under drought. Drought was imposed at the beginning of the grain filling period. Leaf conductance and rate of transpiration were higher in the flag leaf than in the leaves of lower insertion. The rate of transpiration of the awns on a dry weight basis was of similar magnitude to that of the flag leaves. On clear days the rate of transpiration of fully watered barley plants was at a high level during most part of the day. The transpiration only decreased at low light intensities. The rate of transpiration was high despite leaf water potentials falling to rather low values due to high evaporative demands. In water stressed plants transpiration decreased and midday depression of transpiration occurred. Normally, daily accumulated transpirational water loss was lower in high K leaves than in low K leaves and generally the bulk water relations of the leaves were more favourable in high K plants than in low K plants. The factorial dependency of the flag leaf conductances on leaf water potential, light intensity, leaf temperature, and leaf-to-air water vapour concentration difference (ΔW) was analysed from a set of field data. From these data, similar sets of microclimatic conditions were classified, and dependencies of leaf conductance on the various environmental parameters were ascertained. The resulting mathematical functions were combined in an empirical simulation model. The results of the model were tested against other sets of measured data. Deviations between measured and predicted leaf conductance occurred at low light intensities. In the flag leaf, water potentials below-1.6 MPa reduced the stomatal apertures and determined the upper limit of leaf conductance. In leaves of lower insertion level conductances were reduced already at higher leaf water potentials. Leaf conductance was increased hyperbolically as photosynthetic active radiation (PAR) increased from darkness to full light. Leaf conductance as a function of leaf temperature followed an optimum curve which in the model was replaced by two linear regression lines intersecting at the optimum temperature of 23.4°C. Increasing leaf-to-air water vapour concentration difference caused a linear decrease in leaf conductance. Leaf conductances became slightly more reduced by lowered water potentials in the low K plants. Stomatal closure in response to a temperature change away from the optimum was more sensitive in high K plants, and also the decrease in leaf conductance under the influence of lowered ambient humidity proceeded with a higher sensitivity in high K plants. Thus, under conditions which favoured high conductances increase of evaporative demand caused an about 10% larger decrease in leaf conductance in the high K plants than in the low K plants. Stomatal sizes and density in the flag leaves differed between low and high K plants. In plants with partially open stomata, leaf conductance, calculated from stomatal pore dimensions, was up to 10% lower in the high K plants than in the low K plants. A similar reduction in leaf conductance in high K plants was measured porometrically. It was concluded that the beneficial effect of K supply on water use efficiency reported in former studies primarily resulted from altered stomatal sizes and densities.     

Does transpiration control stomatal responses to water vapour pressure deficit?

Three types of observations were used to test the hypothesis that the response of stomatal conductance to a change in vapour pressure deficit is controlled by whole-leaf transpiration rate or by feedback from leaf water potential. Varying the leaf water potential of a measured leaf by controlling the transpiration rate of other leaves on the plant did not affect the response of stomatal conductance to vapour pressure deficit in Glycine max. In three species, stomatal sensitivity to vapour pressure deficit was eliminated when measurements were made at near-zero carbon dioxide concentrations, despite the much higher transpiration rates of leaves at low carbon dioxide. In Abutilon theophrasti, increasing vapour pressure deficit sometimes resulted in both decreased stomatal conductance and a lower transpiration rate even though the response of assimilation rate to the calculated substomatal carbon dioxide concentration indicated that there was no ‘patchy’ stomatal closure at high vapour pressure deficit in this case. These results are not consistent with stomatal closure at high vapour pressure deficit caused by increased whole-leaf transpiration rate or by lower leaf water potential. The lack of response of conductance to vapour pressure deficit in carbon dioxide-free air suggests that abscisic acid may mediate the response.     

The responses of stomata and leaf gas exchange to vapour pressure deficits and soil water content

