Psychrometric Field Measurement of Water Potential Changes following Leaf Excision

In situ measurement of sudden leaf water potential changes has not been performed under field conditions. A laboratory investigation involving the measurement of leaf water potential prior to and 2 to 200 minutes after excision of citrus leaves (Citrus jambhiri) showed good linear correlation (r = 0.99) between in situ leaf psychrometer and Scholander pressure chamber measurements. Following this, a field investigation was conducted which involved psychrometric measurement prior to petiole excision and 1 minute after excision. Simultaneous pressure chamber measurements were performed on neighboring leaves prior to the time of excision and then on the psychrometer leaf about 2 minutes after excision. These data indicate that within the first 2 minutes after excision, psychrometer and pressure chamber measurements were linearly correlated (r = 0.97). Under high evaporative demand conditions, the rate of water potential decrease was between 250 and 700 kilopascals in the first minute after excision. These results show that the thermocouple psychrometer can be used as a dynamic and nondestructive field technique for monitoring leaf water potential.     

A model of drop size distribution for a system with evaporation

Abstract. The size and shape of drops on leaf surfaces strongly affect their persistence. The relationship between volume and exposed surface area of drops on wheat leaves and the log-normal drop size distribution in a wheat canopy after rain are used to derive equations to describe how the total volume and drop number change with evaporation. Firstly, the behaviour of a single drop as it evaporates is considered and then equations describing the change in a population of drops with an initial log-normal distribution are derived. The time taken for all the drops to reach complete dryness is about thirty times that for the same volume of water spread uniformly over the surface with the same potential evaporation rate.     

A New Micro-osmometer

This apparatus measures the rate of distillation at room temperaturebetween the unknown solution and a known standard solution undercarefully denned conditions. It is calibrated using solutionsof known osmotic pressure. It is therefore an empirical vapour-pressureosmometer. The sample (0·06 ml.) is placed in the capsuleC (Fig. 1) and the standard solution contained in the pipetteP, where it is held at a fixed position due to surface tensionat the jet. The upper meniscus is located on the eye-piece scaleof the microscope M. The capillary tube above the meniscus andthe space inside the screw plunger S are air filled. By turningthe screw S the standard solution can be extruded to hang asa drop from the tip of the jet. After a measured time (20 min.)the drop is again drawn into the pipette to the fixed positionat which it comes to rest automatically, and the change in volume,due to condensation or evaporation, measured on the microscopescale. If the sample has a surface tension not much lower than thatof water, the osmometer can be used in the reverse way, thesample being drawn into the pipette and the standard in thecapsule. Used in this way only o-ooi ml. of the sample is needed. A separate calibration curve is required for each standard (Fig.2) and each standard has a range of about 9 atm. (=0·75°C). The standard deviation of single determinations is ±0·06atm. (=0·005° C.). This is uniform over the rangeinvestigated (0-28 atm. or=2·3° C). Satisfactory comparisons between the osmometer and the cryoscopicmethod have been obtained with samples of plant sap (Table I). The osmometer has the advantage of needing only a small volumeof solution which can be viscous, opaque, and need not be freefrom cell debris, &c. It operates at physiological temperaturesand needs no very stringent temperature control. It is simpleto construct and manipulate and occupies less than 1 hour perdetermination of which only 15 minutes is taken up in operation.     

Determination of relationships between water potential and water saturation deficit in leaf tissue

The mutual relationship between the water potential and water saturation deficit (w.s.d.) was studied on samples of leaf tissue of fodder cabage. Definite values of water potential were obtained by long-term exposure of plant material to an atmosphere with definite constant pressure of water vapour. The resulting w.s.d. values were determined gravimetrically. Water saturation deficit varies indirectly with the water potential. This dependence was linear for values of water potential from ?4·4 to ?43·9 atm. Since the stabilization of equilibrum of water potential between the leaf tissue and surrounding atmosphere was very slow the relationship between water potential and w.s.d. was influenced by the size of samples and by the length of exposure. Therefore this method was more suitable for relative than for absolute measurement.     

