Quantification of excess water loss in plant canopies warmed with infrared heating

Here we investigate the extent to which infrared heating used to warm plant canopies in climate manipulation experiments increases transpiration. Concerns regarding the impact of the infrared heater technique on the water balance have been raised before, but a quantification is lacking. We calculate transpiration rates under infrared heaters and compare these with air warming at constant relative humidity. As infrared heating primarily warms the leaves and not the air, this method increases both the gradient and the conductance for water vapour. Stomatal conductance is determined both independently of vapour pressure differences and as a function thereof, while boundary layer conductance is calculated using several approaches. We argue that none of these approaches is fully accurate, and opt to present results as an interval in which the actual water loss is likely to be found. For typical conditions in a temperate climate, our results suggest a 12–15% increase in transpiration under infrared heaters for a 1 °C warming. This effect decreases when stomatal conductance is allowed to vary with the vapour pressure difference. Importantly, the artefact is less of a concern when simulating heat waves. The higher atmospheric water demand underneath the heaters reflects naturally occurring increases of potential evapotranspiration during heat waves resulting from atmospheric feedback. While air warming encompasses no increases in transpiration, this fully depends on the ability to keep humidity constant, which in the case of greenhouses requires the presence of an air humidification system. As various artefacts have been associated with chamber experiments, we argue that manipulating climate in the field should be prioritized, while striving to limit confounding factors. The excess water loss underneath infrared heaters reported upon here could be compensated by increasing irrigation or applying newly developed techniques for increasing air humidity in the field.     

A chamber for applying pressure to roots of intact plants

A chamber was designed to apply up to 20 bars pressure to roots of intact plants. The unique features of this chamber are a split top arrangement to permit enclosing roots of intact plants within the chamber, a circulation coil to control temperature of rooting media, and a valve arrangement to permit changing solution without disturbing the plant. Changes in transpiration in response to changes in the pressure applied to roots of intact pepper plants illustrate one use of the equipment. Well watered plants at low light (0.05 langley/min) were observed to exude water from the leaf margins when 5 bars pressure was applied to the roots. When roots were cut off, a 1 bar pressure caused exudation. Plants with cooled roots or plants in dry soil did not exude water when as much as 6 bars pressure was applied. Transient response of transpiration rates to sudden application and release of pressure was observed in pepper and bean plants but not in rhododendron. The magnitude of this transient response was highly dependent upon light intensity and CO₂ concentration in the aerial environment.     

Effects of Humidity on Photosynthesis

It was found for two species that net carbon dioxide uptakerates were reduced at constant intercellular carbon dioxidepartial pressure when single attached leaves were exposed tolarge leaf to air water vapour pressure differences. Leaf temperature,irradiance, and ambient carbon dioxide and oxygen partial pressureswere kept constant. Net carbon dioxide uptake rates decreasedlinearly with increasing vapour pressure difference, even incases where transpiration rates were highest at intermediatevalues of vapour pressure difference. Decreases in net carbondioxide uptake rates were quickly reversible. Different windspeeds across the measured leaf, different vapour pressure deficitsaround the rest of the shoot, and transient responses of netcarbon dioxide uptake rate to abrupt changes in vapour pressuredifference all gave the same response of net carbon dioxideuptake rate to vapour pressure difference. The data show thatthe inhibition of net carbon dioxide uptake rate at a givenvapour pressure difference was not simply related to whole leaftranspiration rate or stomatal conductance. Key words: Vapour pressure difference, CO₂ uptake rate, Leaf temperature     

A system for measuring leaf gas exchange based on regulating vapour pressure difference

