Co-ordination of vapour and liquid phase water transport properties in plants

The pathway for water movement from the soil through plants to the atmosphere can be represented by a series of liquid and vapour phase resistances. Stomatal regulation of vapour phase resistance balances transpiration with the efficiency of water supply to the leaves, avoiding leaf desiccation at one extreme, and unnecessary restriction of carbon dioxide uptake at the other. In addition to maintaining a long-term balance between vapour and liquid phase water transport resistances in plants, stomata are exquisitely sensitive to short-term, dynamic perturbations of liquid water transport. In balancing vapour and liquid phase water transport, stomata do not seem to distinguish among potential sources of variation in the apparent efficiency of delivery of water per guard cell complex. Therefore, an apparent soil-to-leaf hydraulic conductance based on relationships between liquid water fluxes and driving forces in situ seems to be the most versatile for interpretation of stomatal regulatory behaviour that achieves relative homeostasis of leaf water status in intact plants. Components of dynamic variation in apparent hydraulic conductance in intact plants include, exchange of water between the transpiration stream and internal storage compartments via capacitive discharge and recharge, cavitation and its reversal, temperature-induced changes in the viscosity of water, direct effects of xylem sap composition on xylem hydraulic properties, and endogenous and environmentally induced variation in the activity of membrane water channels in the hydraulic pathway. Stomatal responses to humidity must also be considered in interpreting co-ordination of vapour and liquid phase water transport because homeostasis of bulk leaf water status can only be achieved through regulation of the actual transpirational flux. Results of studies conducted with multiple species point to considerable convergence with regard to co-ordination of stomatal and hydraulic properties. Because stomata apparently sense and respond to integrated and dynamic soil-to-leaf water transport properties, studies involving intact plants under both natural and controlled conditions are likely to yield the most useful new insights concerning stomatal co-ordination of transpiration with soil and plant hydraulic properties.     

The effect of atmospheric humidity on photosynthesis,transpiration and water use efficiency of leaves of several plant species

The effect of humidity on the gas exchange of leaves of the dicotyledons soybean (Glycine max (L.) Merrill), sunflower (Helianthus annuus L.), jojoba (Simmondsia chinensis (L.) Schneider), and saltbush (Atriplex halimus L.) and the monocotyledons wheat (Triticum aestivum L.), barley (Hordeum vulgare L.) sorghum (Sorghum bicolor (L.) Moench) and barnyard grass (Echinochloa crus-galli (L.) Beauv.) was examined under conditions of adequate soil moisture in a controlled environment. Photosynthesis and stomatal and internal diffusion resistances of whole, attached, single leaves were not affected by changes in humidity as the vapour pressure deficit between the leaf and atmosphere ranged from 8 to 27 mb. Transpiration increased linearly with increasing vapour pressure deficit. Whole plants of barley exhibited a different response. As humidity was increased, photosynthesis increased, transpiration expressed per unit of vapour pressure difference increased, and diffusion resistances became smaller. Reasons for the different behaviour of single leaves and whole plants are suggested. An index for water use efficiency, expressed per millibar of vapour pressure deficit, was calculated for single leaves of each species used in the experiments. This showed that water use efficiency was highest in the C₄ xerophytes and lowest in the C₃ mesophytes. The effect of environment on water use efficiency is examined using data from the literature.     

In Situ Estimates of Variable Plant Resistance to Water Flux in Ilex opaca Ait

Xylem pressure potentials and stomatal diffusion resistances were measured in the field in Ilex opaca Ait. during days which differed in temperature and vapor pressure deficit. Water flux into leaves was calculated by combining the field data with laboratory determinations of the relation between tissue water deficit and water potential. Estimates of apparent plant resistance were then calculated from fluxes and differences between soil water potential and xylem tension. The resistance depended strongly on water flux, dropping by a factor of over 7 from low to high water flow rates. This extends the generality of variable plant resistances measured in controlled environment studies to I. opaca as it occurs naturally in the field. The relation of apparent plant resistance to water flux as estimated in this study can be useful in simulation models which calculate water uptake to leaves as a flux driven by a difference in soil and leaf water potentials across a resistance between the bulk soil and the leaf.     