Effect of Water Stress on Turgor Differences and C-Assimilate Movement in Phloem of Ecballium elaterium

The effects of water stress on pressure differences and ¹⁴C-assimilate translocation in sieve tubes of squirting cucumber Ecballium elaterium A. Rich were studied. Water stress was induced by transfer of plants from culture solution to a polyethylene glycol 6,000 solution having an osmotic potential of −18.2 atm. Sieve tube turgor, turgor differences between source and sink, and translocation rate were decreased. After 260 minutes of translocation, only 19% of the total fixed ¹⁴CO₂ had moved out of the leaf, compared to the control value of 62% after the same period of time. The results suggest that water stress slows translocation by lowering sieve tube turgor differences, which are essential for the pressure flow mechanism of conduction.     

Translocation of C Sucrose in Sugar Beet during Darkness

The time-course of arrival of ¹⁴C translocate in a sink leaf was studied in sugar beet (Beta vulgaris L. cultivar Klein Wanzleben) for up to 480 minutes of darkness. Following darkening of the source leaf, translocation rapidly declined, reaching a rate approximately 25% of the light period rate by 150 minutes. Comparison of data from plants that were girdled 1 cm below the crown with data from ungirdled plants indicates that after about 150 minutes darkness the beet root becomes a source of translocate to the sink leaf. After about 90 minutes darkness, starch-like reserve polysaccharide from the source leaf begins to contribute ¹⁴C to ethanol soluble pools in that leaf. Because of a 15% isotope mass effect, sucrose, at isotopic saturation, reaches a specific activity which is about 85% of the level of the supplied CO₂. The source leaf sucrose specific activity remains at the isotopic saturation level for about 150 minutes of darkness, after which time input from polysaccharide reserves causes the specific activity to drop to about 55% of that of the supplied CO₂. Sucrose specific activity determinations, polysaccharide dissolution measurements, and pulse labeling experiments indicate that following partial depletion of the sucrose pool, source leaf polysaccharide contributes to dark translocation. Respired CO₂ from the source leaf appears to be derived from a pool which, unlike sucrose, remains at a uniform specific activity.     

Leaf water potential-leaf water deficit relationship for ten species of a semiarid grassland community

The relationship between water potential and relative water content (water content in percentage of full hydration) is a characteristic of plant tissues, that may vary with environmental conditions. It is used here to compare leaf water relations of ten species coexisting in a semiarid grassland community (Festucetum vaginatae danubiale) in Hungary. Three groups of species can be distinguished. In two of these leaf water potential changes only moderately with decreasing leaf water content. These are either short-lived, drought escaping spring plants relying on seasonally favourable water supply (group 1) or xerophytes with very deep root system having access to permanent water resources (group 2, only one species studied here). Xerophytes with moderately deep roots (group 3) display a rapid drop of leaf water potential with increasing leaf water deficit. This generates a steep water potential gradient in the soil-plant continuum that in turn enhances water uptake by roots. There is a positive correlation between the rate of water potential decline and degree of sclerophylly (proportion of dry material in the water-saturated leaf), and both variables show seasonal change in perennial species.     

The Control of Water and Carbon Dioxide Flux in Water-stressed Sugar Beet

Changes in leaf and canopy water potential of sugar beet growingin soil of decreasing water content depended on soil water potentialand were independent of water flux from the plant when thiswas varied by changing the water vapour content of the air.The calculated hydraulic conductance of the plant increasedas flux increased and decreased as leaf water potential decreasedand as the plant aged. The conductances to water vapour of individualleaves and of the canopy decreased as leaf water potential decreasedand increased with increasing humidity of the air. The lattereffect was independent of changes in leaf water potential. Theconductances were not affected by the rate of evaporation orleaf temperature. The rate of photosynthesis was directly relatedto leaf conductance except in severely stressed, mature leavesin which leaf water potential had a more direct effect on photosynthesis.Stomatal conductances, transpiration, and photosynthesis weregreater in young leaves than mature leaves on the same plantand at the same leaf water potential. These results are discussedin relation to current agricultural irrigation practices usedfor sugar beet.     