A system for measurement of leaf gas exchange while regulating leaf to air vapour pressure difference has been developed; it comprises an assimilation chamber, leaf temperature controller, mass flow controller, dew point controller and personal computer. A relative humidity sensor and air and leaf temperature sensors, which are all used for regulating the vapour pressure difference, are mounted into the chamber. During the experiments, the computer continuously monitored the photosynthetic parameters and measurement conditions, so that accurate and intenstive measurements could be made.When measuring the light-response curve of CO₂ assimilation for single leaves, in order to regulate the vapour pressure difference, the leaf temperature and relative humidity in the chamber were separately and simultaneously controlled by changing the air temperature around the leaf and varying the air flow rate through the chamber, respectively. When the vapour pressure difference was regulated, net CO₂ assimilation, transpiration and leaf conductance for leaves of rice plant increased at high quantum flux density as compared with those values obtained when it was not regulated.When measuring the temperature-response curve of CO₂ assimilation, the regulation of vapour pressure difference was manipulated by the feed-forward control of the dew point temperature in the inlet air stream. As the vapour pressure difference was regulated at 12 mbar, the maximum rate of and the optimum temperature for CO₂ assimilation in rice leaves increased 5 molCO₂ m^–2 s^–1 and 5°C, respectively, as compared with those values obtained when the vapour pressure difference took its own course. This was reasoned to be due to the increase in leaf conductance and the decrease in transpiration rate. In addition, these results confirmed that stomatal conductance essentially increases with increasing leaf temperature under constant vapour pressure difference conditions, in other words, when the influence of the vapour pressure difference is removed.This system may be used successfully to measure inter- and intra-specific differences and characteristics of leaf gas exchange in plants with a high degree of accuracy.Abbreviations A CO₂ assimilation rate - A_max Maximum rate of CO₂ assimilation - A_opt Optimum teperature for CO₂ assimilation - CTWB Controlled-temperature water bath - DPC Dew point controller - E Transpiration rate; gl, leaf conductance - HCC Humidity control circuit - IRGA Infrared gas analyzer - LT Leaf temperature - LTC Leaf temperature controller - MFC Mass flow controller - QFD Quantum flux density - RH Relative humidity - RHC Relative humidity controller - VPD Vapour pressure difference - CO₂ Difference of CO₂ concentration between inlet and outlet air     

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.     

Relationships between nitrogen uptake and carbon assimilation in whole plants of tall fescue

Abstract. The present study investigates the relationships between nitrogen uptake, transpiration, and carbon assimilation. Plants growing on nutrient solution were enclosed for 10–16 d in a growth chamber, where temperature, photon flux density, vapour saturation deficit and CO₂ concentration were controlled. One of these factors was modified every 4 to 5 d. Shoot photosynthesis and root and shoot respiration were recorded every half-hour. Nitrogen uptake from the root medium and plant transpiration were measured daily. In most cases, an increase in photon flux density led to increases in transpiration, net daily carbon assimilation, and nitrogen uptake. By modifying transpiration rate without changing photosynthesis (varying vapour saturation deficit), or by modifying transpiration and carbon assimilation in opposite ways (varying CO₂ air concentration), it was shown that nitrogen uptake does not follow transpiration, but is linked to the carbon uptake of the plant. When light was increased from low to intermediate levels, the N uptake/C assimilation ratio remained constant. At higher photon flux density, this ratio declined markedly. It is proposed that in the first case, growth is limited by carbohydrate availability, thus any increase in carbon assimilation leads to a proportional increase in nitrogen uptake, in contrast to the second situation where carbohydrates may accumulate in the plant without further nitrogen requirement.     

Effect of air and soil temperature on water balance of jojoba growing under controlled conditions

Jojoba [ Simmondsia chinensis (Link) Schneider] cuttings were grown in pots under constant light intensity and vapour pressure deficit at wir temperatures of 18 and 27°C in climate-controlled cabinets. Leaf conductance and transpiration rate decreased exponentially as the xylen water potential (Ψ_x) decreased concurrently with the drying out of the soil. At high Ψ_x'leaf conductance and transpiration rate were much higher at the higher air temperature, and as Ψ_x declined both parameters decreased more rapidly at 27°C than at 18°C. When soil temperatures were decreased from 27 to 13°C, leaf water potential was not affected at either air temperatures, but transpiration rate was reduced. A linear negative correlation was found between transpiration rates and soil temperatures. It is suggested that the low soil temperature may restrict reducion of water flux in turn reduces stomatal conductance and transpiration without affecting the water potential in the shoot. The releavance of the response to changes in soil or air temperature to the performance of the plant in its semi-arid habitat is discussed.     

Novel Evidence for a Lack of Water Vapour Saturation Within the Intercellular Airspace of Turgid Leaves of Mesophytic Species