土壤-植物-大气连续体水流阻力分布规律的研究

本文依据田间实测资料,分析了土壤-植物-大气连续体水流阻力的相对重要性,结果表明在连续体中的水流阻力主要分布于从叶气孔腔到大气的扩散过程和根系的吸水过程。叶-气之间的水流阻力比土-根之间要大50倍。最后,讨论了控制连续体水流运动的气孔阻力的变化规律及其与环境因素之间的关系。     

Temperature and Water Relations for Sun and Shade Leaves of a Desert Broadleaf, Hyptis emoryi

The temperature and water relations of sun versus shade leavesof Hyptis emoryi Torr. were evaluated from field measurementsmade in late summer. Throughout most of the day sun leaves hadhigher temperatures and higher resistances to water vapour diffusion,but lower transpiration rates and lower stem water potentials,than did shade leaves. Leaf absorptivity to solar irradiationwas less for 1.5-cm-long sun leaves (0.44) than for 4.0-cm shadeleaves (0.56). For both leaf types the stomatal resistance increasedas the water vapour concentration drop from the leaf to theair increased. Energy balance equations were used together with the measuredtemperature dependence of photosynthesis to predict the effectof variations in leaf absorptivity, length, and resistance onnet photosynthesis. The influence of leaf dimorphism on wholeplants was determined by calculating daily photosynthesis andtranspiration for plants with various percentages of sun andshade leaves. A hypothetical plant with all sun leaves in thesun had about twice the photosynthesis and half the transpirationratio as did plants with sun leaves in the shade or shade leavesin the sun or shade. Plants with both sun and shade leaves hadthe highest predicted photosynthesis per unit ground area. Thepossible adaptive significance of the seasonal variation insun and shade leaf percentages observed for individual H. emoryibushes is discussed in terms of water economy and photosynthesi     

Water loss from cut grass with special reference to hay-making

Water loss from cut grass was studied to determine factors limiting the drying process. An apparatus, used to measure water loss at 28±1°C from blotting paper and from leaves and stem internodes of cocksfoot, consisted of four channels in which air speed was controlled at 25–80 cm s^-1 and relative humidity at 7–68%. The maximum rate of water loss from wet blotting paper was 10500 mg water dm^-2 h^-1 but from leaves and stem internodes supplied with water it was less than 250 mg dm^-2 h^-1. The rate of loss from both plant specimens and blotting paper was linearly related to the vapour pressure differences between the specimen and the surrounding air but was not increased when air speed was changed from 40 to 80 cm s^-1. Grass specimens supplied with water had lower rates of water loss than wet blotting paper because of tissue resistances which were calculated for (a) untreated leaf and stem specimens, (b) rubbed leaves, (c) cut leaves, (d) leaves exposed to steam for 60 s. Treatments (b)-(d) greatly reduced tissue resistances. The rates of drying of leaves and stem internodes not supplied with water changed only slightly in response to faster air speeds but were significantly increased by treatments (b), (c), (c_s) (split stems), (d) and (e) (exposure to petroleum vapour for 60 s). The most effective treatments trebled the drying rates of leaves and increased the drying rates of stem internodes by 10 times. Reductions in relative humidity had little effect on drying rate following treatments (a), (b), (c) and (d), but when treatments (c_s) and (e) were given, additional significant increases in drying rates were obtained when the relative humidity was reduced. Grass specimens given the most effective treatments and dried under the most favourable conditions did not utilize the full drying capacity of the environment, for the rates of water loss from these specimens were at least three times lower than those from wet blotting paper. The results indicate that high rates of drying could be achieved at 28 ^oC or similar temperatures if practical treatments were developed to remove or greatly reduce the high resistance to water loss in cut grass.     

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.     