The responses of leaf conductance, leaf water potential and rates of transpiration and net photosynthesis at different vapour pressure deficits ranging from 10 to 30 Pa kPa^-1 were followed in the sclerophyllous woody shrub Nerium oleander L. as the extractable soil water content decreased. When the vapour pressure deficit around a plant was kept constant at 25 Pa kPa^-1 as the soil water content decreased, the leaf conductance and transpiration rate showed a marked closing response to leaf water potential at-1.1 to-1.2 MPa, whereas when the vapour pressure deficit around the plant was kept constant at 10 Pa kPa^-1, leaf conductance decreased almost linearly from-0.4 to-1.1 MPa. Increasing the vapour pressure deficit from 10 to 30 Pa kPa^-1 in 5 Pa kPa^-1 steps, decreased leaf conductance at all exchangeable soil water contents. Changing the leaf water potential in a single leaf by exposing the remainder of the plant to a high rate of transpiration decreased the water potential of that leaf, but did not influence leaf conductance when the soil water content was high. As the soil water content was decreased, leaf conductances and photosynthetic rates were higher at equal levels of water potential when the decrease in potential was caused by short-term increases in transpiration than when the potential was decreased by soil drying.As the soil dried and the stomata closed, the rate of photosynthesis decreased with a decrease in the internal carbon dioxide partial pressure, but neither the net photosynthetic rate nor the internal CO₂ partial pressure were affected by low water potentials resulting from short-term increases in the rate of transpiration. Leaf conductance, transpiration rate and net photosynthetic rate showed no unique relationship to leaf water potential, but in all experiments the leaf gas exchange decreased when about one half of the extractable soil water had been utilized. We conclude that soil water status rather than leaf water status controls leaf gas exchange in N. oleander.     

Photosynthetic responses of two heliotropic legumes from contrasting habitats

Abstract Photosynthetic responses to light, temperature and leaf-to-air water vapour concentration deficit for Strophostyles helvola (L.) Ell. in an open beach site, and Amphicarpa bracteata (L.) Ell. in two deciduous forest sites were quantified. Photosaturated rates of net CO₂ assimilation were 52.1 ± 4.6, 11.0 ± 1.6 and 4.1 ± 0.3 μmol m^?2s^?2 for plants in beach, roadside and closed canopy sites, respectively. In terms of photosynthesis, plants in the beach site were more tolerant of higher leaf temperatures and water vapour concentration deficits than were plants in forested sites. Heliotropic leaf movements in the beach site reduced calculated total daily transpiration by 2%, increased total daily carbon gain by 8% and reduced the transpiration ratio by 9% relative to an horizontal leaf. During long-term sunflecks in forested sites, heliotropism reduced transpiration by 15%, increased carbon gain by 71% and reduced the transpiration ratio by 50% relative to an horizontal leaf. We hypothesize that heliotropic leaf movements in mesic, high-light, low-nitrogen habitats may increase carbon return on nitrogen investment in photosynthesis, while heliotropism in canopy gaps may represent a morphological mechanism to reduce damage to low-light acclimated photosynthetic systems during long-term sunflecks.     

Stomatal and environmental control of transpiration in a lowland tropical forest tree

Stomatal control of crown transpiration was studied in Anacardium excelsum, a large-leaved, emergent canopy species common in the moist forests of Central and northern South America. A construction crane equipped with a gondola was used to gain access to the uppermost level in the crown of a 35-m-tall individual. Stomatal conductance at the single leaf scale, and transpiration and total vapour phase conductance (stomatal and boundary layer) at the branch scale were measured simultaneously using the independent techniques of porometry and stem heat balance, respectively. This permitted the sensitivity of transpiration to a marginal change in stomatal conductance to be evaluated using a dimensionless coupling coefficient (1-ω) ranging from zero to 1, with 1 representing maximal stomatal control of transpiration. Average stomatal conductance varied from 0.09 mol m^?2 s^?1 during the dry season to 0.3 mol m^?2 s^?1 during the wet season. Since boundary layer conductance was relatively low (0.4 mol m^?2 s^?1), 1-ω ranged from 0.46 during the dry season to only 0.25 during the wet season. A pronounced stomatal response to humidity was observed, which strongly limited transpiration as evaporative demand increased. The stomatal response to humidity was apparent only when the leaf surface was used as the reference point for measurement of external vapour pressure. Average transpiration was predicted to be nearly the same during the dry and wet seasons despite a 1 kPa difference in the prevailing leaf-to-air vapour pressure difference. The patterns of stomatal behaviour and transpiration observed were consistent with recent proposals that stomatal responses to humidity are based on sensing the transpiration rate itself.     

Transpiration rate and intercellular diffusive resistance in the tobacco leaf

The increase in the measured transpiration rate in tobacco leaves due to the experimentally decreased humidity of the bulk air was found to be significantly lower than the theoretical value calculated from the change of water vapour concentration gradients. Boundary layer and stomatal diffusive resistances remained unchanged under experimental conditions with no change of net photosynthetic CO₂ uptake. This suggests an increase in intercellular diffusive resistance with an increase in water vapour concentration gradient which is the driving force of water vapour diffusive part of transpiration flux. The increase can be ascribed to the lengthening of intercellular diffusive pathway as steeper water vapour concentration gradient in intercellular spaces results in an increased evaporating surface of intercellular cells thus moving the effective plane of vaporization in leaf mesophyll further inwards. Due to different and independent changes of concentration gradients for water vapour and CO₂, different length of intercellular diffusive pathways for CO₂ and water vapour may be expected.     