Concurrent comparisons of stomatal behavior, water status, and evaporation of maize in soil at high or low water potential

Concurrent measurements of evaporation, leaf conductance, irradiance, leaf water potential, and osmotic potential of maize (Zea mays L. cv. Pa602A) in soil at either high or low soil water potential were compared at several hours on two consecutive days in July. Hourly evaporation, measured on two weighing lysimeters, was similar until 1000 hours Eastern Standard Time, but thereafter evaporation from the maize in the dry soil was always less than that in the wet soil; before noon it was 62% and by midafternoon, only 35% of that in the wet soil. The leaf water potential, measured with a pressure chamber, was between −1.2 and −2.5 bars and between −6.8 and −8 bars at sunrise (about 0530 hours Eastern Standard Time) in the plants in the wet and dry soil, respectively, but decreased quickly to between −8 and −13 bars in the plants in the wet soil and to less than −15 bars in the plants in the dry soil by 1100 to 1230 hours Eastern Standard Time. At this time, the leaf conductance of all leaves was less than 0.1 cm sec⁻¹ in the maize in the dry soil, whereas the conductance was 0.3 to 0.4 cm sec⁻¹ in the leaves near the top of the canopy in the wet soil. The osmotic potential, measured with a vapor pressure osmometer, also decreased during the morning but to a smaller degree than leaf water potential, so that by 1100 to 1230 hours Eastern Standard Time the leaf turgor potential was 1 to 2 bars in all plants. Thereafter, leaf turgor potential increased, particularly in the plants in soil at a high water potential, whereas leaf water potential continued to decrease even in the maize leaves with partly closed stomata. Evidently maize can have values of leaf conductance differing 3- to 4- fold at the same leaf turgor potential, which suggests that stomata do not respond primarily to bulk leaf turgor potential. Evidence for some osmotic adjustment in the plants at low soil water potential is presented. Although the degree of stomatal closure in the maize in dry soil did not prevent further development of stress, it did decrease evaporation in proportion to the decrease in canopy conductance.     

The behavior of human neutrophils during flow through capillary pores

The passage times of individual human neutrophils through single capillary-sized pores in polycarbonate membranes were measured with the resistive pulse technique, and results were compared to those obtained from the micropipette aspiration of entire cells. Pore transit measurement serves as a useful means to screen populations of cells, and allows for protocols that measure time dependent changes to the population. Neutrophils exhibited a highly linear pressure/flow rate relationship at aspiration pressures from 200 Pa to 1,500 Pa in both the pore and pipette systems. Cellular viscosity, as determined by the method of Hochmuth and Needham, was 89.0 Pa.s for the pore systems and 134.9 Pa.s for the pipette systems. These results are in general agreement with recent values of neutrophil viscosity published in the literature. Extrapolation of the observed linear flow response revealed an apparent minimum pressure for whole cell aspiration significantly above the threshold pressure predicted by Evans' liquid drop model. However, whole cell aspiration was achieved in both the pore and pipette systems at pressures below this extrapolated minimum, although the calculated cellular viscosity was greatly increased. The implications of these two regimes of cell deformation is unclear. This behavior could be explained by shear thinning of the material in the cell body. However the origin of this phenomenon may be in the cortical region of the cell, which exhibits an elastic tension that may be deformation rate dependent.     

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.     

Influence of temperature gradients on leaf water potential

Water potential was monitored at nine locations along single maize (Zea mays L.) leaf blades with aluminum block in situ thermocouple hygrometers. Water potential showed a continuous decrease toward the tip, with a 2- to 4-bar difference between leaf base and tip under both moist and dry soil conditions. The water potential difference between the soil and the leaf base was about 4 bars. Water potentials decreased during the day and during a drying cycle, and increased at night and after irrigation. Heating a band of a leaf to 40 C or cooling it to 7 C had no influence on the water potential of the affected portion when this was corrected for hygrometer output over standard calibrating solutions at the respective temperatures. Heating or cooling a portion of a leaf had neither short nor long term effects on water potential of more distal leaf portions continuously monitored by hygrometers in dew point readout. Water potential fluctuated with an amplitude of about 1.5 bars and an irregular period of 10 to 30 minutes. Measurements with silver foil in situ psychrometers gave similar results.     