Long-term hydraulic acclimation to soil texture and radiation load in cotton

The concept of root contact hypothesizes that the absorbing roots grown in sandy soil are only partially effective in water uptake. Co-ordination of water supply and demand in the plant requires that the capacity for water uptake from the soil should correspond to an operational rate of water loss from the leaves. To examine how the plant hydraulic system responds to variations in soil texture or evaporative demand through long-term acclimation, an experiment was carried on cotton plants (Gossypium herbaceum L.), where three grades of soil texture and three grades of evaporative demand were applied for the whole life cycle of the plants. Plants were harvested 50 and 90 d (fully grown) after sowing and root length and leaf area measured. At 90 d hydraulic conductance was measured as the ratio of sap flow (measured with sap flow sensors or gravimetrically) and water potential. Results showed that for plants grown at the same evaporative demand, those in sandy soil, where root-specific hydraulic conductance was low, developed more absorbing roots than those grown in heavy-textured soil, where root specific conductance was high. This resulted in the same leaf specific hydraulic conductance (1.8 × 10⁻⁴ kg s⁻¹ Mpa⁻¹ m⁻²) for all three soils. For plants grown in the same sandy soil, those subjected to strong evaporative demand developed more absorbing roots and higher leaf-specific hydraulic conductance than those grown under mild evaporative demand. It is concluded that when soil texture or atmospheric evaporative demand varies, plants co-ordinate their capacities for liquid phase and vapour phase water transport through long-term acclimation of the hydraulic system, or plastic morphological adaptation of the root/leaf ratio.     

Nutrient status and mycorrhizal enhancement of water transport in soybean

Mycorrhizal soybean (Glycine max L. Merr. var. Harosoy-63) plants have lower resistances to water transport than nonmy-corrhizal plants after 4.5 weeks of growth. Although resistances of whole plants differ by 40%, there were no differences in the resistances of stems plus leaves, indicating that the major effect of the mycorrhizae was to reduce the resistance of the roots. Since the fungitoxicant, p-chloronitrobenzene, had no effect on resistances to water transport, reduced resistances were probably not caused by a direct modification of the transport pathway by the fungus. Differences in resistance between mycorrhizal and nonmycorrhizal soybean were essentialy eliminated by the application of nutrients to the soil. Thus, lowered resistances of mycorrhizal roots growing in soil with low levels of nutrients probably resulted from the enhanced nutrient status of the plant brought about by the fungus. Mycorrhizal infection increased growth at both low and high nutrient levels.     

Comparative resistance of the soil and the plant to water transport

The resistances to liquid water transport in the soil and plant were determined directly and simultaneously from measurements of soil, root, and leaf water potentials and the flux of water through the soil-plant system to the sites of evaporation in the leaf. For soybean (Merr.) transporting water at a steady rate, water potential differences between soil and root were smaller than between root and leaf over the range of soil water potentials from −0.2 to −11 bars. As soil water was depleted, water flow through the soil and plant decreased to one-tenth the maximum rate, but both the soil resistance and plant resistance increased. The plant resistance remained larger than the soil resistance over the entire range of soil water availability. Previous suggestions that the soil is the major resistance have ignored the increase in plant resistance and/or assumed root densities that were too low.     

Water Vapour and Heat Transfer in Leaves

Factors connected with the formation of water droplets in leavesby distillation from the mesophyll to the epidermis were investigatedin a number of species. It was concluded that in illuminatedleaves water droplets form principally on the inner walls ofguard and subsidiary cells, and sometimes below the anticlinalwalls of epidermal cells, because these sites are cooler thanthe rest of the leaf. Under more isothermal conditions any waterdroplets that had formed disappeared. With increasing waterstress water droplets did not form so readily, though distillationwas occurring. Few water droplets were observed in leaves outof doors that had open stomata. Significant temperature gradientswere measured across leaves with thermocouples, but these werelarger than were gradients calculated from measured thermalconductivities of leaves. The evaporation resistances of theinner walls of the epidermis and of the mesophyll were foundto be similar. This led to the conclusion that the hydrophobicityof the surfaces of these tissues is similar. Water transferin leaves in the vapour phase was found to be more responsiveto temperature than to water stress gradients. leaf, evaporation, distillation, heat loss, transpiration     

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.     