The kinetics of stomatal responses to VPD in Vicia faba: electrophysiological and water relations models

Abstract. An Ohm's law analogy is frequently employed to calculate parameters of leaf gas exchange. For example, resistance to water vapour loss is calculated as the quotient of vapour pressure difference (VPD) and vapour loss by transpiration. In the present research, this electrical analogy was extended. Steady-state transpiration as a function of VPD, assayed in leaflets of Vicia faba using gas exchange techniques, was compared with steady-state K⁺ current magnitude as a function of voltage in isolated guard cell protoplasts of Vicia faba, assayed using the patch clamping technique in the whole cell configuration. An electrophysiological model originally developed to explain the kinetics of current changes following step changes in voltage across a cell membrane was used to fit the kinetics of transpiration changes following step changes in VPD applied to leaflets of Vicia faba. Following step increases in VPD, transpiration exhibited an initial increase, reflecting the increased driving force for water loss and, for large step increases in VPD, a transient decrease in stomatal resistance. Transpiration subsequently declined, reflecting stomatal closure. By analogy to electrophysiological responses, it is hypothesized that the humidity parameter that is sensed by guard cells is VPD. Two models based on epidermal water relations were also applied to transpiration kinetics. In the first model, the transient increase in transpiration following a step increase in VPD was attributed partially to an increase in the Physical driving force (VPD) and partially to a transient decrease in stomatal resistance resulting from reduced epidermal backpressure. In the second model, the transient decrease in stomatal resistance was attributed to a direct response of the guard cells to VPD. Both models based on water relations gave good fits of the data, emphasizing the need for further study regarding the metabolic nature of the guard cell response to humidity.     

Hydraulic architecture of sugarcane in relation to patterns of water use during plant development*

Hydraulic conductance was measured on leaf and stem segments excised from sugarcane plants at different stages of development. Maximum transpiration rates and leaf water potential (Ψ_L) associated with maximum transpiration were also measured in intact plants as a function of plant size. Leaf specific hydraulic conductivity (L_sc) and transpiration on a unit leaf area basis (E) were maximal in plants with approximately 0.2 m² leaf area and decreased with increasing plant size. These changes in Fand L_sc were nearly parallel, which prevented φ_L in larger plants from decreasing to levels associated with substantial loss in xylem conductivity caused by embolism formation. Coordination of changes in E and leaf hydraulic properties was not mediated by declining leaf water status, since φ_L increased with plant size. Hydraulic constrictions were present at nodes and in the node-leaf sheath-leaf blade pathway. This pattern of constrictions is in accord with the idea of plant segmentation into regions differing in water transport efficiency and would tend to confine embolisms to the relatively expendable leaves at terminal positions in the pathway, thereby preserving water transport through the stem.     

Gas-exchange and sap flow measurements of Salix viminalis trees in short-rotation forest

Simultaneous field measurements of transpiration and sap flow were performed on short-rotation Salix viminalis trees ranging in diameter from 1.5 to 3.5 cm (2-year-old shoots on 8-year-old stumps). Transpiration was measured using an open-top ventilated chamber enclosing the whole foliage of a tree. Sap flow was measured using a tree-trunk heat balance (THB) technique with a constant temperature difference and variable heat input. Both the instantaneous and daily values of water flux measured by the two absolute techniques agreed well with a difference of up to about 5%. In July, the hourly transpiration reached a maximum of about 0.2 kg m^–2 (leaf area) or 0.45 kg tree^–1, whereas maximum daily integrals reached 4 kg tree^–1. The response of sap flow rate to abrupt flux change when inducing emboli by cutting-off the stem was very rapid: the registered signal dropped by 85% within 10 min for a specimen with a projected leaf area of 2 m². For S. viminalis trees, transpiration was linearly correlated with stem cross-sectional area and with leaf area.     