Ward, D. A. and Bunce, J. A. 1986. Novel evidence for a lackof water vapour saturation within the intercellular airspaceof turgid leaves of mesophytic species—J. exp. Bot. 37:504– By utilizing a dual-surface leaf chamber evidence was obtainedsuggesting that the water vapour pressure within the intercellularairspace of turgid leaves of mesophytic species can deviatesignificantly from the saturation vapour pressure at the leaftemperature. When the water vapour pressure of the air surroundingthe lower leaf surface of sunflower was maintained constantand high, suddenly exposing the upper leaf surface to air witha low water vapour content caused the lower leaf surface toexhibit a negative rate of transpiration (i.e. an apparent uptakeof water vapour). Since the vapour pressure of the air surroundingthe lower (moist) surface was less than the saturation vapourpressure at the leaf temperature, the occurrence of negativetranspiration indicated that the vapour pressure of the leafairspace deviated from saturation under the conditions of measurementused. For both soybean and sunflower it was also found that if thehumidity around the upper surface was maintained high and constant,a stepwise decrease in lower surface humidity caused substantialreductions in the transpiration rate and apparent conductanceof the upper surface without any concomitant change in its photosyntheticrate. In contrast, both the photosynthetic rate and conductanceof the lower surface were greatly reduced. The relative reductionsof photosynthetic rate and conductance at the lower surfacewere the same. These responses are most easily explained interms of a deviation from water vapour saturation within theintercellular airspace, which gives rise to spurious valuesof conductance. Key words: Intercellular space, water vapour pressure, turgid, leaves, mesophyte     

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.     

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.     

Changes in diffusion resistance of apple leaves undergoing rapid fluctuations in leaf temperature

Transpiration was measured in apple leaves (Malus sylvestris Mill.) which were enclosed in a leaf chamber and subjected to rapid changes in leaf temperature. Fluctuations in leaf temperature produced parallel fluctuations in transpiration. The change in transpiration rate with change in temperature was found to be less than the theoretical value calculated from the change in water vapour density gradient from leaf to air. The results suggest the presence of a small and rapidly varying resistance to water vapour loss from the leaf. The magnitude of this additional resistance increased to a maximum value of approximately 1.5 s cm^-1 as the magnitude of the temperature change increased to a maximum of approximately 12°C.     

A new stem hygrometer, corrected for temperature gradients and calibrated against the pressure bomb

Abstract A simple stem hygrometer for attachment to a bared section of sapwood or a cross-sectional cut end of a shoot is described. Two welded chromelconstantan thermocouples inside the chamber, one touching the sample and the other in the chamber air, allowed measurement of and correction for the temperature gradient between the sample and the dewpoint measuring junction. The instrument was attached to the cut end of an apical shoot of Thuja occidentalis L. protuding from a Scholander-Hammel pressure bomb. Cut-end water potential (ψ_hyg), measured using the stem hygrometer, was compared to xylem pressure potential (ψ_xp) while the latter was manipulated in the pressure bomb. After an initial equilibration time of 3–4 h, hygrometer equilibrium values were achieved within 1.5–4.0 min of changing ψ_xp in the pressure bomb. The half-time (ψ_1/2) for vapour pressure equilibration was 15–40 s. Stable temperature gradients between the sample and dewpoint measuring junction of 0.01–0.1°C were measured. Correcting ψ_hyg for the temperature gradient resulted in excellent agreement with ψ_xp.     

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.     

Spatial distribution of leaf nitrogen and photosynthetic capacity within the foliage of individual trees: disentangling the effects of local light quality,leaf irradiance,and transpiration

There is presently no consensus about the factor(s) driving photosynthetic acclimation and the intra-canopy distribution of leaf characteristics under natural conditions. The impact was tested of local (i) light quality (red/far red ratio), (ii) leaf irradiance (PPFD(i)), and (iii) transpiration rate (E) on total non-structural carbohydrates per leaf area (TNC(a)), TNC-free leaf mass-to-area ratio (LMA), total leaf nitrogen per leaf area (N(a)), photosynthetic capacity (maximum carboxylation rate and light-saturated electron transport rate), and leaf N partitioning between carboxylation and bioenergetics within the foliage of young walnut trees grown outdoors. Light environment (quantity and quality) was controlled by placing individual branches under neutral or green screens during spring growth, and air vapour pressure deficit (VPD) was prescribed and leaf transpiration and photosynthesis measured at branch level by a branch bag technique. Under similar levels of leaf irradiance, low air vapour pressure deficit decreased transpiration rate but did not influence leaf characteristics. Close linear relationships were detected between leaf irradiance and leaf N(a), LMA or photosynthetic capacity, and low R/FR ratio decreased leaf N(a), LMA and photosynthetic capacity. Irradiance and R/FR also influenced the partitioning of leaf nitrogen into carboxylation and electron light transport. Thus, local light level and quality are the major factors driving photosynthetic acclimation and intra-canopy distribution of leaf characteristics, whereas local transpiration rate is of less importance.     