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.     

A Quantitative Study of the Resistances to Transpirational Water Movement in Sunflower (Helianthus annuus L.)

Hydroponic sunflower plants were used in a quantitative studyof the relationship between total plant and leaf resistancesto transpirational water movement and transpiration rate. Theresults demonstrate that both resistances are flux-dependentand decline 5–6-fold during a comparable increase in transpiration.The resistance of excised leaves including the petiole was approximatelyhalf the total plant resistance. Subsequent analyses of the water potential gradients and transpirationalfluxes in whole plants permitted calculation of the magnitudeof the partial resistances imposed by roots, stem, petiole,and leaf. The root and leaf resistances were approximately 50%and 30% of the total resistance respectively. Stem and petiolarresistances were relatively small and both influenced watermovement to the upper leaves similarly. The values obtainedare compared with previous published results obtained usingdiverse experimental techniques.     

The loss of electrolytes from leaves treated with hydrocarbons and their derivatives

Indices of the effects on the leaves of a number of plant species of members of homologous series of hydrocarbons and certain of their oil-soluble derivatives were based on changes in conductance of aqueous systems when hydrocarbon-treated leaves were added. The results were compared with previous phytotoxicity studies where hydrocarbons were applied to plants in vapour and in aqueous treatments. A relationship between the conductance changes and the physical properties of the applied hydrocarbons is suggested. The presumed effects on cell membrane permeability indicated by the conductance changes agreed with known susceptibility of different plant species towards hydrocarbons of different homologous series applied in the liquid state.     

Changes in minimal diffusive resistances of leaf epidermes during ageing of primary leaves of phaseolus vulgaris L.

Minimal diffusive resistances of both leaf epidermes increased during normal and retarded ageing (moving secondary leaves aside, plant decapitation). The retarded ageing of primary leaves slowed down increase in epidermal resistance (r_ep), and was also reflected in the prolongation of their growth, increase of leaf area, size of epidermal and guard cells and stomatal pores. Decrease of stomata density was to some extent balanced by an increase in the pore size; the later rapid rise in diffusion resistance was induced by the loss of stomata ability to open fully.     

Validation of leaf and microbial pectinases: commercial launching of a new platform technology

Almost all current genetically modified plant commercial products are derived from seeds. The first protein product made in leaves for commercial use is reported here. Leaf pectinases are validated here with eight liquid commercial microbial enzyme products for textile or juice industry applications. Leaf pectinases are functional in broad pH/temperature ranges as crude leaf extracts, while most commercial enzyme products showed significant loss at alkaline pH or higher temperature, essential for various textile applications. In contrast to commercial liquid enzymes requiring cold storage/transportation, leaf pectinase powder was stored up to 16 months at ambient temperature without loss of enzyme activity. Commercial pectinase products showed much higher enzyme protein PAGE than crude leaf extracts with comparable enzyme activity without protease inhibitors. Natural cotton fibre does not absorb water due to hydrophobic nature of waxes and pectins. After bioscouring with pectinase, measurement of contact‐angle water droplet absorption by the FAMAS videos showed 33 or 63 (leaf pectinase), 61 or 64 (commercial pectinase) milliseconds , well below the 10‐second industry requirements. First marker‐free lettuce plants expressing pectinases were also created by removal of the antibiotic resistance aadA gene. Leaf pectinase powder efficiently clarified orange juice pulp similar to several microbial enzyme products. Commercial pilot scale biomass production of tobacco leaves expressing different pectinases showed that hydroponic growth at Fraunhofer yielded 10 times lower leaf biomass per plant than soil‐grown plants in the greenhouse. Pectinase enzyme yield from the greenhouse plants was double that of Fraunhofer. Thus, this leaf‐production platform offers a novel, low‐cost approach for enzyme production by elimination of fermentation, purification, concentration, formulation and cold chain.     