Diurnal Changes in Citrus Leaf Thickness, Leaf Water Potential and Leaf to Air Temperature Difference

Diurnal changes in leaf water potential and leaf thickness ofwell-watered citrus trees were found to be highly correlated.Midday decreases in leaf thickness of about 30–35 µm reflected midday decreases in leaf water potential of about1.1–1.3 MPa from predawn values. Leaf water potentialwas also correlated with changes in leaf-to-air temperaturedifference and ambient vapour pressure deficit. Leaf thicknessas well as leaf to air temperature difference could possiblybe used to monitor leaf water status continuously as an indicatorof citrus tree water stress.     

A non-destructive determination of leaf chlorophyll in Vitis vinifera

A portable leaf greenness meter (SPAD-501) has been used to provide a rapid and non-destructive measurement of leaf chlorophyll in Vitis vinifera. Leaf extracted chlorophyll was related linearly to SPAD readings. It is suggested that separate linear equations should be developed for each cultivar so as to maximise the accuracy of estimating leaf chlorophyll content as a function of SPAD readings.     

On the shape of trees

Answers to questions concerning the broad-scale characteristics of trees are sought in the analytical development of a simple model in which the rate of photosynthesis is controlled by leaf water potential and by access to direct solar radiation. These concepts are introduced first in a simple discussion of the single isolated tree. Later they are applied to the forest-stand situation. Expressions are derived which relate growth rate (per tree and per unit area of ground) to the height, shape, spatial separation and lifetime of the trees; to the environmental conditions such as the average solar elevation, potential evaporation E_p and soil moisture content; and to two main physiological factors—the proportionality factor z₀ relating potential drop per unit length of trunk or branch to potential evaporation, and the critical leaf water potential ψ₀ at which net photosynthesis is zero.Accepting the assumptions in the model, the following are examples of its predictions. It is shown that at optimum tree spacing the photosynthesis per unit area of ground may be greatest for the shortest trees (grass ?). It is shown that for a given environment there may be an optimum tree spacing yielding maximum photosynthesis per unit area of ground averaged over the lifetime of the trees; that this maximum decreases with increasing ultimate tree height (which in turn is determined by E_p, z₀ and ψ₀); and that this maximum and this optimum spacing decrease with decreasing average soil moisture. It is shown that there can be an optimum leaf distribution which in general is such that leaf density increases radially from the trunk. It is shown how the optimum shape of trees in a forest might be expected to alter with their size.In general it appears that the concepts discussed here may be useful in explaining the evolutionary development of many of the broad features of tree growth.     

Osmotic Response of Sugar Beet Source Leaves at CO(2) Compensation Point

As sugar beet source leaves lowered the CO₂ concentration to compensation point in a closed atmosphere, leaf thickness and relative water content decreased. Leaf water potential declined rapidly from −0.5 to −1.4 megapascals. At 340 microliters CO₂ per liter, water potential and sucrose, glucose, and fructose contents were steady in photosynthesizing source leaves. Within 90 minutes after leaves were exposed to a CO₂ concentration at the compensation point, leaf sucrose content declined to 60% of the preteatment level, rapidly in the first 30 minutes and then more slowly. During the subsequent 200 minutes, sucrose content increased to 180% of pretreatment level. Glucose and fructose remained unchanged during the treatment. Degradation of starch was sufficient to account for the additional sucrose that accumulated. Labeled carbon lost from starch appeared in sucrose and several other compounds that likely contributed to the recovery in leaf water content.     

Influence of Seismic Stress on Photosynthetic Productivity, Gas Exchange, and Leaf Diffusive Resistance of Glycine max (L.) Merrill cv Wells II

Relative growth rate (RGR), leaf water potential (Ψw), transpiration rate (Tr), photosynthetic rate (Pn), and stomatal and mesophyll resistances to CO₂ exchange were measured or calculated to determine how periodic seismic (shaking) stress decreased dry weight accumulation by soybean (Glycine max [L.] Merrill cv Wells II). Seismic stress was applied with a gyratory shaker at 240 to 280 revolutions per minute for 5 minutes three times daily at 0930, 1430, and 1930 hours. Fifteen days of treatment decreased stem length 21%, leaf area 17%, and plant dry weight 18% relative to undisturbed plants. Seismic stress also decreased RGR 4%, which was due entirely to decreased net assimilation rate. Transpiration decreased 17% and leaf Ψw increased 39% 30 minutes after treatment. A reduction in Pn began within seconds after the onset of treatment and had declined 16% after 20 minutes, at which time gradual recovery began. Assimilation rate recovered fully before the next seismic treatment 5 hours later. Resistance analysis and calculation of leaf internal CO₂ levels indicated that the transitory decrease in Pn caused by periodic seismic stress was due to increased stomatal resistance on the abaxial leaf surface.     