Involvement of root ABA and hydraulic conductivity in the control of water relations in wheat plants exposed to increased evaporative demand

We studied the possible involvement of ABA in the control of water relations under conditions of increased evaporative demand. Warming the air by 3°C increased stomatal conductance and raised transpiration rates of hydroponically grown Triticum durum plants while bringing about a temporary loss of relative water content (RWC) and immediate cessation of leaf extension. However, both RWC and extension growth recovered within 30 min although transpiration remained high. The restoration of leaf hydration and growth were enabled by increased root hydraulic conductivity after increasing the air temperature. The use of mercuric chloride (an inhibitor of water channels) to interfere with the rise on root hydraulic conductivity hindered the restoration of extension growth. Air warming increased ABA content in roots and decreased it in shoots. We propose this redistribution of ABA in favour of the roots which increased the root hydraulic conductivity sufficiently to permit rapid recovery of shoot hydration and leaf elongation rates without the involvement of stomatal closure. This proposal is based on known ability of ABA to increase hydraulic conductivity confirmed in these experiments by measuring the effect of exogenous ABA on osmotically driven flow of xylem sap from the roots. Accumulation of root ABA was mainly the outcome of increased export from the shoots. When phloem transport in air-warmed plants was inhibited by cooling the shoot base this prevented ABA enrichment of the roots and favoured an accumulation of ABA in the shoot. As a consequence, stomata closed.     

Effects of polyethylene-glycol-induced osmotic stress on transpiration and photosynthesis in pinto bean leaf discs

A new leaf disc chamber allows measurements of chlorophyll fluorescence and CO₂ and H₂O vapor exchanges during infusion of solution into the cut edge of the disc. Polyethylene glycol (molecular weight, 6000) was used to apply a mild external osmotic stress to the leaf disc within this chamber. This stress rapidly caused a temporary increase in transpiration. This increase was reversed (5-6 minutes later) and after 20 to 25 min, the stomates nearly completely closed. Internal CO₂ (calculated) and leaf temperature followed the transpiration measurements. However, chlorophyll fluorescence (small rise) followed internal CO₂ (small rise). This complete sequence of events resembles those caused by exposure of leaves to certain air pollutants which have been seen to cause such a transient increase followed by a decrease in stomatal closure.     

Hydraulic control of stomatal conductance in Douglas fir [Pseudotsuga menziesii (Mirb.) Franco] and alder [Alnus rubra (Bong)] seedlings

Experiments were conducted on 1-year-old Douglas fir [Pseudotsuga menziesii (Mirb.) Franco] and 2- to 3-month-old alder [Alnus rubra (Bong)] seedlings growing in drying soils to determine the relative influence of root and leaf water status on stomatal conductance (g_c). The water status of shoots was manipulated independently of that of the roots using a pressure chamber that enclosed the root system. Pressurizing the chamber increases the turgor of cells in the shoot but not in the roots. Seedling shoots were enclosed in a whole-plant cuvette and transpiration and net photosynthesis rates measured continuously. In both species, stomatal closure in response to soil drying was progressively reversed with increasing pressurization. Responses occurred within minutes of pressurization and measurements almost immediately returned to pre-pressurization levels when the pressure was released. Even in wet soils there was a significant increase in g_c with pressurization. In Douglas fir, the stomatal response to pressurization was the same for seedlings grown in dry soils for up to 120 d as for those subjected to drought stress over 40 to 60 d. The stomatal conductance of both Douglas fir and alder seedlings was less sensitive to root chamber pressure at higher vapour pressure deficits (D), and stomatal closure in response to increasing D from 1.04 to 2.06 kPa was only partially reversed by pressurization. Our results are in contrast to those of other studies on herbaceous species, even though we followed the same experimental approach. They suggest that it is not always appropriate to invoke a ‘feedforward’ model of short-term stomatal response to soil drying, whereby chemical messengers from the roots bring about stomatal closure.     