The long-distance abscisic acid signal in the droughted plant: the fate of the hormone on its way from root to shoot

Abscisic acid (ABA) is a potent molecule that certainly modifies stomatal behaviour and plant water loss and probably acts to modify the growth of leaves. The hormone is synthesized both in the leaves and the roots of the plant and in the soil and may move freely from plant to soil and soil to plant. It can also move rapidly through the plant in both the xylem and the phloem and will partition between different compartments in different tissues largely as a function of pH. It is described here how perturbations in soil conditions around the roots and the water status of the air can modify the fluxes of ABA around the plant and its accumulation in different compartments and different tissues. These fluxes can be interpreted as signals of different stresses imposed on the plant and consideration is given to how different perturbations can exert subtle changes which are manifest as modified shoot growth rates and functioning. Most emphasis in the discussion is placed upon the plant's responses to the imposition of soil and atmospheric drought.     

The temperature dependence of shoot hydraulic resistance: implications for stomatal behaviour and hydraulic limitation

Recent soil pressurization experiments have shown that stomatal closure in response to high leaf–air humidity gradients can be explained by direct feedback from leaf water potential. The more complex temperature‐by‐humidity interactive effects on stomatal conductance have not yet been explained fully. Measurements of the change in shoot conductance with temperature were made on Phaseolus vulgaris (common bean) to test whether temperature‐induced changes in the liquid‐phase transport capacity could explain these temperature‐ by‐humidity effects. In addition, shoot hydraulic resistances were partitioned within the stem and leaves to determine whether or not leaves exhibit a greater resistance. Changes in hydraulic conductance were calculated based on an Ohm’s law analogy. Whole‐plant gas exchange was used to determine steady‐ state transpiration rates. A combination of in situ psychrometer measurements, Scholander pressure chamber measurements and psychrometric measurements of leaf punches was used to determine water potential differences within the shoot. Hydraulic conductance for each portion of the pathway was estimated as the total flow divided by the water potential difference. Temperature‐induced changes in stomatal conductance were correlated linearly with temperature‐induced changes in hydraulic conductance. The magnitude of the temperature‐induced changes in whole‐plant hydraulic conductance was sufficient to account for the interactive effects of temperature and humidity on stomatal conductance.     

Stomatal diffusion resistances and leaf growth during droughting of Lolium perenne plants selected for contrasting epidermal ridging

Groups of Lolium perenne plants selected for either deep or shallow adaxial epidermal ridging were grown in a 16 h day of 70 W m^-2 at 25°C, and either watered daily to 33% or allowed to dry to and then watered daily to 21% or to 16% soil moisture. During a 9 day experimental period, adaxial leaf resistances (r₁) were measured with a diffusion porometer four times daily, transpiration was estimated gravimetrically, and daily rates of leaf extension were recorded. Measurements were also made of minimum abaxial resistances, stomatal frequencies and lengths, and relative leaf water content (RLWC). At 33%, 21% or 16% soil moisture, leaf extension rates of deep ridged leaves were, respectively, slower, the same, and more rapid than those with shallow ridges. At 21% or 16% soil moisture, the adaxial r_l of deep-ridged was much lower than that of shallow-ridged leaves at all four sampling times. This difference was most marked on leaves below the youngest fully expanded, and was observed among older leaves even when plants were well watered. At low RLWC (< 85%), leaf resistance was greatest in leaves with shallow ridges. There was no significant difference between the leaf types in the calculated contributions of stomatal frequency or of morphology at any one pore opening, to r_l but deep-ridged leaves had more stomataonthe abaxial surface. Daily rate of plant water loss was directly correlated (r=+ 0.86, P < 0.01) with mean daily maximum stomatal conductance (1/r_l), and rate of leaf extension negatively with maximum r_l. It is suggested that stomata operating in the concavity formed by deep ridges open wider and are less responsive to internal changes in, for example, leaf water status, than those on shallow-ridged leaves because of a more humid microenvironment at the epidermal surface. The results are discussed in relation to the concept of ‘water-savers’ and ‘water-spenders’ and its application to breeding for dry conditions.