The Measurement of Stomatal Responses to Stimuli in Leaves and Leaf Discs

A comparison has been made of stomatal responses in intact leaves,leaf discs supplied with water via their cut edges and leafdiscs floating on water. Xanthium pennsylvanicum leaf discswatered via their cut edges appeared to be more turgid thanintact leaves; this considerably slowed down the rate of stomatalopening but it slightly increased the final steady-state stomatalopening. When the water potential of such leaf discs was loweredby pre-treatment with mannitol solutions rates of stomatal openingincreased whereas maximum steady-state openings decreased. In tobacco leaf discs floating on water the stomata in contactwith water were wider open than those in contact with normalair and they did not respond to treatment with carbon dioxide-freeair. The rate of photosynthesis was severely reduced in tobaccoleaf discs floating with the lower epidermis on water, mostprobably owing to the slow rate of diffusion of carbon dioxidein water. By floating such discs on osmotica the degree of stomatalopening was increased, however, a response to treatment withcarbon dioxide-free air was still not measurable. It is postulatedthat, on account of the relative unavailability of carbon dioxidefrom the water, the carbon dioxide concentration in the substomatalcavities of the lower surface is abnormally low, irrespectiveof whether ordinary air or carbon dioxide-free air is availableto the upper surface. A comparison between porometer readings and measurements ofsiliconerubber impressions of stomatal pores taken from insidethe porometer cup confirmed that the silicone-rubber impressionmethod of assessing stomatal responses to stimuli has severelimitations, especially at small stomatal apertures.     

A Model for Estimation of Water Flow Resistance in Soil-Leaf Pathway under Dynamic Conditions

The present study was carried out to establish a model for estimatingwater flow resistance in a soil-leaf pathway under field conditions.In this model, the change in leaf water content is taken intoconsideration; the model is based on the assumption that waterflow resistance is essentially constant for relatively shortperiods. Resistance was estimated for three subtropical woodyspecies growing on shallow-soiled ridges. For the estimation,transpiration rate and leaf water potential were measured directlyin the field, and leaf water content was estimated based onthe relationship between leaf water potential and relative leafwater content as observed in the laboratory. Resistance showedlittle variation with change in leaf water potential and transpirationrate by day but was particularly high in the evening in allspecies. The reason for this is not known but was perhaps dueto inaccurate measurement for transpiration rate. It was consideredto be pertinent to compare daily values of resistance so asto assess plant adaptation to drought. The model was shown tobe useful for estimating resistance from conventional measurementsin the field. Key words: Leaf water content, leaf water potential, transpiration rate, water flow resistance     

Isotopic composition of transpiration and rates of change in leaf water isotopologue storage in response to environmental variables

During daylight hours, the isotope composition of leaf water generally approximates steady‐state leaf water isotope enrichment model predictions. However, until very recently there was little direct confirmation that isotopic steady‐state (ISS) transpiration in fact exists. Using isotope ratio infrared spectroscopy (IRIS) and leaf gas exchange systems we evaluated the isotope composition of transpiration and the rate of change in leaf water isotopologue storage (isostorage) when leaves were exposed to variable environments. In doing so, we developed a method for controlling the absolute humidity entering the gas exchange cuvette for a wide range of concentrations without changing the isotope composition of water vapour. The measurement system allowed estimation of ¹⁸O enrichment both at the evaporation site and for bulk leaf water, in the steady state and the non‐steady state. We show that non–steady‐state effects dominate the transpiration isoflux even when leaves are at physiological steady state. Our results suggest that a variable environment likely prevents ISS transpiration from being achieved and that this effect may be exacerbated by lengthy leaf water turnover times due to high leaf water contents.