Experimental Studies of the Factors Controlling Transpiration: III. THE INTERRELATIONS BETWEEN TRANSPIRATION RATE, STOMATAL MOVEMENT, AND LEAF-WATER CONTENT

Transpiration rates of single leaves of Pelargonium and wheatwere measured under constant conditions of light, temperature,and air flow. Concurrently, stomatal movement was followed withthe resistance porometer during cycles of changing water contentof the leaf and changes induced by light and darkness. Stomatalmovement was found to exert a large controlling influence onthe transpiration rate, whereas water content had an extremelysmall or negligible effect. An approximately inverse linearrelation between transpiration rate and logarithm of resistanceto viscous flow through the leaf is believed to be the resultantof an inverse curvilinear relationship between the diffusiveconductance of the stomata and log. leaf resistance and thedecreasing difference of vapour pressure arising from the highertranspiration rates with increasing stomatal conductances. Nevertheless,the relation demonstrates that the transpiration rate is influencedby the degree of stomatal opening throughout its entire range. There was some evidence of lower transpiration rates duringand after recovery from wilting than before wilting. This isattributed to a decrease in a cell-wall conductance, the evaporatingsurface being located within the cell wall. During wilting partiallyirreversible contraction of the cell wall occurs. There wasalso evidence of slow changes in cell volume at full turgidityattributable to plastic flow. These occurred when the leaf wastransferred from environments of a high to low potential forevaporation. Extensive movement of the stomata followed changes in leaf water,passive opening resulting from decrease and closure from increaseof leaf water. It is suggested that the direction and extentof stomatal changes induced by water deficits is a consequenceof the rate of change of leaf water content and not of the absolutevalues. The stomata also showed an enhanced tendency to closein dry moving air following a period of wilting even after theleaf had regained turgidity.     

A Method for Separating Cuticular and Stomatal Components of Gas Exchange by Amphistomatous Leaves: II. EXPERIMENTAL SOLUTION

The apparent cuticular component of transpiration of stomatabearing leaf epidermis was estimated by restricting stomataldiffusion by mass flow of air in the opposite direction. Thiswas achieved by applying an air pressure gradient across theamphistomatous leaf. Some assumptions of the previously suggestedmethod (antrcek and Slav?k, 1990) were experimentally verifiedusing maize leaves. The technique makes possible a quantitativeestimation of cuticular water loss including that of the externalperistomatal (i.e. vapour not passing through the pores) andthe respective conductance when the stomata are partially open. In addition to the fact that the cuticular portion of the totalleaf vapour loss (i.e. relative cuticular transpiration) dependson stomatal opening, even the absolute value of apparent cuticulartranspiration was (1) increased by lower vapour pressure deficitand (2) decreased with closing stomata. These changes, inducedby variations in a vapour pressure deficit of 2.45?0.35 kPa,ranged between 0.66?0.14µg cm ^–2 s^–1. Theabsolute value of apparent cuticular transpiration changed onaverage by a factor of 2.3 due to stomata opening or closingwhich was induced by turning the light on or by exogenous ABAapplication. Possible interference by residual vapour diffusingthrough the stomatal pore was evaluated by the model application.An attempt was also made to assess the cuticular component ofCO₂-uptake rate. Experimental results are discussed in contextwith the feedforward response of stomata to air humidity. Key words: Cuticular transpiration, cuticular CO₂-uptake, feedforward response, maize     

Transpiration does not control growth and nutrient supply in the amphibious plant Mentha aquatica

Mentha aquatica L. was grown at different nutrient availabilities in water and in air at 60% RH. The plants were kept at 600 mmol m^?3 free CO₂ dissolved in water (40 times air equilibrium) and at 30 mmol m^?3 CO₂ in air to ensure CO₂ saturation of growth in both environments. We quantified the transpiration-independent water transport from root to shoot in submerged plants relative to the transpiration stream in emergent plants and tested the importance of transpiration in sustaining nutrient flux and shoot growth. The acropetal water flow was substantial in submerged Mentha aquatica, reaching 14% of the transpiration stream in emergent plants. The transpiration-independent mass flow of water from the roots, measured by means of tritiated water, was diverted to leaves and adventitious shoots in active growth. The plants grew well and at the same rates in water and air, but nutrient fluxes to the shoot were greater in plants grown in air than in those that were submerged when they were rooted in fertile sediments. Restricted O₂ supply to the roots of submerged plants may account for the smaller nutrient concentrations, though these exceeded the levels required to saturate growth. In hydroponics, the root medium was aerated and circulated between submerged and emergent plants to minimize differences in medium chemistry, and here the two growth forms behaved similarly and could fully exploit nutrient enrichment. It is concluded that the lack of transpiration from leaf surfaces in a vapour-saturated atmosphere, or under water, is not likely to constrain the transfer of nutrients from root to shoot in herbaceous plants. Nutrient deficiency under these environmental conditions is more likely to derive from restricted development and function of the roots in waterlogged anoxic soils or from low porewater concentrations of nutrients.