Photosynthesis and Transpiration as a Function of Gaseous Diffusive Resistances in Orange Leaves

The CO₂ and H₂O vapour exchange of single attached orange, Citrus sinensis (L.), leaves was measured under laboratory conditions using infrared gas analysis. Gaseous diffusive resistances were derived from measurements at a saturating irradiance and at a leaf temperature optimum for photosynthesis. Variation in leaf resistance (within the range 1.6 to 60 s cm^-1) induced by moisture status, or by cyclic oscillations in stomatal aperture, was associated with changes in both photosynthesis and transpiration. At low leaf resistance (ri less than 10 s cm^-1) the ratio of transpiration to photosynthesis declined with reduced stomatal aperture, indicating a tighter stomatal control over H₂O vapour loss than over CO₂ assimilation. At higher leaf resistance (ri greater than 10 s cm^-1) changes in transpiration and photosynthesis were linearly related, but leaf resistance and mesophyll resistance were also positively correlated, so that strictly stomatal control of photosynthesis became more apparent than real. This evidence, combined with direct measurements of CO₂ diffusive resistances (in a -O₂ gas stream) emphasised the presence of a significant mesophyll resistance; i.e., an additional and rate limiting resistance to CO₂ assimilation over and above that encountered by H₂O vapour escaping from the leaf.     

Alteration of transpiration rate, by changing air vapour pressure deficit, influences leaf extension rate transiently in Miscanthus

A controlled environment chamber for whole plants is described in which vapour pressure deficit (VPD) and temperature can be controlled independently. Plant responses to changes in VPD at constant temperature were measured in terms of leaf extension and plant transpiration rates. Manipulation of VPD independently of temperature was shown to be capable of altering the leaf extension rates of the C4 grass Miscanthus x giganteus grown in hydroponics. The effects of VPD on leaf extension are attributed to changes in transpiration rate and hence leaf water status. It was found that, at a temperature of 20C, the influence of a fixed change in VPD was proportionally less than those observed at temperatures which are close to the threshold for growth (between 6 and 10C). These responses are discussed in relation to our current understanding of the mechanisms of cell growth. The fact that the VPD effects on leaf expansion rates were largely transient suggest that simple models driven by temperature alone are adequate to predict leaf expansion within the temperature range 6-20°C, for this genotype of Miscanthus, in the field.Key words: Leaf growth, Miscanthus x giganteus, temperature, vapour pressure deficit, C4 plants.      

Stomatal and hydraulic conductance in growing sugarcane: stomatal adjustment to water transport capacity*

Abstract Stomatal conductance per unit leaf area in well-irrigated field- and greenhouse-grown sugarcane increased with leaf area up to 0.2 m² plant ¹, then declined so that maximum transpiration per plant tended to saturate rather than increase linearly with further increase in leaf area. Conductance to liquid water transport exhibited parallel changes with plant size. This coordiantion of vapour phase and liquid phase conductances resulted in a balance between water loss and water transport capacity, maintaining leaf water status remarkably constant over a wide range of plant size and growing conditions. The changes in stomatal conductance were not related to plant or leaf age. Partial defoliation caused rapid increases in stomatal conductance, to re-establish the original relationship with remaining leaf area. Similarly, pruning of roots caused rapid reductions in stomatal conductance, which maintained or improved leaf water status. These results suggest that sugarcane stomata adjusted to the ratio of total hydraulic conductance to total transpiring leaf area. This could be mediated by root metabolites in the transpiration stream, whose delivery per unit leaf area would be a function of the relative magnitudes of root system size, transpiration rate and leaf area.     

Contributions of evaporation, isotopic non-steady state transpiration and atmospheric mixing on the delta18O of water vapour in Pacific Northwest coniferous forests

Changes in the ²H and ¹⁸O of atmospheric water vapour provide information for integrating aspects of gas exchange within forest canopies. In this study, we show that diurnal fluctuations in the oxygen isotope ratio (δ¹⁸O) as high as 4‰ were observed for water vapour (δ¹⁸O_vp) above and within an old‐growth coniferous forest in the Pacific Northwest region of the United States. Values of δ¹⁸O_vp decreased in the morning, reached a minimum at midday, and recovered to early‐morning values in the late afternoon, creating a nearly symmetrical diurnal pattern for two consecutive summer days. A mass balance budget was derived and assessed for the ¹⁸O of canopy water vapour over a 2‐d period by considering the ¹⁸O‐isoflux of canopy transpiration, soil evaporation and the air entering the canopy column. The budget was used to address two questions: (1) do δ¹⁸O values of canopy water vapour reflect the biospheric influence, or are such signals swamped by atmospheric mixing? and (2) what mechanisms drive temporal variations of δ¹⁸O_vp? Model calculations show that the entry of air into the canopy column resulted in an isotopically depleted ¹⁸O‐isoflux in the morning of day 1, causing values of δ¹⁸O_vp to decrease. An isotopically enriched ¹⁸O‐isoflux resulting from transpiration then offset this decreased δ¹⁸O_vp later during the day. Contributions of ¹⁸O‐isoflux from soil evaporation were relatively small on day 1 but were more significant on day 2, despite the small H₂¹⁶O fluxes. From measurements of leaf water volume and sapflux, we determined the turnover time of leaf water in the needles of Douglas‐fir trees as ≈ 11 h at midday. Such an extended turnover time suggests that transpiration may not have occurred at the commonly assumed isotopic steady state. We tested a non‐steady state model for predicting δ¹⁸O of leaf water. Our model calculations show that assuming isotopic steady state increased isoflux of transpiration. The impact of this increase on the modelled δ ¹⁸O_vp was clearly detectable, suggesting the importance of considering isotopic non‐steady state of transpiration in studies of forest ¹⁸O water balance.     

Soil Water Stress and Photosynthesis in Cotton

An experiment was conducted in SPAR systems at Florence, S.C., to obtain a data set for use in the simulation of the effect of drying soil on photosynthetic rates in cotton. The plant water status was monitored using leaf water potential and stem diameter meaurements. Reductions were noted in apparent photosynthesis rates after only 5 days of soil drying, and as anticipated, there was uniform displacement of the diurnal cycle of leaf water potential, and corresponding decreases in transpiration and CO₂ uptake. The photosynthesis-light response curves indicated that an average two-fold reduction in photosynthesis rates occurred for solar radiation greater than 250 W/m². Stem diameter change (from a nonstress pre-sunrise value) and integrated stem stress were found to be good indicators of maximum daily plant water stress. The integrated stem stress gave a measure of the duration of the stress along with its magnitude. A simulation method for predicting leaf water potential from stem diameter measurements was used to show that the magnitude and duration of plant water stress increased uniformly during the experiment. This increase was representative of the decreased rates of photosynthesis measured. These data will be used in the simulation of cotton growth and yield.     

On the water balance ofPhytoseiulus persimilis A.-H. and its ecological significance

The net water vapour exchange ofPhytoseiulus persimilis A.-H. is described. Water loss by transpiration increases progressively with ambient temperature. The transpiration rate is directly proportional to the saturation deficit of the air (15 to 30° C) and at constant temperature linearly dependent on the water vapour activity: m_T=–0.81 a_v+0.91 (for a_v 0.0 to 0.85 at 20°C). Phytoseiulus persimilis is able to absorb water vapour from the unsaturated atmosphere. This occurs above a certain threshold (critical equilibrium activity, CEA), which is a_v=0.9 at 15 to 25°C and increases to a_v=0.935 at 30°C.The environmental humidity conditions influencingP. persimilis on the leaf surface are described. The diurnal water vapour profile within the laminar layer at the leaf surface includes periods with water vapour values high enough for these mites to utilize their water vapour sorption capability and to restore a previously-suffered water deficit. In addition,P. persimilis shows a positive hygrotactic behaviour when in a state of water deficit.The survival time of starvingP. persimilis is at least doubled when a possibility to absorb water vapour is available. The water balance at limited food resources is discussed. With a food supply (one prey mite, containing about 5.5 g water) every 3 days and a water vapour activity of a_v=0.76 (20°C), water balance is achieved and the survival time is maximal (approximately 120 days).     

Water Balance of Birch and Larch Leaves and Their Resistance to Short and Progressive Soil Drought

In a greenhouse experiment, which imitated a short (4-day-long) and progressive (3-week-long) soil drought, an infrared gas analyzer was employed to assess transpiration (mol/(m² s)) and leaf transpiration conductivity (g _w, mol H₂O/(m² s)) in intact one-year-old plants of Betula platyphylla Suk. and Larix gmelini (Rupr.) Rupr. grown at the saturation levels of photosynthetically active radiation. The drought duration did not affect the adaptation of leaf water balance, it was determined only by leaf temperature and water supply. At ample water supply and with all other conditions being equal, the transpiration of birch exceeded that of larch by the factor of 3; however, birch was less tolerant to soil drought. The authors conclude that water supply and leaf temperature determine plant resistance to soil drought. The species-specific drought resistance depends on the g _w value.